Zoogeography of Arachnida

Petar Beron

The classical zoogeographical subdivision of the land fauna, done by Sclater and Wallace in the nineteenth century, is still “official” in our time. This subdivision has been based on vertebrates, mostly mammals and birds. The ocean of tiny creatures (insects, arachnids, etc.) has been (and still is) largely neglected. Some attempts have been done to restructure the old scheme (Lopatin, Krizhanovskiy, Morrone, etc.), but nothing has been done for the total of Arachnida – old groups of non-flying invertebrates, quite important for zoogeography and already relatively well known. The classical zoogeographical subdivision of the land fauna, done by Sclater and Wallace in the nineteenth century, is still “official” in our time. This subdivision has been based on vertebrates, mostly mammals and birds. The ocean of tiny creatures (insects, arachnids, etc.) has been (and still is) largely neglected. Some attempts have been done to restructure the old scheme (Lopatin, Krizhanovskiy, Morrone, etc.), but nothing has been done for the total of Arachnida – old groups of non-flying invertebrates, quite important for zoogeography and already relatively well known. By venturing into this mammoth task, to put together the information about all orders in Arachnida and to elaborate on their distribution could be explained (besides with my long experience as a lecturer of zoogeography) with the words of one of the prominent arachnologists of our time, the Brazilian Adriano Kury “In a moment of deprivation of common sense” (Kury, on line). Quot potui – feci! Some 60 years ago, such analysis would have been much more incomplete, much more difficult, and on completely different bases. Since the middle of the twentieth century, so many new descriptions have appeared, revisions on the whole cladistics taxonomy and zoogeography, so we have now a picture much closer to reality. At the same time, the huge amount of information became very hard to digest, especially for a single analyzer. It is fortunate that some competent specialists provided catalogues, bibliographies, and checklists of whole groups of Arachnida and of particular countries or regions. We must mention some (not all) of these very helpful arachnological manuals, published in the twentieth and twenty-first centuries: The “smaller orders”: Harvey (2003) Palpigradi: Kraepelin (1901), Rowland and Sissom (1980), Condé (1996)

Monographiae Biologicae 94 Petar Beron Zoogeography of Arachnida Monographiae Biologicae Volume 94 Series editor Henri J. Dumont, Ghent University, Department of Biology, Ghent, Belgium Aims and Scope The Monographiae Biologicae provide a forum for top-level, rounded-off monographs dealing with the biogeography of continents or major parts of continents, and the ecology of well individualized ecosystems such as islands, island groups, mountains or mountain chains. Aquatic ecosystems may include marine environments such as coastal ecosystems (mangroves, coral reefs) but also pelagic, abyssal and benthic ecosystems, and freshwater environments such as major river basins, lakes, and groups of lakes. In-depth, state-of-the-art taxonomic treatments of major groups of animals (including protists), plants and fungi are also elegible for publication, as well as studies on the comparative ecology of major biomes. Volumes in the series may include single-author monographs, but also multi-author, edited volumes. More information about this series at http://www.springer.com/series/6304 Petar Beron Zoogeography of Arachnida Petar Beron National Museum of Natural History, Bulgarian Academy of Sciences Sofia, Bulgaria ISSN 0077-0639 ISSN 2215-1729 (electronic) Monographiae Biologicae ISBN 978-3-319-74417-9 ISBN 978-3-319-74418-6 (eBook) https://doi.org/10.1007/978-3-319-74418-6 Library of Congress Control Number: 2018932984 © Springer International Publishing AG, part of Springer Nature 2018 This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. The publisher, the authors and the editors are safe to assume that the advice and information in this book are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the editors give a warranty, express or implied, with respect to the material contained herein or for any errors or omissions that may have been made. The publisher remains neutral with regard to jurisdictional claims in published maps and institutional affiliations. Printed on acid-free paper This Springer imprint is published by the registered company Springer International Publishing AG part of Springer Nature. The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland E. Haeckel, Kunstformen der Natur, 1904 “One of the primary suppositions of sciences is that world is orderly, that there are patterns, and that patterns need causal explanation. The same can be said of the science of historical biogeography.” J. Cracraft (1988) To the memory of Vassil B. Guéorguiev Bulgarian entomologist, biospeleologist, and zoogeographer, for our friendship and his ideas Contents 1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 2 Builders of Arachnology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bibliography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 33 3 Systems of Arachnida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bibliography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35 41 4 The Fossil Arachnida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bibliography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45 51 5 Factors Determining the Distribution of Arachnida . . . . . . . . . . . . . . 5.1 Some Basic Notions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.1.1 Zoogeography and Biogeography: The New Approach . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.1.2 Centers of Origin and Dispersal . . . . . . . . . . . . . . . . . . . 5.1.3 Development of Atlantic Ocean . . . . . . . . . . . . . . . . . . . 5.1.4 Development of the Indian Ocean . . . . . . . . . . . . . . . . . 5.2 Paleogeography and the Past Distribution of Arachnida . . . . . . . 5.2.1 Age of the Faunas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.3 Passive Dispersal and Phoresy of Arachnida: Ballooning, Rafting, and Continental Drift . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.3.1 Dispersal (Natural and by Man Activities) . . . . . . . . . . . 5.4 Ecological Factors . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5.4.1 Climatic and Other Barriers and Bridges . . . . . . . . . . . . 5.4.2 Real and “Fake” Disjunctions. . . . . . . . . . . . . . . . . . . . . 5.4.3 Old World-New World Disjunctions. . . . . . . . . . . . . . . . 5.4.4 West African-Indomalayan Disjunction . . . . . . . . . . . . . 5.4.5 Strange (Old?) Disjunctions . . . . . . . . . . . . . . . . . . . . . . 5.4.6 Disjunctions in the Southern End of the World (New Zealand, Australia, South Africa, Patagonia) . . . . 5.4.7 Boreomontane and Arctic-Alpine Disjunctions . . . . . . . 5.4.8 Transpacific Disjunctions . . . . . . . . . . . . . . . . . . . . . . . . 59 59 59 60 61 64 65 65 72 72 76 76 77 79 81 81 81 81 82 xi xii Contents 5.5 Endemics and Relicts in Arachnida . . . . . . . . . . . . . . . . . . . . . . . 83 5.5.1 Endemism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83 5.5.2 Relicts and Relictness . . . . . . . . . . . . . . . . . . . . . . . . . . . 94 5.6 The Northern Limits of Arachnida. . . . . . . . . . . . . . . . . . . . . . . . 97 Bibliography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100 6 Review of the Classification of the Orders of Arachnida . . . . . . . . . . 6.1 Palpigradi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.2 Schizomida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.3 Thelyphonida (Uropygi). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.4 Amblypygi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.5 Ricinulei . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.6 Scorpiones . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.7 Pseudoscorpiones . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.8 Opiliones . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.9 Solifugae (Solpugida, Solifugi) . . . . . . . . . . . . . . . . . . . . . . . . . . 6.10 Araneae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.11 Opilioacarida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.12 Trombidiformes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.13 Sarcoptiformes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.14 Holothyrida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.15 Ixodida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6.16 Mesostigmata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bibliography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135 135 136 138 140 143 144 146 149 159 170 185 189 193 194 198 199 199 7 Regional Arachnogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.1 Holarctic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.1.1 Zoogeography and Arachnozoogeography. . . . . . . . . . . 7.2 Europe (General) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.2.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.2.2 Arachnogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.3 Balkan Peninsula and Aegaeis . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.3.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.3.2 Arachnogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.4 Apennine Peninsula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.4.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.4.2 Arachnogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.5 Iberian Peninsula and the Pyrenees . . . . . . . . . . . . . . . . . . . . . . . 7.5.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.5.2 Arachnogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.6 Mediterranean Sea and Its Islands . . . . . . . . . . . . . . . . . . . . . . . . 319 319 319 320 320 323 328 328 333 342 342 343 344 344 345 348 Contents 7.7 7.8 7.9 7.10 7.11 7.12 7.13 7.14 7.15 7.16 7.17 xiii Balearic Islands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.7.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.7.2 Arachnogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sicily, Sardinia, Corsica, and Elba . . . . . . . . . . . . . . . . . . . . . . . . 7.8.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.8.2 Arachnogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Malta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.9.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.9.2 Arachnogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Central and Northern Europe, Great Britain, Ireland, Island, and Faroe Islands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.10.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.10.2 Arachnogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Arachnida of the Atlantic Islands (Macaronesia, St. Helena, Ascension) . . . . . . . . . . . . . . . . . . . . . 7.11.1 Canary Islands, Azores, and Madeira . . . . . . . . . . . . . . . Cabo Verde . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.12.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.12.2 Arachnogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . St. Helena and Ascension. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.13.1 Geography and Paleogeography (General). . . . . . . . . . . 7.13.2 Arachnogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Russia (North of Caucasus), Belarus, Ukraine, Siberia, Altai, China (North of Yangtze), and Mongolia . . . . . . . 7.14.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.14.2 Arachnogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sakhalin and Kuril Islands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.15.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.15.2 Arachnogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Caucasus and Transcaucasia . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.16.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.16.2 Arachnogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Turkey, Cyprus, Sinai, the Dodecanese Islands, Iran, Afghanistan, the Near East, Middle Asia, and the Arabian Peninsula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.17.1 Geography, General Zoology, and Paleogeography . . . . 7.17.2 Arachnogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 350 350 351 352 352 354 357 357 358 359 359 359 363 363 367 367 367 368 368 368 370 370 371 374 374 374 375 375 376 379 379 380 xiv Contents 7.18 7.19 7.20 7.21 7.22 7.23 7.24 7.25 7.26 7.27 7.28 Socotra . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.18.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.18.2 Arachnogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Karakorum, Hindu Kush, Pamir, Tian Shan, Himalaya, and Tibet . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.19.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.19.2 Arachnogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Indian Peninsula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.20.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.20.2 Arachnogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ceylon (Sri Lanka) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.21.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.21.2 Arachnogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Andaman and Nicobar Islands . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.22.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.22.2 Arachnogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Southeast Asia (Burma, Thailand, Indochina, Peninsular Malaysia, China South of Yangtze, and Hainan) . . . . 7.23.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.23.2 Arachnogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Taiwan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.24.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.24.2 Arachnogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . The Malay Archipelago . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.25.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.25.2 Arachnogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . The Philippines. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.26.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.26.2 Arachnogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Japan (the Ryukyus Excluded) . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.27.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.27.2 Arachnogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ryukyu Islands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.28.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.28.2 Arachnogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 391 391 391 393 393 394 397 397 397 400 400 400 403 403 404 404 404 405 421 421 421 423 423 424 431 431 431 435 435 436 438 438 439 Contents 7.29 Korea (North and South) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.29.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.29.2 Arachnogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.30 Africa, North of 20oN . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.30.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.30.2 Arachnogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.31 Africa Between 20oN and Zambezi-Kunene (Tropical or Intertropical Africa) . . . . . . . . . . . . . . . . . . . . . . . . . 7.31.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.31.2 Arachnogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.32 Differences Between the Arachnofaunas of East and West Africa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.33 Tropical South America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.33.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.33.2 Arachnogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.34 Galapagos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.34.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.34.2 Arachnogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.35 Juan Fernandez Islands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.35.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.35.2 Arachnogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.36 North America, North of Rio Grande (incl. Greenland) . . . . . . . 7.36.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.36.2 Arachnogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.37 Polynesia and Micronesia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.37.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.37.2 Arachnogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.38 Hawaii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.38.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.38.2 Arachnogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.39 Antarctic and Subantarctic Arachnofauna . . . . . . . . . . . . . . . . . . 7.39.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7.39.2 Arachnogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bibliography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xv 441 441 442 444 444 448 453 453 454 475 476 476 478 502 502 503 505 505 505 506 506 507 514 514 515 519 519 520 522 522 523 528 xvi 8 Contents Some Peculiarities of the Distribution of Arachnida . . . . . . . . . . . . . . 8.1 Mediterranean and Central European Arachnofauna . . . . . . . . . . 8.2 Arachnofauna of the Ancient Mediterranean . . . . . . . . . . . . . . . . 8.3 The Boundary Neotropica-Nearctic . . . . . . . . . . . . . . . . . . . . . . . 8.3.1 Palpigradi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.3.2 Ricinulei . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.3.3 Pseudoscorpiones . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.3.4 Opiliones. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.3.5 Amblypygi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.3.6 Uropygi (Thelyphonida) . . . . . . . . . . . . . . . . . . . . . . . . . 8.3.7 Schizomida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.3.8 Araneae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.3.9 Opilioacarida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.4 Mexico: The Great American Interchange of Fauna . . . . . . . . . . 8.4.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.5 Arachnida of the Antilleans and Mesoamerica . . . . . . . . . . . . . . 8.5.1 Central America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.6 Antillean (Caribbean) Islands . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.6.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.6.2 Comparison Between the Faunas of Central America and the Antilleans . . . . . . . . . . . . . . . . . . . . . . 8.6.3 Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.6.4 Opiliones. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.7 Patagonia: South Chile, Argentina, Uruguay, and Tierra del Fuego . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.7.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.7.2 Arachnogeographical Character of South Chile and Argentina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.8 Falkland Islands (Malvinas) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.8.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.9 The Boundary Between Palearctic and Paleotropical . . . . . . . . . 8.9.1 The Boundary Between Western Palearctic and Afrotropical Arachnofaunas. . . . . . . . . . . . . . . . . . . 8.10 Arachnida in the Afrotropical and Palearctic Regions . . . . . . . . . 8.10.1 Analysis and Comments . . . . . . . . . . . . . . . . . . . . . . . . . 8.11 The Boundary Across China . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.11.1 The Transitional Zone. . . . . . . . . . . . . . . . . . . . . . . . . . . 8.11.2 Arachnogeography of Himalaya. . . . . . . . . . . . . . . . . . . 8.11.3 East Asiatic (Himalayan-Chinese) Region of Lopatin (1980) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.12 The Cape Arachnofauna . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.12.1 Africa South of Zambezi-Kunene. . . . . . . . . . . . . . . . . . 625 625 625 626 626 627 627 627 628 628 628 628 629 629 629 639 639 650 650 669 673 673 674 674 675 679 679 680 680 681 681 688 688 689 690 690 690 Contents 8.13 Arachnofauna of Madagascar and the Adjacent Islands . . . . . . . . 8.14 Madagascar . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.14.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.15 Mascarene Islands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.15.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.16 Seychelles (incl. Aldabra), Comores . . . . . . . . . . . . . . . . . . . . . . 8.16.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.16.2 Remarkable Distributions and Absences . . . . . . . . . . . . 8.17 Evaluation of Wallace’s Line and the Other Lines in Southeast Asia According to the Arachnida. . . . . . . . . . . . . . . 8.17.1 Wallace’s Line . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.17.2 Lydekker’s Line . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.17.3 Weber’s Line . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.17.4 Development of Wallacea and the Surrounding Lands of Southeast Asia and the Malayan Archipelago . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.17.5 The “Lines” and Some Other Groups . . . . . . . . . . . . . . . 8.17.6 Arachnida and the Lines . . . . . . . . . . . . . . . . . . . . . . . . . 8.17.7 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.18 The Arachnida of the Papuan Subregion . . . . . . . . . . . . . . . . . . . 8.18.1 New Guinea and Melanesia . . . . . . . . . . . . . . . . . . . . . . 8.19 Arachnogeography of Australia and Tasmania . . . . . . . . . . . . . . 8.19.1 Australia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.20 Tasmania. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.20.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.21 Parasitiformes (Some Ixodida and Mesostigmata): Australia and New Zealand . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.22 New Zealand . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.22.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.23 New Caledonia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.23.1 Geography, General Zoogeography, and Paleogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.24 Lord Howe Island . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.25 Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8.25.1 Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bibliography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 xvii 701 701 701 711 711 713 713 726 727 728 728 728 730 731 732 743 746 746 753 753 764 764 767 768 768 774 774 777 778 782 783 Cave Arachnida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 833 9.1 Cave Life in Different Orders . . . . . . . . . . . . . . . . . . . . . . . . . . . 833 9.2 Distribution of Cave Arachnida in Different Geographical Areas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 842 xviii Contents 9.3 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 846 Bibliography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 846 10 High-Altitude Arachnida (Partly from Beron 2008, Updated in Beron 2016) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10.1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10.2 Major Groups of Arachnida in the Old World Living At or Above 2200 m . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10.3 Level of Knowledge on Opiliones in the High Mountains and Credibility of Conclusions. . . . . . . . . . . . . . . . . . . . . . . . . . . 10.4 Maximum Altitudes of Spiders Living Above 2200 m in the Old World . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10.5 High-Altitude Arachnida in the Oreal of the Himalayas, Compared with the Alps. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10.6 High-Altitude Arachnida in the Mountains of Central Asia . . . . 10.7 Afrotropical High-Altitude Arachnida . . . . . . . . . . . . . . . . . . . . . 10.8 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10.9 Champions in Altitude (Above 4500 m) in the Old World . . . . . 10.10 Some Personal Notes on the High-Altitude Arachnofauna of the Americas and Elsewhere . . . . . . . . . . . . . . . . . . . . . . . . . . Bibliography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 853 853 860 864 864 870 880 882 884 884 886 888 11 Zoogeographical Notes on Different Orders of Arachnida . . . . . . . . 11.1 Palpigradi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11.2 Schizomida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11.3 Amblypygi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11.4 Thelyphonida (Uropygi). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11.5 Ricinulei . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11.6 Solifugae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11.7 Scorpiones . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11.8 Pseudoscorpiones . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11.9 Opiliones . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11.10 Araneae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11.11 Opilioacarida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bibliography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 893 893 894 894 895 896 896 897 897 898 900 902 902 12 Zoogeographical Subdivisions of the World . . . . . . . . . . . . . . . . . . . . Bibliography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 905 927 13 General Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bibliography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 931 936 Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 939 Chapter 1 Introduction Abstract The classical zoogeographical subdivision of the land fauna, done by Sclater and Wallace in the nineteenth century, is still “official” in our time. This subdivision has been based on vertebrates, mostly mammals and birds. The ocean of tiny creatures (insects, arachnids, etc.) has been (and still is) largely neglected. Some attempts have been done to restructure the old scheme (Lopatin, Krizhanovskiy, Morrone, etc.), but nothing has been done for the total of Arachnida – old groups of non-flying invertebrates, quite important for zoogeography and already relatively well known. The classical zoogeographical subdivision of the land fauna, done by Sclater and Wallace in the nineteenth century, is still “official” in our time. This subdivision has been based on vertebrates, mostly mammals and birds. The ocean of tiny creatures (insects, arachnids, etc.) has been (and still is) largely neglected. Some attempts have been done to restructure the old scheme (Lopatin, Krizhanovskiy, Morrone, etc.), but nothing has been done for the total of Arachnida – old groups of non-flying invertebrates, quite important for zoogeography and already relatively well known. By venturing into this mammoth task, to put together the information about all orders in Arachnida and to elaborate on their distribution could be explained (besides with my long experience as a lecturer of zoogeography) with the words of one of the prominent arachnologists of our time, the Brazilian Adriano Kury “In a moment of deprivation of common sense” (Kury, on line). Quot potui – feci! Some 60 years ago, such analysis would have been much more incomplete, much more difficult, and on completely different bases. Since the middle of the twentieth century, so many new descriptions have appeared, revisions on the whole cladistics taxonomy and zoogeography, so we have now a picture much closer to reality. At the same time, the huge amount of information became very hard to digest, especially for a single analyzer. It is fortunate that some competent specialists provided catalogues, bibliographies, and checklists of whole groups of Arachnida and of particular countries or regions. We must mention some (not all) of these very helpful arachnological manuals, published in the twentieth and twenty-first centuries: The “smaller orders”: Harvey (2003) Palpigradi: Kraepelin (1901), Rowland and Sissom (1980), Condé (1996) © Springer International Publishing AG, part of Springer Nature 2018 P. Beron, Zoogeography of Arachnida, Monographiae Biologicae 94, https://doi.org/10.1007/978-3-319-74418-6_1 1 2 1 Introduction Schizomida: Harvey (1992), Reddell and Cokendolpher (1995) Amblypygi: Mullinex (1975), Quintero (1981 1986) Thelyphonida (Uropygi): Rowland and Cooke (1973) Ricinulei: Tuxen (1974) Solifugae: Kraepelin (1901), Gromov and Savary (Bibliography on line) Scorpiones: Fet, Sissom, Lowe and Braunwalder (2000) Pseudoscorpiones: Chamberlin (1931), Beier (1932a 1932b 1963), Harvey (1990 1992), Schawaller (1980, bibliography) Opiliones: Roewer (1923), Rosas Costa (1950 – Cyphophthalmi), Hallan (2005, checklist), Cokendolpher (1997, bibliography), Giribet (2000, Cyphophthalmi), Kury (2003, catalogue New World Laniatores; 2012, description of catalogues and checklists), Pinto-da-Rocha, Machado and Giribet (Eds.) (2007), Schönhofer (2013, Dyspnoi) Araneae: Roewer (1942), Brignoli (1983), Platnick (The World Spider Catalog) Opilioacarida: Hammen van der (1966), Beron (2014) Holothyrida: Beron (2014) Prostigmata: Beron (2008b, 2016) Mesostigmata: Beron (2016) Regional lists and bibliographies: Old World: Beron (2008a, high altitude Arachnida) New World: Kury (2003, Opiliones Laniatores) Africa: Dupré (2013, Scorpiones), Lamoral and Reynders (1975, Scorpions, Ethiopian Region), Scharff (1990, Linyphiidae), Starega (1984, Phalangiidae; 1992, African Opiliones, except Phalangiidae) East Africa: Probst (1973, Scorpiones) South Africa: Lawrence (1931, Opiliones; 1955, Solifugae, Scorpiones and Pedipalpi, Schizomida, 1969), Kauri (1961, Opiliones), Lotz (2009 – Opiliones) Madagascar: Lawrence (1959, Opiliones), Lourenço (1996, Scorpiones) Australia: Forster (1955 and further), Glauert (1963, Scorpiones, W. Australia), Koch (1977, Scorpiones, Australia and Papua), Harvey (1992) Europe: Beier (1963, Pseudoscorpiones), Stol (1993, 2007, Opiliones, North Europe), Deltshev and Blagoev (2001, Araneae, Bulgaria) North America: Hoff (1958, Pseudoscorpiones), Cokendolpher and Lee (1993, Opiliones) Texas: Rowland and Reddell (1976, Texas) South America: Benavides and Giribet (2007, Opiliones Neogoveidae), MelloLeitão (1945, Scorpiones), Kury (2003, Laniatores), Caporiacco (1948, Guianas, Opiliones), Ringuelet (1959, Argentina, Opiliones) West Indies: Cokendolpher and Camilo-Rivera (1989, Opiliones, bibliography) Mexico: Díaz Nájera (1975, Araneae), Kury and Cokendolpher (2000, Opiliones), Vazquez Rojas (1981 – Solifugae) Madagascar: Emerit (1974, Araneae, Gasteracanthinae), Lawrence (1959, Opiliones), Lourenço (1996, Scorpiones), Araneae (Goodman and Benstead 2005) 1 Introduction 3 New Zealand: Forster (1967–1973, Araneae; 1978, Opiliones), Dumbleton, 1953, Heath, 1977 (Ixodida) Oceania: Chamberlin (1934) Chile: Cekalovic (1975, Solifugae; 1976, Arachnida; 1983, Scorpiones; 1984, Pseudoscorpiones and Palpigradi; 1986, Opiliones), Beier (1964, Pseudoscorpiones) Northern Asia: Eskov (1994, Linyphiidae); Fet (1988, Scorpiones, the former USSR); Marusik, Eskov, and Kim (1992, Araneae); Mikhailov (1998 1999 2000 2002 2013, Araneae, the former USSR); Staręga (1978 – Opiliones of the former USSR) Panama: Fairchild et al. (1966 – Ixodida) Neotropical Region: Guglielmone et al. (2003, Ixodida) Georgia: Djaparidze (1960 – Ixodida) India: Murthy and Ananthakrishnan (1977, Pseudoscorpiones), Pocock (1900, Arachnida), Siliwal, Molur and Biswas (2005, Araneae), Tikader (1987, Araneae) Nepal: Clifford et al. (1975, Ixodida) Iran: Abassian-Lintzen (1960, Ixodida), Mozaffarian and Marusik (2001, Araneae) Arachnida: Blick, Hänggi, and Thaler (2002, Germany, Switzerland, Austria, Belgium, and the Netherlands), Maes et al. (1989, Nicaragua), Zaragoza (2007 – Iberian Peninsula) Palpigradi: Kraepelin (1901), Rowland and Sissom (1980), Condé (1996) Solifugae: Aliev and Gadzhiev (1983, Azerbaydjan), Armas and Teruel (2005, Cuba), Gromov and Savary (bibliography), Levy and Shulov (1964, Israel) Schizomida: Armas 2004 (Cuba, Dominican Rep.), Harvey (1992, 2000, Australia; 2001, Seychelle Isl.); Reddell and Cokendolpher (1995) Amblypygi: Armas (2004, 2009b, Antilleans), Mello-Leitão (1931, Brazil) Thelyphonida (Uropygi): Mello-Leitão (1931, Brazil) Ricinulei: Tuxen (1974, Africa) Scorpiones: Acosta and Maury (1998, Argentina); Acosta and Ochoa (2002, Bolivia); De Armas (2009a, Antilleans); Fet (1988, former USSR; 1994, Tukmenistan; 2010, Europe); Koch (1977, Australia); Lamoral (1979, Namibia); Levy and Amitai (1980, Palestina); Lourenço and Méndez (1984, Panama); Lourenço (1995a, Ecuador; 1996, Madagascar; 1997, Colombia); Maury (1984, Paraguay); Mello-Leitão (1932, Brazil); Zhu, Qi, and Song (2004, China); Ojanguren-Affilastro (2005 – Argentina) Pseudoscorpiones: Agnarsson (1998, Iceland); Beier (1932, 1965a, New Guinea; 1966, Philippines); Chamberlin (1934, Oceania); Ćurčić (1974, Yugoslavia); Ćurčić, Dimitriević, and Legakis (2004, Serbia, Montenegro, Macedonia); de Lessert (1911, Switzerland); El-Hennawy (1988, Egypt); Gardini (2000, Italia); Harvey (1990); Kunt (2008, Turkey); Legg and O’Connor (1997, Ireland); Mahnert (1981–1988, Kenya; 1975, Malta; 2004, Austria); Petrov (1997, Bulgaria); Schawaller (1980, bibliography; 1994b, Thailand; 1995, China); Telnovs (2002a, Latvia); Zaragoza (2000, bibliography, Iberian Peninsula, Baleares, Macaronesia); Legg (1988, Great Britain); Murthi and Anasnthakrishnan (1977, India) 4 1 Introduction Opiliones: Agnarsson (1998, Iceland); Babalean (1992, Romania); Bayram et al. (2010, Turkey); Bezdecka (2008, Czech Republic); Blick and Komposh (2004, Central and Northern Europe); Canals (1936, Chile); Cawley (2002, Ireland); Chevrizov (1980, URSS); Cokendolpher (1990, Egypt); Cokendolpher and Camilo-Rivera (1989, West Indies); Cokendolpher and Lee (1993, Greenland, Canada, USA, Mexico); Gritzenko (1978, Asian part of USSR); Forster (1954, New Zealand); Hillyard and Sankey (1989, British Fauna); Kim, D.-H., J.-W. Lee, and J.-P. Kim (2006, Korea); Klimeş (2000, Czech and Slovak Rep.); Komposch (2004, Hungary); Komposch and Gruber (2005, Austria); Lawrence (1959, Madagascar); Lotz (2009, Southern Africa); Martens (1978, Germany; many papers, Nepal); Mheidze (1964, Georgia); Novak (2004, Croatia; 2005, Bosnia and Herzegowina); Novak et al. (2006, Slovenia); Prendini (2010, Seychelles); Rafalski (1960, 1961, Poland); Rafalski and Staręga (1997, Poland); Rambla (1967, Portugal); Redikorzev (1936 – Soviet Union); Ringuelet (1959, Argentina; 1963, Uruguay); Roewer (1923); Shavanova (2004, Belarus); Šilhavý (1956, Czechoslovakia); Staręga (1976, Bulgaria; 1978, Soviet Union; 1992, Afrotropical Region; 2000, Poland); Suzuki (1985a, 1985b, Thailand); Telnovs (2002b, Latvia); Tsurusaki (1993, Japan) Araneae: Aakra and Hauge (2000, 2003, Norway, Svalbard, Jan Mayen); Agnarsson (1996, Iceland); Bosmans (2009, Belgium); Bosmans and Chadzaki (2005, Greece); Bosmans and de Keer (1985, Pyrenees); Blagoev (2002, Macedonia); Bonnet (1945–1961, bibliography); Brignoli (1983); Cardoso and Morano (2010, Iberian Peninsula); Deltshev (2005, Bulgaria); Deltshev and Blagoev (2001, Bulgaria); Deltshev, Curčić, and Blagoev (2001, Serbia); Drensky (1936, Balkan Peninsula); Evenhuis (2006, Fiji); Forster et al. (1967–1973, New Zealand); Forster and Forster (1973, New Zealand), Gajdos, Svaton, and Sloboda (1999, Slovakia); Kim (1990, Korea); Kostanjšek and Kuntner (2015, Slovenia); Larsen and Scharff (2003, Greenland); Le Peru (2007, France); Marinu and Verneau (2002, Corsica); Marusik, Eskov, Logunov, and Basarukin (1993, Sakhalin and Kuril Isls); Marusik, Eskov, Koponen, and Vinokurov (1993, Yakutia); Kronestedt (2001, Sweden); Marusik, Logunov, and Koponen (2000, Tuva); Merrett, Locket, and Millidge (1985, Great Britain); Mikhailov and Fet (1994, Turkmenistan); Mikhailov (1997, 1998, 1999, 2000, 2002, 2013, the former Soviet Union); Milosević (2002, Croatia); Nikolić and Polenec (1981, f. Yugoslavija); Platnick (2000–2011. The World Spider Catalog, Version 11.5.); Proszynski and Staręga (1971, Poland); Relys and Spungis (Latvia); Roewer (1942); Song, Zhang, and Daigin (2002, Singapore); Starega (2000, Poland); Suman (1964, Hawaii), Tikader (1970, Sikkim); Topcu, Demir, and Seyyar (2005, Turkey); Varol (2003, Turkey); Vilkas (1992, Lithuania); Weiss and Urak (2000, Romania) Opilioacarida: Beron (2014), Vasquez and Klompen (2002, North and Central America; 2010, Madagascar) Holothyrida: Beron (2014), Lehtinen (1995) Ixodida: Drensky (1955, Bulgaria), Feider (1965, Romania), Filippova (1966, Argasidae of the World), Starkoff (1958, Ixodida of Italy), Georgieva and 1 Introduction 5 Gecheva (2013, Ixodidae of Bulgaria), Gregson (1956, Ixodida of Canada), Anastos (1950, Ixodida of Indonesia), Roberts (1964, Ixodida of Tasmania; 1970, Ixodida of Australia), Dumbleton 1953, Heath 1977 (Ixodida of New Zealand), Hoogstraal (1953, Ixodida of Madagascar) Prostigmata: Beron (2008a – Calyptostomatoidea and Erythraeoidea of the world) Oribatida: Shtanchaeva 2003 (Caucasus), Collof and Haliday 1998 (Australia), Bayartogtokh 2010 (Mongolia), Vu Quang Manh (Vietnam) This book would have been impossible without the generous help of many arachnologists and biogeographers since the 1960s, when I started gathering material for it. Some of these people I knew personally, I received regularly their publications and advice, and also I benefited from their hospitality in many institutes and museums; I had also access to their private book and reprint collections. I cannot enumerate all of them, but I will mention some, without their titles. Bulgaria – Ch. Deltshev, B. Gruev, V.B. Guéorguiev, P. Mitov, B. Petrov, S. Lazarov Serbia – B. Ćurčić, I. Karaman, S. Matvejev Slovenia - I. Hadži Croatia – T. Novak Romania – M. Dumitresco, D. Dumitrescu, Z. Feider, S. Avram Hungary – Y. Kontchán, S. Mahunka, Z. Varga Czech Rep. – J. Buchar, M. Daniel, F. Kovařik, J. Kratochvil, M. Krumpál, V. Šilhavý Poland – J. Rafalski, W. Staręga, R. Haitlinger, J. Mąkol, G. Gabryś, W. Weselowska Italy – P.M. Brignoli, G. Callaini, L. Delle Cave, G. Gardini, G. Lazzeroni, I. Marcellino, M. La Greca, A. Vigna Taglianti France – L. Berland, B. Condé, J. Heurtault, C. Juberthie, M. Judson, P. Leclerc, R. Legendre, W.R. Lourenço, M. Vachon, A. Vandel, I. Coineau Great Britain – G. Legg, R. Hall, P.J.A. Pugh Belgium – P. Benoit, R. Bosmans, A. Fain, R. Jocqué Netherlands – L. van der Hammen Austria – M. Beier, E. Christian, J. Gruber, H. Franz, C. Komposch, H. Schatz, K. Schmölzer, K. Thaler, H. Janetschek Switzerland – V. Mahnert Denmark – H. Enghoff, S.L. Tuxen Finland – J. Kaisila, P. Lehtinen Sweden – H. Lohmander Germany – J.A. Dunlop, J. Haupt, J. Martens, P. Müller, W. Schawaller, P. Weygoldt, J. Wunderlich Spain – M. Rambla, J.A. Zaragoza Russia – S. Golovatch, Yu. Marusik, A. Tanasevitch, O.L. Krijanovskij, U. Shtanchaeva, K.G. Mikhailov Kazakhstan – A.V. Gromov Georgia – D. Kobakhidze Iran – A. Saboori Turkey – K.B. Kunt 6 1 Introduction United States – E. Benedict, J.C. Chamberlin, J.C. Cokendolpher, J. Cracraft, V. Fet, O.F. Francke, G. Giribet, J.L. Gressitt (Hawaii), C. C. Hoff, C. and M. Goodnight, F. Howarth (Hawaii), H. Hoogstraal, E. Mayr, R.W. Mitchell, W.B. Muchmore, M. Muma, N. Platnick, L. Prendini, J.R. Reddell, J. M. Rowland, P.A. Selden, W. Shear Canada – S. Peck Cuba – L. de Armas, R. Teruel, M.A. Ituralde-Vinent, G. Alayón Garcia Panama – D. Quintero, Jr. Australia – B. Haloway, M. Harvey, V.V. Hickman, C.J. Humphries, L.E. Koch, D. Lohman, B. Main, P.V. Taylor, R. Southcott New Zealand – R.R. Forster Mexico – A. Hoffmann, G. Halffter, J.J. Morrone Brazil – L.J. de Araújo Feio, A.B. Kury, the family H. and B. Soares Venezuela – M.A. González - Sponga Argentina – V. Vitali-di-Castri Japan – K. Morikawa, M. Shimojana, S. Suzuki, N. Tsurusaki China – J.X. Zhang, Shuqiang Li Korea – Kap Yong Paik Philippines – L. Corpuz Raros India – M.S. Mani South Africa – R.F. Lawrence, H. Martin Israel – F.D. Por My sincere gratitude goes also to the colleagues who helped me a lot in preparing the maps and with other important “technicalities” – Mr. Assen Ignatov, Mrs. Sylvia Tosheva, and others. Chapter 2 Builders of Arachnology Abstract In many older authors (Aristotle and others), we can find information about spiders, scorpions, and other Arachnida. The scientific studies however started with the paper of Clerck (1757), the only one recognized as valid before the system of Linnaeus (1758). In many older authors (Aristotle and others), we can find information about spiders, scorpions, and other Arachnida. The scientific studies however started with the paper of Clerck (1757), the only one recognized as valid before the system of Linnaeus (1758). Before trying to discuss the zoogeographical problems of Arachnida, we owe a tribute to the builders of the arachnology by saying a few words about the life of some (by no means all) founding fathers. Some of them (Beier, Vachon, Brignoli, Kratochvil, Hadži) were known personally to the present author. To the correspondence with others (Chamberlin, Lawrence, Šilhavý, Suzuki, all other “living classics”), he owes much of his devotion to arachnology. Clerck, Carl (1709–1765) – the names in his paper of 1757 Svenska Spindlar were recognized as valid, and it is the oldest recognized paper in the zoological nomenclature. It started the scientific study of Arachnida. Presently, 53 spider species carry names given by Clerck. © Springer International Publishing AG, part of Springer Nature 2018 P. Beron, Zoogeography of Arachnida, Monographiae Biologicae 94, https://doi.org/10.1007/978-3-319-74418-6_2 7 8 2 Builders of Arachnology Linnaeus, Carolus (Carl von Linné) (1707–1778) – professor of medicine and botany at Uppsala (Sweden) and founder of the system of plants and animals (Systema Naturae, Ed. X, 1758) C. Linnaeus (Portrait by A. Roslin, 1775) 2 Builders of Arachnology 9 Thorell, Tord Tamerlan Theodor (1830–1901) – born in Sweden but lived longtime in Genoa and author of numerous important contributions to the system of Arachnida (On European spiders, 1869; Synonym European spiders, 1870–1873, many papers with descriptions of new taxa from Malaysian and Papuan collections, Arctica, Cameroon, etc.). Thorell described 12 families of spiders (Ctenizidae, Theraphosidae, Palpimanidae, Hersiliidae, Uloboridae, Oxyopidae, Amaurobiidae, Zodariidae, Philodromidae, etc.). Important papers are devoted also to Opiliones (13 publications on Opiliones from SE Asia, Indonesia, Burma, Argentina, USA, Europe, and West Asia). T.T.T. Thorell 10 2 Builders of Arachnology Sørensen, William Emil (1848–1916) – Danish arachnologist. Described 157 valid species of Opiliones. Some important papers are as follows: 1898, Arachnida Groenlandica (Acaris exceptis); 191, Opiliones (in I. Sjostedt’s Kilimanjaro – Meru volumes); and 1932, Descriptiones Laniatorum. William Emil Sørensen (1848–1916) 2 Builders of Arachnology 11 Hansen, H.J. (1855–1936) – Danish arachnologist. Some important papers: Hansen & Sorensen (1904) – established the modern system of Opiliones; Described many new taxa in Arachnida. Hans Jakob Hansen 12 2 Builders of Arachnology Mello-Leitão, Cândido Firmino de (1886–1948) – Brazilian arachnologist, professor, and explorer of Opiliones (198 publications, describing 347 valid species), Uropygi, Amblypygi, Scorpiones, Araneae, and some smaller orders in Brazil. Some important papers were Pedipalpos do Brasil e algumas notas sobre a ordem (1931), Opiliões do Brasil (1932, 511 pp.), Escorpiães Sul-Americanos (1945, 468 pp.), and others. Cândido Firmino de Mello-Leitão 2 Builders of Arachnology 13 Pocock, Reginald Innes (1863–1947) – English arachnologist and curator of the collections of Arachnida and Myriapoda from 1885 to 1904. Author of many papers on various groups of Arachnida. R.I. Pocock (From P. Bonnet, 1945) 14 2 Builders of Arachnology Kraepelin, Karl Matthias Friedrich Magnus (1848–1915) – German arachnologist, worked in the Zoology Museum in Hamburg, and author of many descriptions of various groups of Arachnida. Among the publications of Kraepelin are a volume on Palpigradi and Solifugae in Das Tierreich and many others. Karl Kraepelin 2 Builders of Arachnology 15 Roewer, Carl-Friedrich Roewer (1881–1963) – German arachnologist, professor, and director of the Overseas Museum in Bremen (1933–1945). His huge works laid the base of the system of Opiliones and of Solifugae. Since his first paper (Revision der Opiliones Plagiostethi, 1910, 294 pp.), he indicated a very mighty arrival in the field of taxonomy of Opiliones. In 1923 he published the biggest monograph on this group ever written – “Die Weberknechte der Erde” – followed by series of voluminous supplements (Weitere Weberknechte I–XX, 1927–1957) and by numerous publications, describing most of the known genera of Opiliones from all over the world and 2260 valid species. Another big achievement of Roewer was the system of Solifugae (1934, 1941). In his papers he described almost all genera of this order. Roewer wrote (without computer) also a Catalogue of Araneae from 1758 to 1940 (1942) and articles on other Arachnida (mostly Araneae). Some of the localities indicated by him have been questioned (Martens and v. Helversen 1972). His huge collection is preserved in Senckenberg Museum. C.-F. Roewer (From O. Kraus) 16 2 Builders of Arachnology Beier, Max (1903–1979) – his main fields have been the pseudoscorpions and the Mantodea. After several revisions, Beier’s two volumes of the series appeared Das Tierreich (Pseudoscorpionidea I and II, 1932). Together with the monograph of J.C. Chamberlin, published in the previous year, they laid the foundation of Pseudoscorpionology. In his papers on Pseudoscorpiones from all over the world, Beier published many new taxa and contributed greatly to the study of faraway countries. Many of his papers are devoted to Mantodea and Orthoptera. Max Beier has been an associate and director of Naturhistorisches Museum in Vienna. Max Beier 2 Builders of Arachnology 17 Chamberlin, Joseph Conrad (1898–1962) – one of the founders of the system of Pseudoscorpiones and nephew of R.V. Chamberlin. The second part of his thesis, entitled The Arachnid Order Chelonetida (1931), is the most important for the classification of the order. He described many new genera and the 12 of the 26 existent families: Lechytiidae, Bochicidae, Ideoroncidae, Neobisiidae, Menthidae, Atemnidae, Geogarypidae, Pseudochiridiidae, Pseudogarypidae, Hyidae, Syarinidae, and Withiidae. From 1921 to 1962, J.C. Chamberlin published 41 papers on Pseudoscorpiones of many countries and also other papers on Crustacea, systematic and economic entomology, etc. Joseph Conrad Chamberlin 18 2 Builders of Arachnology Brignoli, Paolo Marcelo (1942–1986) – Italian arachnologist, professor in Rome and Aquila, and specialist mainly in world spiders. From 1967 to 1985, he wrote 205 publications, describing 25 new genera and 370 new species. He has many contributions to the fauna of Greece, Mexico, Ethiopia, Nepal, Italy, and other countries, partly as a result of his own expeditions in different continents. Also he has publications in other Arachnida (Opiliones, Schizomida, etc.). He expressed interesting theoretical views on cave spiders. He wrote a catalogue of world spiders (1940–1981) of 755 pages. Paolo Marcelo Brignoli 2 Builders of Arachnology 19 Simon, Eugène (1848–1924) – French zoologist, working mainly on spiders and birds (Trochilidae). His main arachnological works are, 1864–1884, Histoire Naturelle des Araignées; 1875, Les Arachnides de France; and 1879, Arachnides de France (VII volume, analyze); also he had many descriptions of spiders collected by expeditions in Indochina, the Philippines, Greece, Syria, Yemen, Algeria, Egypt, and other countries. Simon described 40 families (out of the total of 117 accepted families in the world), many new genera, and 4650 species of spiders and other Arachnids (about 3790 species are still considered valid) more than any other arachnologist. E. Simon 20 2 Builders of Arachnology Koch Carl Ludwig (1778–1857) – German arachnologist. His principal works, Übersicht des Arachnidensystems, Nürnberg, 1837–1850; Deutschlands Crustaceen, Myriapoden und Arachniden, Regensburg, 1835–1844; and Die Arachniden (1831– 1848) (16 volumes), are the base of European arachnology. He is the father of another prominent arachnologist – Ludwig Carl Christian Koch (1825–1908). C.-L. Koch 2 Builders of Arachnology 21 Banks, Nathan (1868–1953) – American arachnologist and entomologist. Author of many descriptions of new genera and species in all orders of Arachnida, together with studies on Trichoptera and other insects. Nathan Banks 22 2 Builders of Arachnology Šilhavý, Vladimir (1913–1984) – Czech physician and prominent specialist in Opiliones. Working mostly on the fauna of the Caribbean, Mexico, the Balkans, and Central Asia, he described many new genera and species and wrote the book on Opiliones of Czechoslovakia (1956). Vladimir Šilhavý (Contributed by P. Bezdĕčka) 2 Builders of Arachnology 23 Hadži, Jovan (1884–1972) – Slovenian arachnologist and academician. He studied the scorpions, opilions, and pseudoscorpions of former Yugoslavia, describing many new species. His main arachnological works were Opilionidea. In: Catalogus Faunae Jugoslaviae, III/4 (1973b) and Neue Taxa der Weberknechte (Opilionidea) aus Jugoslawien (1973a), both published posthumously. Jovan Hadži (Slovenia Wikipedia) 24 2 Builders of Arachnology Petrunkevitch, Alexander (1875–1964) – American arachnologist of Russian origin (Ukrainian-born). From 1910 to 1939, he described over 130 spider species. He was a professor (since 1917) in Yale University, where he taught from 1910 to 1944. Many important papers on fossil Arachnida: A Monograph of the Terrestrial Paleozoic Arachnida of North America (1913); Tertiary Spiders and Opilionids of North America (1922); Inquiry into the Natural Classification of Spiders, Based on a Study of Their Internal Anatomy (1933); Palaeozoic and Mesozoic Arachnida of Europe (1953); Arachnida. In; Treatise on Invertebrate Paleontology (1955); Amber Spiders in European Collections (1958), papers on the spiders of Puerto Rico, of Mexico, etc. He described also many new families and higher taxa, including the valid fossil families Protosolpugidae, Kustarachnidae, Eotrogulidae, Nemastomoididae, Dolichophoninae, Archaeoctonidae, Centromachidae, Mazoniidae, Isobuthidae, and Calcitronidae. A. Petrunkevitch (Courtesy, Yale University) 2 Builders of Arachnology 25 Kratochvil, Josef (1909–1992) – Czech Zoologist and Academician, worked on Araneae, Opiliones, and Mammalia. He wrote 209 papers, including many on Arachnida (from 1930 to 1970), and several books. Kratochvil described many new taxa, especially from the caves of the Balkan Peninsula, including the genera Paralola and Tranteeva from Bulgarian caves (1958). He was a director of many scientific institutions. Josef Kratochvil 26 2 Builders of Arachnology Lawrence Reginald Frederick (1897–1987) – the most prominent researcher of South African Arachnida and Myriapoda. R. Lawrence published 23 papers on Opiliones (describing 224 valid species) and many new genera and species on all groups of Arachnida and Myriapoda. Reginald Frederick Lawrence 2 Builders of Arachnology 27 Vachon Max (1908–1991) – French specialist in Scorpiones and Pseudoscorpiones. Prof. Vachon was Director of the Laboratory of Invertebrates (Paris Museum of Natural History). Max Vachon 28 2 Builders of Arachnology Suzuki, Seisho (1914–2011) – During the period from 1938 to 1991, Dr. Suzuki wrote important series of 135 papers on Opiliones (Japan, Taiwan, Nepal, Malaysia, Philippines, Thailand, Sakhalin, etc.), with many discoveries and descriptions of valid species and several other taxa, also with zoogeographic analysis. He wrote also some papers of spiders. Suzuki Seisho 2 Builders of Arachnology 29 Gonzalez-Sponga Manuel-Angel (1929–2009) – Venezuelan arachnologist and professor. Described 219 valid species and many other taxa of Opiliones, many scorpions, and other Arachnids. Manuel-Angel Gonzalez-Sponga (Courtesy of R. Pinto-da-Rocha) 30 2 Builders of Arachnology Berlese Antonio (1863–1927) – Italian Acarologist who described thousands of new genera and species of mites. Antonio Berlese 2 Builders of Arachnology 31 Oudemans Anthonie Cornelis (1858–1943) – Dutch acarologist and one of the founders of the actual system of Acari. Anthonie Cornelis Oudemans 32 2 Builders of Arachnology Staręga Wojciech (1939–2015) – Polish arachnologist and specialist in Opiliones and spiders. He published many new taxa, the important revision of Bulgarian Opiliones (1976), catalogues, and articles on African Opiliones and also on the Opiliones of Russia, Central and East Asia, and other countries. W. Starega (Courtesy of J. Cokendolpher) Bibliography 33 Of course, the pioneers were (and still are) followed by many others, on a new level. Now data on Arachnida is snowballing, and the systems change overnight. With the new means of communication, we can now be on course to rapidly exchanging information, thanks also to some of our colleagues, who took care to facilitate our life by preparing catalogues of some orders and/or pages online. Such catalogues are listed in the Bibliography, but here we enumerated some of their authors: Brignoli P.M., Harvey M., Kury A., Giribet G., Cockendolfer J.C., Platnick N., Fet V., Deltshev Hr., Blagoev G., Ćurčić B., Martens J., Morano E., Michailov K., Juberthie Ch., Rambla M., Weigoldt P., Prendini L., Lourenço W., Drensky P., Gardini G., Saaristo M., Raven R.J., Lehtinen P., Rowland M., Dunlop J., Selden A.A., Thaler K., Wunderlich J.W., Shear W., and Muchmore W.B. Bibliography Birabén M (1949) Necrológio – Cândido F. de Mello-Leitão. Revista de la Socedad Entomologica Argentina. Buenos Aires 14(4):238–240 Buchar J (1984) Memory of MUDr. Vladimír Šilhavý, CSc. Acta entomologica bohemosl 81:157–158 Carpenter FM, Darlington PJ Jr (1954) Nathan Banks, a biographic sketch and list of publications. Psyche 61:81–110 Judson M, Chamberlin DC (1998) Joseph C. Chamberlin 1898 – 1962. J Arachnol 26:411–418 Kaltenbach A (1980) Hofrat Professor Dr. Max Beier zum Gedenken. Annalen des Naturhistorischen Museums Wien 83:763–781 Kury AB, Baptista RLC (2004) Arachnological papers published by Cândido Firmino de MelloLeitão (Arachnida). Publicações Avulsas do Museu Nacional, Rio de Janeiro 105:1–17 Muster C, Dunlop JA (2006) Le temps marche si vite – in memory of Konrad Thaler. J Arachnol 34:281–284 Osella G (1987) Professor Dr Paolo Marcello Brignoli (1942–1986). Bull Br Arachnol Soc 7(6):186 Prószynski J (1997) In memoriam Prof. Dr. Jan Rafalski (1909-1995). Proceedings of the 16th European Colloquium of Arachnology, Marek Zabka, editor. Siedlce, 8 – 9 Roesler R (1997/98) Karl Ludwig Koch (1778–1857). In Oberpfälzer Heimat. Auflage 42, Weiden Vigna Taglianti A (1986) Paolo Marcello Brignoli (1942 – 1986). Fragmenta Entomologica Roma 19(1):267–271 Chapter 3 Systems of Arachnida “Traditions die hard: it is difficult to understand why fishes or birds should be divided into dozens of orders, whereas, for most zoologists, a single order appears sufficient for all spiders or all mites. From this it could be inferred that either the arachnologists are more conservative or, in the eyes of most zoologists, small differences between (nice) birds are more significant than great differences between (ugly) spiders.” P.M. Brignoli (1980, The Evolution of Arachnida) Abstract Heymons (1901), the author of the term Chelicerata, united Merostomata (Eurypterids and Xiphosurans) with Arachnida in one monophyletic group. Dunlop and Selden (1997) restricted this term to Lemoneites, Xiphosura, Chasmataspida, Eurypterida, and Arachnida. There were many attempts to create a “natural” system of the monophyletic (Wheeler et al. 1993) class of Arachnida. Presently, it seems well established that the class Arachnida (Lamarck 1801) is composed of 10 recent orders plus 2–6 orders of “Acari” and variable number of fossil taxa. The seemingly undisputed orders are Palpigradi, Solifugae, Amblypygi, Uropygi, Schizomida, Ricinulei, Scorpiones, Pseudoscorpiones, Opiliones, and Araneae. The Acari have been (and still are) subject of various systems (see Beron 2008, 2011), more often considered as two (Acariformes and Parasitiformes) or three (+ Opilioacarida) orders. In the recent Manual of Acarology, Krantz and Walter and eds. (2009) agree on six orders of “Acari,” Mesostigmata, Holothyrida, Ixodida, Opilioacarida, Sarcoptiformes, and Trombidiformes, grouped in two superorders: Acariformes and Parasitiformes. Some authors (Stockwell 1989, Lourenço 2000) still raise some orders into subclasses or even classes (Opilionida, Scorpionida). Heymons (1901), the author of the term Chelicerata, united Merostomata (Eurypterids and Xiphosurans) with Arachnida in one monophyletic group. Dunlop and Selden (1997) restricted this term to Lemoneites, Xiphosura, Chasmataspida, © Springer International Publishing AG, part of Springer Nature 2018 P. Beron, Zoogeography of Arachnida, Monographiae Biologicae 94, https://doi.org/10.1007/978-3-319-74418-6_3 35 36 3 Systems of Arachnida Eurypterida, and Arachnida. There were many attempts to create a “natural” system of the monophyletic (Wheeler et al. 1993) class of Arachnida. Presently, it seems well established that the class Arachnida (Lamarck 1801) is composed of 10 recent orders plus 2–6 orders of “Acari” and variable number of fossil taxa. The seemingly undisputed orders are Palpigradi, Solifugae, Amblypygi, Uropygi, Schizomida, Ricinulei, Scorpiones, Pseudoscorpiones, Opiliones, and Araneae. The Acari have been (and still are) subject of various systems (see Beron 2008, 2011), more often considered as two (Acariformes and Parasitiformes) or three (+ Opilioacarida) orders. In the recent Manual of Acarology, Krantz and Walter and eds. (2009) agree on six orders of “Acari,” Mesostigmata, Holothyrida, Ixodida, Opilioacarida, Sarcoptiformes, and Trombidiformes, grouped in two superorders: Acariformes and Parasitiformes. Some authors (Stockwell 1989, Lourenço 2000) still raise some orders into subclasses or even classes (Opilionida, Scorpionida). The Higher and Highest Arrangements. The arrangement of Mello-Leitão (1931) in Portuguese: Arachnideos Subclass Ctenophoros Order Escorpioes Subclass Lipoctenos Sectio Patelados Susectio Megoperculados Order Pedipalpos Order Aranhas Dubinin (1959, announced first in 1957) divided the “superclass” Chelicerata into five classes: Merostomata, Scorpionomorpha, Solifugomorpha (with subclasses Holactinochitinosi and Pedipalpides), Arachnida (subclasses Opiliomorphae, Soluta, and Araneae), and Acaromorpha. Van der Hammen (1977) proposed to subdivide his new phylum Chelicerata into seven classes: Epimerata, Cryptognomae, Opilionidea, Apatellata, Arachnidea, Merostomata, and Scorpionidea. There is also a proposal by Savory (1977) to promote the suborder Cyphophthalmi into full order. This was not accepted. The Systems of Some Orders “Pedipalpi” – under this name have been united three orders of Arachnida: Amblypygi, Uropygi, and Schizomida. Amblypygi Millot (1949) – two families: Charontidae and Tarantulidae Quintero (1986) – created the suborders Apulvillata and Pulvillata and the family Charinidae 3 Systems of Arachnida 37 Suborder Apulvillata – families Phrynidae and Phrynichidae Suborder Pulvillata – families Charinidae and Charontidae Thelyphonida (Uropygi). Rowland and Cooke (1973) elevated the ancient Pocock’s (1899) subfamilies into families, thus three genera (Hypoctonus, Labochirus, and Thelyphonellus) in Hypoctonidae and the remaining 13 genera in Thelyphonidae. Presently, only one family is accepted, Thelyphonidae, which is divided into three subfamilies (Thelyphoninae, Hypoctoninae, and Typopeltinae). Scorpiones Stockwell (1989) – until this paper traditionally up to 8–9 families have been recognized. He raised the group to class with three orders (Protoscorpiones, Palaeoscorpiones, and Scorpiones). Lourenço (2000) – based on the classification of Stockwell (1989), he kept the scorpions as a class. Class Scorpionida Order Protoscorpiones Petrunkevich, 1949 Order Palaeoscorpiones Stockwell, 1989 Order Scorpiones C.L. Koch, 1837 Suborder Mesoscorpionina Stockwell, 1989 Suborder Neoscorpionina Thorell et Lindström, 1885 Infraorder Palaeosterni Stockwell, 1989 Infraorder Orthosterni Pocock, 1911 – includes the actual scorpions (20 recent families of the superfamilies Bothriuroidea, Buthoidea, Chaeriloidea, Chactoidea, Vaejovoidea, and Scorpionoidea) Fet et al. (2000) – their world catalogue includes 16 recent families and one fossil family in the suborder Neoscorpionina, infraorder Orthosternina, superfamily Scorpionidea, and the fossil suborder Branchioscorpionina. Prendini and Wheeler (2005) – 18 families. Prendini (2011a, b, c, d, e, f, g) – Order Scorpiones, one recent suborder Neoscorpionina with 23 families (four fossil, but there is a chance Akravidae to be found alive), so 19–20 recent families. Pseudoscorpiones (Chelonethi) – see the table in Harvey (1992, p.1376). Chamberlin (1931) – three suborders: Heterosphyronida, Monosphyronida, and Diplosphyronyda Beier (1932a, b) – three suborders: Chthonioidea, Neobisioidea, and Cheliferinea Harvey (1992, 2011, 2013a, b) – two suborders Epiocheirata and Iocheirata, with 24 families in 7 superfamilies, in total 26 families Suborder Epiocheirata Superfamily Chthonioidea (four families – Chthoniidae, Lechytiidae, Pseudotyrannochthoniidae, Tridenchthoniidae) 38 3 Systems of Arachnida Superfamily Feaelloidea (two families – Feaellidae, Pseudogarypidae) Suborder Iocheirata Infraorder Hemictenata Superfamily Neobisioidea (seven families – Neobisiidae, Bochicidae, Gymnobisiidae, Hyidae, Ideoroncidae, Parahyidae, Syarinidae) Infraorder Panctenata Superfamily Garypoidea (six families – Garypidae, Garypinidae, Geogarypidae, Larcidae, Menthidae, Olpiidae) Superfamily Cheiridioidea (two families – Cheiridiidae, Pseudochiridiidae) Superfamily Sternophoroidea (one familes – Sternophoridae) Superfamily Cheliferoidea (four families – Cheliferidae, Atemnidae, Chernetidae, Withiidae) Opiliones Thorell (1876a, b) – divided Opiliones into Palpatores and Laniatores Loman (1902) – proposed the term Insidiatores (new suborder including Triaenonychidae) Pocock (1902) – introduced the name Plagiostethi (= Palpatores), divided into Apagosterni and Eupagosterni. This was followed by Roewer (1910). Hansen and Sørensen (1904) – established a system of three suborders: Cyphophthalmi, Palpatores, and Laniatores. Palpatores were divided into the tribes Eupnoi and Dyspnoi. Mello-Leitão (1944) – proposed a phylogenetic tree of the Opiliones, noticing that there was a sister-group relationship between Cyphophthalmi and Palpatores Šilhavý (1961) – proposed subordinal rank of Eupnoi and Dyspnoi plus two more suborders Gonyleptomorphi and Oncopodomorphi Martens (1976) – The Palpatores are replaced by four superfamilies and the Laniatores by three: Palpatores – Troguloidea, Ischyropsalidoidea, Phalangoidea, and Caddoidea Laniatores – Travunioidea, Oncopodoidea, Gonyleptoidea Martens (1980 and further) – introduced the term Cyphopalpatores (Cyphophthalmi plus Palpatores). It was followed by Cokendolpher and Lee (1993) and other authors. Shear (1980, 1993) – divided suborder Cyphophthalmi into two infraorders with three superfamilies and six families: Suborder Cyphophthalmi Infraorder Temperophthalmi Superfamily Sironoidea Family Troglosironidae 3 Systems of Arachnida 39 Family Sironidae Family Pettalidae Infraorder Tropicophthalmi Superfamily Stylocelloidea Family Stylocellidae Superfamily Ogoveoidea Family Ogoveidae Family Neogoveidae This arrangement was accepted by Giribet (2000) and used since. Cokendolpher and Lee (1993) – used the term Cyphopalpatores but in their catalogue retained the names Cyphophthalmi and Palpatores Giribet et al. (2002) – introduced the term Dyspnolaniatores (Dyspnoi and its sister group Laniatores) Giribet and Kury (2007) – reviewed the phylogeny and biogeography of Opiliones and agreed to maintain the classical and generally accepted subdivision of the monophylic order Opiliones into Cyphophthalmi, Eupnoi, Dyspnoi, and Laniatores. We shall follow this arrangement in the present book. They used the old term Phalangida as a name of the clade formed by Eupnoi, Dyspnoi, and Laniatores. Their summary tree: Sironidae Pettalidae Troglosironidae Ogoveidae Neogoveidae Stylocellidae Caddoidea Phalangioidea Cyphophthalmi Eupnoi Ischyropsalidoidea Troguloidea Travunioidea Triaenonychoidea Zalmoxoidea Samooidea Phalangodoidea Epedanoidea. Gonyleptoidea. Dyspnoi Laniatores 40 3 Systems of Arachnida System of Opiliones in Kury (2013): Order Opiliones Sundevall, 1833 (4 suborders, 46 extant families, 3 fossil families, 1649 extant genera, 18 fossil genera, 6534 extant species, 37 fossil species) Suborder Cyphophthalmi (six families) Suborder Eupnoi Superfamily Caddoidea (one family, Caddidae) Superfamily Phalangioidea (four extant families Neopilionidae, Phalangiidae, Sclerosomatidae) – Monoscutidae, Suborder Dyspnoi Superfamily Ischyropsalidoidea (three families – Ischyropsalididae, Sabaconidae, Taracidae) Superfamily Troguloidea (four extant families – Dicranolasmatidae, Nemastomatidae, Nipponopsalididae, Trogulidae) Suborder Laniatores Infraorder Insidiatores Superfamily Travunioidea (three families – Nippononychidae, Paranonychidae, Travuniidae) Superfamily Triaenonychoidea (two families – Synthetonychiidae, Tryaenonychidae) Infraorder Grassatores Superfamily Assamioidea (two families – Assamiidae, Pyramidopidae) Superfamily Epedanoidea (five families – Epedanidae, Petrobunidae, Podoctidae, Sandokanidae, Tithaeidae) Superfamily Gonyleptoidea (seven families – Agoristenidae, Cosmetidae, Cranaidae, Gonyleptidae, Manaosbiidae, Stygnidae, Stygnopsidae) Superfamily Phalangodoidea (one family – Phalangodidae) Superfamily Samooidea (five families – Biantidae, Escadabiidae, Kimulidae, Samoidae, Stygnommatidae) Superfamily Zalmoxoidea (four families – Fissiphalliidae, Guasiniidae, Icaleptidae, Zalmoxidae) Araneae Pocock (1892) – basic terms in the classification of spiders Petrunkevitch (1939) – classification of Araneae Dubinin (1959) – subclass Araneae with three orders: Liphistiomorphae, Mygalomorphae, and Araneomorphae Platnick and Gertsch (1976) – order Araneae; their system is now followed. Suborder Mesothelae Pocock Suborder Opisthothelae Pocock Infraorder Mygalomorphae Pocock Infraorder Araneomorphae Smith Bibliography 41 Lehtinen (1978) – three groups without ranking: Theraphosomorpha, Araneomorpha, and Amaurobiomorpha Eskov and Zonshtein (1990) – order Araneida (Araneae) 1. Suborder Theraphosina (Orthognatha) 1.1. Infraorder Liphistiomorphae (family Liphistiidae) 1.2. Infraorder Theraphosomorphae (Mygalomorphae) 1.3. Infraorder Filistatomorphae (family Filistatidae) 2. Suborder Araneina (Labidognatha) 2.1. Infraorder Geralycosomorphae (labidognathous carboniferous “liphistiomorphs”) 2.2. Infraorder Dysderomorphae 2.3. Infraorder Hypochilomorphae (Hypochiloidea and Austrochiloidea) 2.4. Infraorder Araneomorphae Eskov and Zonshtein (1990) also group the seven infraorders into three grades distinguished by symplesiomorphies: Arthrabdominata, Sigillata, and Asigillata. Other sources: Bristowe (1938), Dubinin (1957 1959), Giribet et al. 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Zoologische Mededelingen, Leiden 51:307–319 van der Hammen L (1989) An introduction to comparative arachnology. SPB Academic Publishing, The Hague Weygoldt P, Paulus HF (1979a) Untersuchungen zur Morphologie, Taxonomie und Phylogenie der Chelicerata. 1. Morphologische Untersuchungen. Zeitschrift für Zoologische Systematik und Evolutionsforschung 17:85–116 Weygoldt P, Paulus HF (1979b) Untersuchungen zur Morphologie, Taxonomie und Phylogenie der Chelicerata. 2. Cladogramme und die Entfaltung der Chelicerata. Zeitschrift für Zoologische Systematik und Evolutionsforschung 17:177–200 Wheeler WC, Hayashi CY (1998) The phylogeny of the extant chelicerate orders. Cladistics 14:173–192 Wheeler WC, Cartwright P, Hayashi CY (1993) Arthropod phylogeny: a combined approach. Cladistics 9:1–39 World Spider Catalog (2017) World spider catalog. Natural history museum bern, online at http:// wsc.nmbe.ch, version 18.5, accessed on 27 Aug 2017. doi:10.24436/2 Chapter 4 The Fossil Arachnida Abstract Concerning the fossil orders, Dunlop (1996a, b) presents a revised concept of the formerly accepted (Weygoldt and Paulus 1979; Shultz 1990) five fossil orders of Arachnida. The subclasses of Petrunkevitch (1949) have been rejected, Anthracomartida – synonymized with Trigonotarbida. It was concluded that the position of Phalangiotarbida and Haptopoda remains obscure. Kustarachnida, according to Dunlop (1996a, b), represents misidentified opilionids. Concerning the fossil orders, Dunlop (1996a, b) presents a revised concept of the formerly accepted (Weygoldt and Paulus 1979; Shultz 1990) five fossil orders of Arachnida. The subclasses of Petrunkevitch (1949) have been rejected, Anthracomartida – synonymized with Trigonotarbida. It was concluded that the position of Phalangiotarbida and Haptopoda remains obscure. Kustarachnida, according to Dunlop (1996a, b), represents misidentified opilionids. According to the table in the paper of Dunlop et al. (2008), by March 2008, there were 1593 spp. of fossil Arachnida, including 111 Scorpiones, 25 Opiliones, 38 Pseudoscorpiones, 5 Solifugae, 294 “Acari,” 1 Palpigradi, 15 Ricinulei, 979 Araneae, 9 Amblypygi, 7 Uropygi, and 4 Schizomida (actually the Schizomida are 6). Extremly rich fossil fauna has been found in the Burmese amber. “Fossils from the mid-Cretaceous (c.99 Ma) Myanmar (Burma) amber include all extant orders of Arachnida, including the earliest representatives of Schizomida, Parasitiformes, and Palpigradi…The most abundant and diverse arachnid order is the Araneae, with 38 families, 93 genera, and 165 species recorded tom date” (Selden and Dong Ren 2017). Going through the recent analytical work on the position of different (recent) orders in the system of Arachnida, the following summary appears: Palpigradi Palaeokoenenia mordax Rowland et Sissom, 1980, from the late Tertiary (?Pliocene) of Arizona and the Electrokoenenia yaksha Engel et Huang from the Cenomanian Burmese amber remain the only known fossil Palpigrades. © Springer International Publishing AG, part of Springer Nature 2018 P. Beron, Zoogeography of Arachnida, Monographiae Biologicae 94, https://doi.org/10.1007/978-3-319-74418-6_4 45 46 4 The Fossil Arachnida Solifugae Only five fossil species of these large arachnids, living mostly in dry areas, are known (from Poland, USA, Brazil, and two from the Baltic and Dominican amber, respectively) (Poinar and Santiago-Blay 1989; Dunlop et al. 2004). Ricinulei Selden (1992) revised the fossil ricinuleids, describing 15 valid fossil species in 4 genera and 2 families from the Pennsylvanian Coal Measures of Europe and North America. Amblypygi Eight valid species of fossil whip spider are accepted by Dunlop et al. (2008): four from Europe and North America; two from Mexican and Dominican amber, respectively; another one from Mexican amber (Poinar and Brown 2004); and one from the Crato Formation in Brazil (Dunlop and Martill 2002). According to Dunlop and Mrugalla (2015), there are already 11 spp. The Late Carboniferous whip spider Graeophonus anglicus Pocock 1911, was redescribed by Dunlop et al. (2007a, b). The species Electrophrynus mirus Petrunkevitch 1971, was considered representing new genus and family, but both have been strongly contested by Armas (2006) and Dunlop and Mrugalla (2015). According to Selden et al. (1991), Amblypygi originated at least by the mid-Devonian (ca. 380 Ma). Presently, there are no Amblypygi living in Europe, and very few are known from North America, north of Rio Grande. Thelyphonida (Uropygi) According to Harvey (2003), there are nine species of fossil Uropygi, but after the recalculation of Tetlie and Dunlop (2008) and Dunlop et al. (2008), their number has been reduced to six to seven. Cai and Huang (2017) described from Upper Cretaceous Burmese amber the new genus and species of Thelyphonida Mesothelyphonus parvus - the earliest Thelyphoninae known so far. Schizomida Five fossil Schizomida genera have been described so far: Antillostenochrus Armas et Teruel and Rowlandius Reddell et Cokendolpher from the Miocene amber of the Dominican Republic; Calcitro Petrunkevitch from China, Oligocene, and the USA, Pliocene (Calcitronidae); and Onychothelyphonus Pierce and Calcoschizomus Pierce from the USA, Pliocene (Hubbardiidae) (Petrunkevitch 1945; Pierce 1951). From them, only Rowlandius contains extant species. Scorpiones The phylogenetic position of the scorpions remains a key question to resolve in studies of Chelicerate evolution (Weygoldt 1998; Dunlop and Braddy 2001). Scorpions are considered since longtime as the oldest known arachnids (Selden 1993a, b). Pocock (1893) divided the Arachnida into Ctenophora (scorpions) and Lipoptena (the remaining Arachnida). Dubinin (1962) unites Scorpiones and Eurypterida in a class (now clade) Scorpionomorpha. As scorpions are known as (sea ?) forms from the Silurian and the Devonian, very different from the extant forms, the dispute is between paleontologists and neontologists. The familial and suprafamilial classification of the scorpions is very controversial (see Soleglad and Fet 2003; Prendini and Wheeler 2005). The extant scorpions are assigned to the Orthosterni – a group or “parvorder” (Soleglad and Fet 2003) – known from the Carboniferous to the present. The number of superfamilies is not definitely established. 4 The Fossil Arachnida 47 Despite their hard cover, the fossil scorpions are relatively few in numbers. Fet et al. (2000), based on the monograph of Kjellesvig-Waering (1986), quote 78 spp. The further transformations and new descriptions from Baltic amber (Lourenco 2012), Burmese amber (Santiago-Blay et al. 2004), etc. increased this number to 111 by 2008 (Dunlop et al. 2008). The table in the last mentioned paper indicates one curious (and unique among Arachnida) particularity of fossil scorpions: 79 spp. are from Paleozoic age, 16 from Mesozoic, and 16 from Cenozoic. The older concept that scorpions are very primitive (partly because are very ancient) and are the ancestors of the other arachnids is strongly denied by cladists, which find (Shultz 1990) that they are derived arachnids forming a Dromopoda clade together with Opiliones, Solifugae, and Pseudoscorpiones. Some authors (Uchida 1966) still support the primitive character of the scorpions. The cladistic analysis of Dunlop and Braddy (2001) did not provide an undisputable end of the contest between the three main hypotheses: (1) Scorpions are the sister group of all other arachnids; (2) scorpions are derived group of arachnids as part of Dromopoda clade (and perhaps sister group of opilionids); and (3) scorpions are sister group of eurypterids. Support exists for any of the alternative hypotheses, based on morphological and molecular data and their interpretation. We can only admire the perception of the older authors (Pocock, Dubinin), which had in hand much less instruments for comparative analysis. Gess (2013) published “the earliest record of terrestrial animals in Gondwana” from South Africa. This was the new genus and species Gondwanascorpio emzantsiensis, a scorpion from a Famennian (Late Devonian) formation. Pseudoscorpiones In the catalogue of Harvey (1990) were listed 32 valid fossil pseudoscorpions from amber (Burmese, Chinese, Baltic, and Dominican). Schwaller et al. (1991) published the oldest pseudoscorpion from the mid-Devonian of Gilboa, New York State (the first Paleozoic pseudoscorpion). More details could be found in Harvey (1990) and Spahr (1993). Dunlop et al. (2008). After the new data of Hendericks (2005) and Judson (2007), respectively, from Baltic and Dominican amber, the total number of fossil pseudoscorpions amounts on 38 species (Dunlop et al. 2008). According to Harms and Dunlop (2017), overall 16 of the 26 [27?] families of living pseudoscorpions have been documented from fossils and 49 currently valid species are recognized. “The present-day distributions of some group (Faellidae and Pseudogarypidae) is relictual and highlights past extinction events” (Harms and Dunlop 2017). Harvey et al. (2017) described from the Burmese amber (mid-Cretaceous, ca. 90 Mya) the oldest member of Chthoniidae Weygoldtiella plausus Harvey et al. Opiliones The fossil record of this important group has been summarized by Dunlop (2007a, b). As we can see from the summary table in Dunlop’s paper, listing all known by this time fossil Opiliones, the fossil record of this group, containing now more than 6500 described species, consists of only 31 entries, part of them unnamed. Eight of them are from the Paleozoic (from Lower Devonian in Scotland to Upper Carboniferous in France). Only two taxa are recognized as genuine Mesozoic harvestmen (from the Lower Cretaceous in Australia and from Upper Cretaceous 48 4 The Fossil Arachnida Myanmar amber). None of them belongs to the modern families. The remaining 21 entries are Cenozoic and belong to the present-day families Trogulidae, Phalangiidae, Sclerosomatidae, Nemastomatidae, Caddidae, Kimulidae, Samoidae, and Cladonychiidae and are even assigned to the modern genera Trogulus, Kimula, Caddo, Dicranopalpus, Leiobunum, Nemastoma, Sabacon, etc. Sometimes these findings cast new light over the present-day distribution of families like Caddidae or Kimulidae and help in restoring the paleodistribution of the Opiliones. Actually, only 25 species are recognized as valid (Dunlop et al. 2008), with only one sp. from the Mesozoic. Ten of the findings come from the Baltic amber, considered to be of Eocene age (some 38–54 million years ago), mostly old information, summarized by Koch and Berendt (1854) and Menge (1854). The modern research in the Dominican amber by Cokendolpher (1986) and Cokendolpher and Poinar (1982, 1998) brought another four species. They belong to the family of Phalangodidae and Kimulidae, still living in the Antillean area. The age of this fauna is not very different from the age of Baltic amber fauna, but the research in the Dominican amber has much shorter history (for the harvestmen since 1978), and there is a lot more to expect. Only 14 fossil Opiliones species have been recorded before 1955, so the task of Petrunkevitch (1955), who summarized the existing scant information concerning fossil Opiliones, seems to be relatively simple. But it is amazing that these widespread animals are so rare in the fossil record (the reasons are explained by Dunlop, op. cit.). The oldest known Devonian harvestmen look very much like the modern forms, and it is a well-founded guess that there are also Silurian or other pre-Devonian Opiliones-like creatures, which are to be discovered. By Devonian (the Opiliones are considered to be at least 400 million years old), most of the known arachnid orders already existed as terrestrial, air-breathing animals. However, Garwood et al. (2014) described a new fossil suborder and concluded that “The discovery of Tetrophthalmi alters molecular divergence time estimates, supporting Carboniferous rather then Devonian diversification for extant suborders and directly impacting inference of terrestrialization history and biogeography.” The peculiar suborder Cyphophthalmi is represented by Siro platypedibus Dunlop et Giribet, 2003, a single specimen, described from Tertiary Bitterfeld amber, and by a new genus and species Palaeosiro burmanicum Poinar, 2008, from the Early Cretaceous Burmese amber (Upper Albian (100 to 105 m.y.B.P.) Araneae The bulk of Arachnida (the Acari excluded), both living over 40 000 spp. and fossil (979 spp.), belong to the order of spiders. A general review of the spider fossil record is provided by Penney and Selden (2006a, b). The oldest record is from Gilboa, New York (mid-Devonian, 380 Ma) (Selden et al. 1991). According to Dunlop et al. (2008), the vast majority of fossil spiders (c. 820 spp.) originate from amber: 540 spp. from the Baltic amber and 170 spp. from the Dominican amber. There are also findings in the younger (Cenozoic) amber and copal of France, Germany, Ukraine, China, etc. Spiders are described recently also from Cretaceous ambers in Siberia, Canada, Burma, the USA, and Spain (see Dunlop et al. 2008). Significant findings from the Cretaceous occurred also in Siberia and Mongolia (Eskov and Zonshtein 1990) and Spain (Selden and Penney 2003) and from the Crato Formation in Brazil (Selden et al. 2006). 4 The Fossil Arachnida 49 Acari As tiny creatures, mites are not many in the fossil record. After the publication of Krivolutsky (1973) about finding fossil Oribatida (including some recent genera (Cultroribula, Achipteria) from the Jurassic and Cretaceous deposits in Russia, it was accepted (Hammer and Wallwork 1979) that “families of oribatids that we can recognize today were present during the Jurassic and undoubtedly earlier.” It was proved some time later by Norton et al. (1988) with the finding of oribatids in the terrestrial Devonian deposit near Gilboa, New York. Describing the second fossil Opilioacarid, Dunlop et al. (2010a) analyzed the status of the fossil Acari by this time. According to them and to Dunlop et al. (2010b), 292 fossil actinotrichid species (including recent forms found as subfossils) have been recorded. Actinotrichids were certainly present by the Early Devonian (ca.410 Ma) and potentially much earlier. There are only 14 fossil species of anactinotrichids, and their record goes back to Cretaceous (Poinar and Buckley 2008) – two opilioacarids, four gamasids, no holothyrids, and eight ticks. The small number of fossil gamasids is really “puzzling.” Most genera of the fossil Gamasida and Ixodida are the same as in the extant fauna. Only in the Burmese amber have been found two new genera of Ixodidae (Cornupalpatum Poinar et Brown, 2003, and Compluriscutula Poinar et Buckley, 2008). Many years since this time, the fossil record of Oribatida is still poor (Dunlop et al. 2008). According to the table of Dunlop et al. (2008), by March 2008, there were known 294 “Acari.” Obviously, Oribatida are privileged as being preserved in the older rock layers (not amber), because of their hard armor. Such armor, even harder, has also the caeculids, but they are much smaller group, and only two species have been described recently from Baltic and Dominican amber (Eocene). From the Canadian amber has been described as new genus and species Proterythraeus southcotti Vercammen-Grandjean, 1972. From the Baltic amber have been recorded another new genus (Arythaena Menge, 1854) and eight other Erythraeidae species, assumed to belong to the present-day widespread genera Erythraeus, Balaustium, and Leptus. Provided that the material still exists, these identifications should be revised (Zacharda and Krivolutsky 1985). The fossil Mesostigmata are rare. From the Baltic amber (Eocene, ca. 44–49 Ma), Dunlop, Kontschán, and Zwanzig (2013) recorded Uropodina and Dunlop et al. (2014) Laelapidae of the extant genus Myrmozercon. Opilioacarida The first fossil opilioacarid to be described is Paracarus pristinus Dunlop, Wunderlich et Poinar, 2004, from Eocene Baltic amber. It has been attached to the genus Paracarus Chamberlin et Mulaik, 1942, represented by a modern species from Central Asia, and represents the northernmost species of Opilioacarida. The second fossil opilioacarid (? Opilioacarus aenigmus Dunlop, Sempf et Wunderlich, 2010a) was described again from Baltic amber (Eocene, 45–50 Ma). However, it has been assigned tentatively to the now Mediterranean genus Opilioacarus With, 1903. Paleozoogeography of Arachnida Important information concerning the past distribution of the Arachnida comes from the amber – Baltic, Dominican, Burmese, Lebanese, Chinese, etc. The amber is of different geological age. Other findings of 50 4 The Fossil Arachnida fossil Arachnida are from the Crato Member of the Santana Formation in Brazil (Martill et al., Eds, 2007), Kazakhstan, etc. Here are some data showing the differences between the past and present distribution of Arachnida: Palpigradi Remaining are the findings of Palaeokoenenia mordax Rowland et Sissom from the late Tertiary (?Pliocene) of the USA (Arizona) and of the Electrokoenenia yaksha Engel et Huang from the Cenomanian Burmese amber (Myanmar). Ricinulei The findings of the ancient group in the Pennsylvanian Coal Measures of Europe and North America (Nam.- Steph., 327–290 Ma) (Selden 1992) show a much broader distribution in the distant past. Actually, no ricinuleid lives in the Palearctic and (practically) in the Nearctic (only one sp. Pseudocellus dorotheae from Texas, on the border of Mexico). There are no extant Ricinulei also in Asia, but Wunderlich (2012) described? Poliocera cretacea from Burmese Cretaceous amber. Scorpiones There are at least 45 fossil families of scorpions (almost entirely from the suborder Branchioscorpionina). Solifugae Fossil species have been described not only from countries, where there are Solifugae even now (USA, Brazil), but also from Poland and the Baltic amber. Amblypygi Most of the known fossil Amblypygi come from countries where the order is represented now (Brazil, Mexico, Dominican Republic). However, four species are recorded from Europe and North America (Nam.-Steph., 327–290 Ma). Thelyphonida (Uropygi) Different genera from the now existing have been described from the Carboniferous of North America and Europe. Presently, Uropygi are not known from Europe (even closely) and from most of North America. Schizomida One representative of the order was found from the Oligocene of China. Presently, the only Schizomida known from this country is Bamazomus siamensis from Hong Kong, within the present range of the order. Opiliones There are fossil families of Opiliones, belonging to four recent suborders: Cyphophthalmi (Sironidae), Eupnoi (Phalangiidae), Dyspnoi (Caddidae, Nemastomatidae), and Laniatores. The material from Baltic amber (Eocene, ca. 44–49 Ma) has been presented by Koch and Berendt (1854) and supplemented or corrected by Roewer (1939), Dunlop and Barov (2005), and Dunlop and Mitov (2009). From the Bitterfeld amber (Germany, Oligocene–Chattian, 25.3–25.8 Ma) have been described the first fossil cyphophthalmid Siro platypodibus Dunlop et Giribet, 2003, and many other Opiliones, belonging to extant genera (Lacinius Thorell, Amilenus Martens, Mitostoma Roewer, ?Histricostoma Kratochvil, Caddo Banks, Dicranopalpus Doleschall, and even species like Lacinius erinaceus Staręga, known from Caucasus) (Dunlop and Mitov 2009). It is supposed that Opiliones of the extant genera and even species existed some tens of millions year ago (in the Paleogene). However, the Opiliones are much older and, as specialists say, can be Bibliography 51 traced back to at least the mid-Paleozoic. With the discovery of Caddo in North Europe, it was supposed even that the now discontinuous species of this genus (Japan and North America) have been spread in the Tertiary much broader. The fact that extant species have been recorded in the Oligocene, more than 20 Ma ago, raises the problem of the pace of evolution. Cokendolpher and Cokendolpher (1982) reexamined the tertiary harvestmen from Colorado. Araneae As it was said, the age of the fossil spiders is mostly from the Mesozoic, the oldest known being at 380 Ma (mid-Devonian of Gilboa). The Russian arachnologist Eskov (1984) used the finding of a member of “Gondwanan” family Archaeidae (Jurarchaea zherikhini) in the Upper Jurassic of Kazakhstan, as an argument against the attempts to explain many distributions with Gondwana. The actual distribution of the family (South Africa, Madagascar, Australia, Tasmania, New Zealand, and Patagonia) was considered by Legendre (1977) for “typically Gondwanan.” These spiders are known also from Baltic amber (seven species) and probably their present kind Gondwanan distribution is a relict one from a much more cosmopolitic distribution in the past. About the same time was published also a revision by Forster and Platnick (1984), leaving within Archaeidae only four genera from the Baltic amber, Australia, South Africa, and Madagascar. This does not change very much the conclusions of Eskov. Also, as Wunderlich (1995) pointed, Cyatholipidae are not “representing former parts of the Gondwanaland” (Griswold 1987) but rather a relict distribution of a very wide distribution in the Lower Tertiary period. Opilioacarida Paracarus pristinus from Baltic amber has been ascribed by Dunlop, Wunderlich, and Poinar (2004) to the genus Paracarus, represented today only by one species in Central Asia. The other fossil Opilioacarid (Opilioacarus (?) aenigmus Dunlop, Sempf et Wunderlich, 2010a) is conditionally included in the genus Opilioacarus, living today in the south of Europe (Italy, Greece) and in North Africa. Trombidiformes From Canadian amber has been described as new genus and species Proterythraeus southcotti Vercammen-Grandjean, 1972. 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Společnosti Zoologicke 49:147–152 Zherikin VV (1978) Development and succession of Cretaceous and Cenozoic faunistic complexes (Tracheata and Chelicerata). In: Proceedings of Paleontological Institute 165, 200 p (in Russian) Chapter 5 Factors Determining the Distribution of Arachnida Abstract As factors of distribution of Arachnida are outlined paleogeography and paleodistribution, age of groups, barriers, bridges, ability to overcome them, phoresy, dispersal, climate, orography and many other fundamental concepts. Zoogeography appears to be one of the most amusing and Stimulation of the natural sciences: every few years its fundamental concepts change and one can begin anew. (P.M. Brignoli 1983) 5.1 5.1.1 Some Basic Notions Zoogeography and Biogeography: The New Approach In the article of Posadas et al. (2006), a review is made of the development of the historical biogeography and the basic concepts of this science. The Argentinian authors never mention the word “zoogeography.” This is also the approach of most Western biogeographers. They also neglect entirely Russian and any other publication in non-Western languages. Most contemporary authors use the term “biogeography”; while the notions are generally the same in zoogeography and phytogeography, I think that when we have to deal with animals, it is entirely justified to use the word “zoogeography.” Basic review of the main concepts and ideas would be useful to the readers of this text. Most information derives from the analysis of Posadas et al. (2006). © Springer International Publishing AG, part of Springer Nature 2018 P. Beron, Zoogeography of Arachnida, Monographiae Biologicae 94, https://doi.org/10.1007/978-3-319-74418-6_5 59 60 5.1.2 5 Factors Determining the Distribution of Arachnida Centers of Origin and Dispersal Species have center of origin, from which some individuals disperse by chance and then evolve through natural selection (Darwin-Wallace traditional approach) (Matthew 1915). x x x Trying to explain the distribution of various animals, the older zoogeographers “discovered” many land bridges between continents and islands, even entire continents. With the development of modern explanations of these distributions (continental drift, etc.), most of these bridges became unnecessary and now have mostly historic value. They are explained in details by Müller (1974a, b). Paleantarctic This is a term created by Jeannel (1942) and means a continent, appearing in the Jurassic during the first breakup of Gondwana and the beginning of the “opening up” of Indian Ocean. Schuchert Land After Schuchert and Ihering, a North American-Pacific mountain range, connecting North with South America during the Cretaceous (from West to East from the Pacific Coast to the Missouri Basin). Lemuria Imaginary land bridge across the Indian Ocean, used by the older authors (Sclater, Ihering) to explain the connection of the fauna of Madagascar and India in the Cretaceous or early Tertiary. Beringia A land bridge (actually the notion is used to indicate a much larger dry land), connecting several times during the Tertiary and during the cold periods of the Pleistocene (Müller 1974a, b) Eurasia and North America. Through it massif exchange occurred of faunas and, finally, the penetration of Man in the Americas. Archiplata According to Ihering (1927), this was a land bridge in the place of today’s Andean South America. In Cretaceous time it connected the Archinotis with Schuchert land. This bridge was an attempt to explain the interchange of fauna between North and South America in the Tertiary. Archiguiana Supposed Cretaceous island in the region of Venezuela and the Guianas. 5.1 Some Basic Notions 61 Mesosoic Archhelenis Land bridge, connecting South America with South West Africa by way of Tristan da Cunha. Archatlantis Ihering (1927) coined this land bridge to connect the Antilles and Florida with North Africa and southern Spain and including the Azores, Canaries, and Cabo Verde islands. Archinotis Land bridge constructed by the older biogeographers to connect New Zealand and Australia with southern South America through Antarctica and the South Pacific islands (Cretaceous and early Tertiary). Canary Bridge Connection of the Eastern Canary Islands (Lanzarote) with the African mainland. Some biogeographers support this idea, under variations. After Sauer and Rothe (1972), “Separation of the eastern Canaries from Africa might have been by rifting and a land connection might still have existed in the lower Pliocene.” The shells found on the islands indicate that until about 12 million years ago, they were inhabited by large flightless birds. Tyrrhenian Bridge Supposed Riss-Glacial land bridge between Tuscany and Corsica and Sardinia. This bridge was invented in the efforts to explain mainly the distribution of herpetofauna. Galita Bridge Connecting Sardinia and Corsica with Africa. Balearic Bridge Connecting Corsica, Sardinia, and Balearic Islands with Spain. Provence Bridge Connecting Sardinia and Corsica with Provence. Buffon’s Law of Endemism Different areas, no matter how similar their physical conditions, generally house different species. 5.1.3 Development of Atlantic Ocean According to Sclater et al. (1977), the development of Atlantic Ocean is as follows: 165 Ma (Middle Jurassic) – The Atlantic Ocean is closed. The Baffin Bay is completely closed, and the shelf of Greenland fits neatly into the shelf of Labrador. Iberia is overlapping slightly onto the Atlas Mountains. To the South there is almost a perfect fit of the outline of South America and Africa. 62 5 Factors Determining the Distribution of Arachnida Map 5.1 The reconstructed position of the Atlantic continent 165 Ma (After Sclater et al. 1977, modified) 140 Ma (Late Jurassic–Early Cretaceous) – The main event between 165 and 140 Ma is the opening up of North Atlantic. 125 Ma (Early Cretaceous) – Between 165 and 140 Ma, Africa is still drifting away from North America along the Newfoundland fracture zone. By 125 Ma Iberia and Africa separate, and the North Atlantic is connected with Tethys. The protoCaribbean arc system has been as high and active as it is today. 110 Ma (boundary between Lower and Middle Cretaceous) – South America and Africa start to separate. The North and South Atlantic are still separated by part of Africa and South America. The elevated Walvis and Rio Grande Rise split the South Atlantic into two separate basins. 95 Ma (Middle Cretaceous) – Africa and Iberia continue to separate from North America. The rise between both parts of South Atlantic starts to sink, and the water will be able to flow from the South to the North through the Vema Gap. Europe starts separating from North America. 5.1 Some Basic Notions 63 80 Ma (Late Cretaceous–Senonian) – Greenland starts to move away both from Europe and North America. Iberia is well separated from Africa and is far from North America. 65 Ma (boundary between Cretaceous and Tertiary) – Europe is already far from North America, and Iberia is fixed to Europe. 53 Ma (boundary Paleocene–Eocene) – Baffin Bay is starting to open; Europe is separating much faster from Greenland and North America. The South Atlantic is already wide and deep. 36 Ma (boundary Eocene–Oligocene) – The Atlantic Ocean acquired almost its present shape. 21 Ma (Early Miocene), 10 Ma (Late Miocene) – Between the Oligocene and the Miocene, the Cabo Verde Plateau is uplifted, and the Azores-Gibraltar Ridge starts to form. 3.5 Ma – Uplift of the Isthmus of Panama and the waters of Pacific and Atlantic Oceans are separated. Map 5.2 The present Atlantic Ocean (After Sclater et al. 1977, modified) 64 5 Factors Determining the Distribution of Arachnida Conclusions 1. Since the Middle Jurassic (165 Ma) till the Lower Cretaceous (125 Ma), Northern Atlantic was completely closed basin. 2. Since the Lower Cretaceous (125 Ma) till Upper Cretaceous (110–95 Ma), Southern Atlantic was also completely closed basin. 5.1.4 Development of the Indian Ocean With a surface of 73,556,000 km², the Indian Ocean covers 20% of the total water surface of the planet. There are many scientific and semi-scientific attempts to analyze the India-Madagascar paleoconnections, the history of the massifs, and the remains of them as archipelagoes with most interesting fauna (Seychelles, Mascarene and others), the speculations concerning the hypothetic continent Lemuria, and many others. The arbitrary boundary between the Indian and Atlantic Oceans is following the 20°E meridian; along the meridian 146°55′ E runs the boundary with the Pacific. The average depth of Indian Ocean is 3890 m and the maximal 8047 m (Diamantina Trench). Speaking of “Sudamadie,” Jeannel (1965) is basing his conclusions on the two landmasses – the bloc South Africa-Madagascar (the continental part) and the remains of the so-called Subantarctic Plateau (the islands Crozet, Prince Edward, Kerguelen, and McDonald). Jeannel is sticking to the term Lemuria as a firm land, remains of which are the Comoro, Seychelles, and Mascarene Islands. He also accepts unconditionally the idea of a former bridge (land connection) between South Africa and South America as such connection is needed to explain the distribution of some groups of insects between the two continents. But the riddle still stays, as the analysis of Jeannel is based on the paleogeography of Furon (1959). Since this time many new data was obtained, modifying the zoogeographical conclusions. Now it is accepted that the African, Indian, and Antarctic plates converge in the Indian Ocean in the so-called Triple Point Rodriges. Lemuria of Jeannel is called now Mascarene Plateau (2000 km long underwater plateau East of Madagascar). Concerning the mid-ocean underwater ridges, there are several such active ridges, keeping in mind that Indian Ocean is the youngest of all oceans. In his works Jeannel pays special attention to Kerguelen Islands, where deep layers of fossil plants are found from Eocene and Oligocene. According to Jeannel, this area has been an extensive land with rivers and forests, similar to those existing now in Tierra del Fuego. Now is considered that Kerguelen Archipelago (7215 km2, almost the size of Corsica) is situated on a sunken microcontinent having a surface of 2.2 million of km2. The development of Indian Ocean has been outlined in details according to modern geophysical data by Ali and Aitchison (2008). 5.2 Paleogeography and the Past Distribution of Arachnida 5.2 5.2.1 65 Paleogeography and the Past Distribution of Arachnida Age of the Faunas Here seems useful to remind the reader the time span of the different periods (BP). The oldest and the predominant record of the different groups of arachnids is as follows: Palpigradi Palaeokoenenia mordax Rowland et Sissom is from the late Tertiary (?Pliocene). The Pliocene extended from 5.332 million to 2.588 million years before present. Engel et al. (2016) published from Cenomanian (mid-Cretaceous) Burmese amber a much older (94–97 My old) member of Palpigradi (Electrokoenenia yaksha Engel et Huang), the first Mesozoic fossil of this order. Solifugae Five fossil species known (from Poland, USA, Brazil and from the Baltic and Dominican amber, respectively). The age of Baltic amber is 44 million years ago (Eocene), of Dominican amber – 25–15 Ma. Ricinulei Known from the Pennsylvanian Coal Measures of Europe and North America (Nam.- Steph., 327–290 Ma). Wunderlich (2012) described a new fossil species from the Cretaceous of Burma (Myanmar). Amblypygi There are nine valid species of fossil whip spider: – – – – Four from Europe and North America (Nam.-Steph., 327–290 Ma) Three from Mexican amber (30 million years) One from Dominican amber (15–45 million years) One from the Crato Formation of Brazil (Early Cretaceous, mostly during the early Albian age, about 108 million years ago) Thelyphonida (Uropygi) There are 7–8 fossil Uropygi from Brazil (Mesoproctus rowlandi Dunlop, 1998, Lower Cretaceous (Aptian) of the Crato Member of the Santana Formation) and from Carboniferous of North America and Europe (Geralinura, Proschizomus, Parageralinura, Protelyphonus). A new genus and species (Mesothelyphonus parvus Cai et Huang, 2017) has been described from Burmese amber (Cretaceous). Schizomida Three fossil Schizomida have been described from Arizona, USA, one from the Oligocene of China (the Oligocene epoch is from 34 to 23 million years BP) and two from Dominican amber (Krüger and Dunlop 2010). Scorpiones Known from the Silurian and the Devonian (sea forms ?), scorpions are considered since longtime as the oldest known arachnids. The extant scorpions are assigned to the Orthosterni, known from the Carboniferous to present. By 2008 there were known 111 spp. of fossil scorpions (79 spp. are of Paleozoic age, 16 of Mesozoic, and 16 of Cenozoic). According to Prendini (2011), fossil scorpions include 45 families, 86 genera, and 120 species (Akravidae being considered by the present author as recent). They belong to the extinct suborder 66 5 Factors Determining the Distribution of Arachnida Branchioscorpionina Kjellesvig-Waering, 1986 (42 families), and to suborder Neoscorpionina Thorell et Lindström, 1885 (three families). Pseudoscorpiones Most of the 38 fossil species of pseudoscorpions come from amber (Burmese, Chinese, Baltic, and Dominican). Schawaller et al. (1991) published the oldest pseudoscorpion from the mid-Devonian (Paleozoic) of Gilboa, New York State (380 Ma), the only fossil family of Pseudoscorpiones (Dracochelidae; Schawaller, Shear et Bonamo, 1991). Opiliones Eight entries in the table of Dunlop (2007) are from the Paleozoic (from Lower Devonian in Scotland to Upper Carboniferous in France). Two taxa are Mesozoic (from the Lower Cretaceous in Australia and from Upper Cretaceous Myanmar amber). The remaining 21 entries are Cenozoic and belong to the present-day families. Ten of the findings come from the Baltic amber of Eocene age (some 38–54 Ma years ago), mostly old information. The different beds of Dominican amber have been dated as Miocene (15–20 million years) to Upper Eocene (ca. 30–45 million years). So, the age of this fauna is not very different from the age of Baltic amber fauna, but the research in the Dominican amber has much shorter history (for the harvestmen since 1978), and there is a lot more to expect. It is of note the description of a new genus and species of Cyphophthalmi (Palaeosiro burmanicum) by Poinar (2008) from the Burmese amber of Upper Albian (100–105 my BP). The oldest known Devonian harvestmen, considered to be at least 400 million years old, look very much like the modern forms, and it is a well-founded guess that there were also Silurian or other pre-Devonian Opiliones-like creatures, which are to be discovered. Garwood et al. (2014) described a new fossil suborder Tetrophthalmi, based on two fossil species – the new Carboniferous harvestman Hastocularis argus and the Devonian Eophalangium sheari. This finding supports Carboniferous rather than Devonian diversification for extant suborders of Opiliones. Araneae The vast majority of fossil spiders (c. 820 spp.) originate from amber: 540 spp. from the Baltic amber and Dominican amber c.170 spp. (Miocene to Upper Eocene (15–20 million years to ca. 30–45 million years). There are also findings in the younger (Cenozoic) amber and copal of France, Germany, Ukraine, China, etc. Spiders have been described recently also from Cretaceous ambers in Siberia, Canada, Burma, the USA, Spain, and Mongolia and from the Crato Formation of Brazil. It means that the age of the fossil spiders is mostly from the Mesozoic, the oldest known being at 380 Ma (mid-Devonian of Gilboa). Acari Most of the Acari (except of Opilioacarida and Holothyrida) are not so important zoogeographically, many of them being parasitic, and the others are not adequately studied and are not treated in details in this publication. 5.2 Paleogeography and the Past Distribution of Arachnida 67 Opilioacarida Two fossil spp. have been described from Baltic amber (44 million years ago, Eocene): Opilioacarus (?) aenigmus Dunlop, Sempf et Wunderlich, 2010, and Paracarus pristinus Dunlop, Wunderlich et Poinar, 2004. Acariformes Trombidiformes Among the other Arthropoda from Baltic amber by Koch and Berendt (1854) have been recorded eight spp. of Erythraeidae (ascribed to “Rhyncholophus,” now Erythraeus, Balaustium, and Leptus). Menge (1854) described from Baltic amber the new genus and species Arythaena troguloides; Vercammen-Grandjean (1972) found in Canadian amber (Cretaceous) another new genus and species: Proterythraeus southcotti. Zacharda (in Zacharda and Krivolutsky 1985) described Mesoanystis taymirensis from Taymyr Peninsula, Upper Cretaceous amber (the oldest fossil anystid mite known), and Palaeoerythracarus sachalinensis from Sachalin (Paleogene amber). Bartel et al. (2015) described three new spp. of Fessonia (Smarididae) from Baltic amber, and Calyptostoma katyae Koniktyiewicz, Wohltmann et Makol, 2016, was the first fossil Calyptostoma (described also from Baltic amber). Dunlop (2007) described from the Early Cretaceous (Aptian) Crato Formation from Ceará State, Brazil, a new large, fossil mite (Arachnida: Acari, Erythraeoidea), Pararainbowia martilli. Sarcoptiformes Oribatida From many places have been recorded fossil oribatid mites, because of the hard tegument and their massif presence (up to 75%) in the soil fauna (Krivolutsky and Druck 1986). Norton et al. (1988) recorded 14 fossil Oribatid spp. (including the new genera Protochthonius and Devonacarus and the new family Devonacaridae) from terrestrial Devonian deposit near Gilboa, New York. An acarid belonging to the still-living genus Hydrozetes Berlese, 1902, was found in the Lower Sinemurian (Jurassic) in a core drilling from Sweden (Sivhed and Wallwork 1978). Parasitiformes Mesostigmata Inclusions assignable to the tortoise mites (Mesostigmata, Uropodina) have been described by Dunlop et al. (2013) from Baltic amber (Eocene, ca. 44–49 Ma). This is the oldest record of Uropodina. “Fossil mesostigmatid mites (Acari: Parasitiformes: Mesostigmata) are extremely rare, and specimens from only nine families, including four named species, have been described so far. A new 68 5 Factors Determining the Distribution of Arachnida record of Myrmozercon sp. described here from Eocene (ca 44–49 Myr) Baltic amber represents the first – and so far only – fossil example of the derived, extant family Laelapidae”. Ixodida In the amber from Cretaceous has been recorded a larval argasid tick (Carios jerseyi Klompen et Grimaldi, 2001). Another Mesozoic tick (Amblyomma, Ixodidae) has been described from Dominical amber (Lane and Poinar Jr. 2009), the first in the New World. Ixodes tertiarius Scudder, 1885, was described from Oligocene amber (Wyoming, USA). Another tick (Ixodes succineus Weidner, 1964) is known from the Baltic amber. We borrowed from the paper of Giribet, Sharma et al. (2012) this chronology: Geological intervals: – 0–35 Ma (separation of Mediterranean plate from Western Europe, separation of Japan from Eurasia, connection of Iberia to Eurasia) – 35–45 Ma (separation of Borneo and Indomalay Archipelago from Eurasia) – 45–60 Ma (Balkans connected to Western Europe, Iberia connected to Mediterranean plate, Balkans, and Japan). – 60–75 Ma (Iberia separated from Mediterranean plate, Balkans, and Japan, North America separated from Western Europe, emergence of Indomalay Archipelago) – 75–110 Ma (Mediterranean plate separated from North America, Iberia connected to Western Laurasia, Balkans separated from North America and Western Europe) – 110–120 Ma (Iberia disconnected from other landmasses; Western Europe, Mediterranean plate, and North America separated from Eastern Laurasia; emergence of Borneo; Indomalay Archipelago nonexistent) – 120–180 Ma (Iberia disconnected from other landmasses; Western Europe, Mediterranean plate, and North America separated from Eastern Laurasia; Borneo and Indomalay Archipelago nonexistent) – 180–250 Ma (Thai-Malay Peninsula disconnected from other landmasses; Eastern Himalaya disconnected from North America, Western Europe, and Iberia; Borneo and Indomalay Archipelago nonexistent) – 250–296 Ma (Borneo and Indomalay Archipelago nonexistent, other landmasses connected) Here seems useful to remind the reader the time span of the different periods (BP). Silurian – 444–416 Ma (roughly, other figures exist in different sources). Known scorpions (sea forms?) Devonian – 416–359 Ma. Known scorpions, spiders, and pseudoscorpions (midDevonian, 380 Mya) Carboniferous – 359–299 Ma. Known scorpions Orthosterni (the presently existing scorpions) Permian – 299–251 Ma Triassic – 251–200 Ma Jurassic – 200–146 Ma 5.2 Paleogeography and the Past Distribution of Arachnida 69 Cretaceous – 146–66 Ma Tertiary – 66–2.6 Ma Paleogene (Paleocene, Oligocene, Eocene) – 66–23 Ma Neogene (Miocene, Pliocene) – 23 – 2.6 Ma Quaternary – 2.6 Ma to the present Pleistocene – 2.6–0.012 Ma Holocene – 0.012 Ma to the present Palaeokoenenia mordax Rowland et Sissom is from the late Tertiary (?Pliocene). The Pliocene extended from 5.332 million to 2.588 million years before present. Engel et al. (2016) published from Cenomanian (mid-Cretaceous) Burmese amber a much older (94–97 My old) member of Palpigradi (Electrokoenenia yaksha Engel et Huang), the first Mesozoic fossil of this order. Solifugae Five fossil species known (from Poland, USA, and Brazil and from the Baltic and Dominican amber, respectively). The age of Baltic amber is 44 million years ago (Eocene), of Dominican amber – 25–15 Ma. Ricinulei Known from the Pennsylvanian Coal Measures of Europe and North America (Nam.- Steph., 327–290 Ma). Wunderlich (2012) described a new fossil species from the Cretaceous of Burma (Myanmar). Amblypygi There are nine valid species of fossil whip spider: – – – – Four from Europe and North America (Nam.-Steph., 327–290 Ma) Three from Mexican amber (30 million years) One from Dominican amber (15–45 million years) One from the Crato Formation of Brazil (Early Cretaceous, mostly during the early Albian age, about 108 million years ago) (Uropygi) Known from the Silurian and the Devonian (sea forms ?), scorpions are considered since longtime as the oldest known arachnids. The extant scorpions are assigned to the Orthosterni, known from the Carboniferous to present. By 2008 there were known 111 spp. of fossil scorpions (79 spp. are of Paleozoic age, 16 of Mesozoic, and 16 of Cenozoic). According to Prendini (2011), fossil scorpions include 45 families, 86 genera, and 120 species (Akravidae being considered by the present author as recent). They belong to the extinct suborder Branchioscorpionina Kjellesvig-Waering, 1986 (42 families) and to suborder Neoscorpionina Thorell et Lindström, 1885 (three families). Eight entries in the table of Dunlop (2007) are from the Paleozoic (from Lower Devonian in Scotland to Upper Carboniferous in France). Two taxa are Mesozoic (from the Lower Cretaceous in Australia and from Upper Cretaceous Myanmar amber). 70 5 Factors Determining the Distribution of Arachnida The remaining 21 entries are Cenozoic and belong to the present-day families. Ten of the findings come from the Baltic amber of Eocene age (some 38–54 Ma years ago), mostly old information. The different beds of Dominican amber have been dated as Miocene (15–20 million years) to Upper Eocene (ca. 30–45 million years). So, the age of this fauna is not very different from the age of Baltic amber fauna, but the research in the Dominican amber has much shorter history (for the harvestmen since 1978), and there is a lot more to expect. Amber of Upper Albian (100 to 105 my BP). The oldest known Devonian harvestmen, considered to be at least 400 million years old, look very much like the modern forms, and it is a well-founded guess that there were also Silurian or other pre-Devonian Opiliones-like creatures, which are to be discovered. Garwood et al. (2014) described a new fossil suborder Tetrophthalmi, based on two fossil species – the new Carboniferous harvestman Hastocularis argus and the Devonian Eophalangium sheari.This finding supports Carboniferous rather than Devonian diversification for extant suborders of Opiliones. Araneae The vast majority of fossil spiders (c. 820 spp.) originate from amber: 540 spp. from the Baltic amber and Dominican amber c.170 spp. (Miocene to Upper Eocene, 15–20 million years to ca. 30–45 million years). There are also findings in the younger (Cenozoic) amber and copal of France, Germany, Ukraine, China, etc. Spiders are described recently also from Cretaceous ambers in Siberia, Canada, Burma, the USA, Spain, and Mongolia and from the Crato Formation in Brazil. It means that the age of the fossil spiders is mostly from the Mesozoic, the oldest known being at 380 Ma (mid-Devonian of Gilboa). Acari Most of the Acari (except of Opilioacarida and Holothyrida) are not so important zoogeographically, many of them being parasitic, and the others are not adequately studied and are not treated in details in this publication. Opilioacarida Two fossil spp. have been described from Baltic amber (44 million years ago, Eocene): Opilioacarus (?) aenigmus Dunlop, Sempf et Wunderlich, 2010, and Paracarus pristinus Dunlop, Wunderlich et Poinar, 2004. Trombidiformes Among the other Arthropoda from Baltic amber by Koch and Berendt (1854) have been recorded eight spp. of Erythraeidae (ascribed to “Rhyncholophus,” now Erythraeus, Balaustium, and Leptus). Menge (1854) described from Baltic amber the new genus and species Arythaena troguloides; Vercammen-Grandjean (1972) found in Canadian amber (Cretaceous) another new genus and species: Proterythraeus southcotti. Zacharda (in Zacharda and Krivolutsky 1985) described Mesoanystis taymirensis from Taymyr Peninsula, Upper Cretaceous amber (the oldest fossil anystid mite known), and Palaeoerythracarus sachalinensis from Sachalin (Paleogene amber). Bartel et al. (2015) described three new spp. of Fessonia (Smarididae) from Baltic Amber, and Calyptostoma katyae Koniktyiewicz, Wohltmann et Makol, 2016, was the first fossil Calyptostoma (described also from Baltic amber). Dunlop (2007) described from the Early Cretaceous (Aptian) Crato 5.2 Paleogeography and the Past Distribution of Arachnida 71 Formation from Ceará State, Brazil, a new large, fossil mite (Arachnida: Acari, Erythraeoidea), Pararainbowia martilli. Sarcoptiformes Oribatida From many places have been recorded fossil oribatid mites, because of the hard tegument and their massif presence (up to 75%) in the soil fauna (Krivolutsky and Druck 1986). Norton et al. (1988) recorded 14 fossil oribatid spp. (including the new genera Protochthonius and Devonacarus and the new family Devonacaridae) from terrestrial Devonian deposit near Gilboa, New York. An acarid belonging to the stillliving genus Hydrozetes Berlese, 1902, was found in the Lower Sinemurian (Jurassic) in a core drilling from Sweden (Sivhed and Wallwork 1978). Parasitiformes Mesostigmata Inclusions assignable to the tortoise mites (Mesostigmata, Uropodina) have been described by Dunlop et al. (2013) from Baltic amber (Eocene, ca. 44–49 Ma). This is the oldest record of Uropodina. “Fossil mesostigmatid mites (Acari: Parasitiformes: Mesostigmata) are extremely rare, and specimens from only nine families, including four named species, have been described so far. A new record of Myrmozercon sp. described here from Eocene (ca 44–49 Myr) Baltic amber represents the first - and so far only - fossil example of the derived, extant family Laelapidae”. Ixodida In the amber from Cretaceous has been recorded a larval argasid tick (Carios jerseyi Klompen et Grimaldi, 2001). Another Mesozoic tick (Amblyomma, Ixodidae) has been described from Dominical amber (Lane and Poinar Jr. 2009), the first in the New World. Ixodes tertiarius Scudder, 1885 was described from Oligocene amber (Wyoming, USA). Another tick (Ixodes succineus Weidner, 1964) is known from the Baltic amber. We borrowed from the paper of Giribet, Sharma et al. (2012) this chronology: Geological intervals: – 0–35 Ma (separation of Mediterranean plate from Western Europe, separation of Japan from Eurasia, connection of Iberia to Eurasia) – 35–45 Ma (separation of Borneo and Indomalay Archipelago from Eurasia) – 45–60 Ma (Balkans connected to Western Europe, Iberia connected to Mediterranean plate, Balkans, and Japan) – 60–75 Ma (Iberia separated from Mediterranean plate, Balkans, and Japan, North America separated from Western Europe, emergence of Indomalay Archipelago) 72 5 Factors Determining the Distribution of Arachnida – 75–110 Ma (Mediterranean plate separated from North America, Iberia connected to Western Laurasia, Balkans separated from North America and Western Europe) – 110–120 Ma (Iberia disconnected from other landmasses; Western Europe, Mediterranean plate, and North America separated from Eastern Laurasia; emergence of Borneo; Indomalay Archipelago nonexistent) – 120–180 Ma (Iberia disconnected from other landmasses; Western Europe, Mediterranean plate, and North America separated from Eastern Laurasia; Borneo and Indomalay Archipelago nonexistent) – Dunlop et al. (2008), Dunlop and Martill (2002), Dunlop et al. (2004), Eskov and Zonshtein (1990), Harvey (1990, 2003), Judson (2007), Kjellesvig-Waering (1986), Krivolutsky and Krasilov (1977), Penney and Selden (2007), Poinar and Brown (2004), Poinar and Santiago-Blay (1989), Rowland and Sissom (1980), Selden (1993), Selden et al. (1991), Soleglad and Fet (2003), Tetlie and Dunlop (2008) 5.3 5.3.1 Passive Dispersal and Phoresy of Arachnida: Ballooning, Rafting, and Continental Drift Dispersal (Natural and by Man Activities) Dispersal and Dispersion The difference between these two notions, sometimes used indiscriminately, was explained by Platnick (1976): “Dispersion would be defined as a property of individuals, the process by which an organism is able to spread from its place of origin to another locality. Dispersal would be defined as a property of taxa, the process by which a species or other group is able to spread from its previously established range to a different range.” Dispersal was and remains one of the major problems of zoogeography and a rather controversial one. From the extreme fixists (stabilists) to the extreme mobilists, one can observe the whole range of approach of the zoogeographer. Despite the numerous achievements of earth science, the extreme fixists still believe that the patterns of the land-sea distribution over the globe remain unchanged since the third day of Creation. Their opponents, it is true, sometime deal freely with entire continents, displacing them to fit some particular distribution. The fixists like to refer to the saying of L. Vialleton (1929, L’Origine des êtres vivants): “On a joué des dislocations géologiques ou de l’existence de continents supposés au point de pouvoire faire dire que certains biogéographes n’hesiteraient pas à inventer un continent pour expliquer un genre de Coléoptère.” Many and many cases of distribution, however, can be explained only by admitting direct land connections. To say that the Galapagos and Aldabra tortoises or the moa of New Zealand have reached these islands riding logs is, to put it mildly, not serious. The “stepping stones” could have had significance with some groups, but the present author is firmly convinced that 5.3 Passive Dispersal and Phoresy of Arachnida: Ballooning, Rafting, and Continental… 73 even a gap of 10 meters of salt water is enough to prevent the dispersal of many animal species and even groups. The land invertebrates are often thought to disperse easier than the land vertebrates. This is probably true (to some extend) for some groups of insects. Again, how to explain the high endemism of insect fauna of Madagascar? There are many groups not dispersing by floating logs. Yes, tropical rivers do carry sometimes logs and even “floating islands” far into the ocean, but none of these “rafts” has brought Uropygi (abundant in Southeast Asia) or many Papuan groups to Australia. Isopods use to live in rotten logs, but nevertheless the fauna of Cuba is strictly neotropical (the proximity of North America hasn’t affected it) (Vandel 1972a, b), and only two widespread genera of woodlice are common to both Australia and Melanesia. As Vandel (1973b) emphasizes: “Encore que l’Australie et la Nouvelle-Guinée soient, au temps present, fort proche l’une de l’autre, leurs faunes isopodiques ne presentent à peu près aucun point commun.” The remaining 20 genera and all the 53 species of Melanesian Oniscidea have preferred to not undertake the risky rafting across the 150 km-wide Torres Strait (the same is true for the numerous Australian genera of Isopoda not represented in Melanesia). According to Szymkowiak et al. (2007), “Despite similar geological history and close vicinity of both areas, the crab spiders show great distinction between New Guinea and Australia. 36 genera on both areas have been recorded, of which only 13 are common (Amyciaea, Cymbacha, Diaea, Hedana, Loxoporetes, Mastira, Misumena, Porropis, Runcinia, Stephanopis, Tharrhalea, Tmarus, and Xysticus), and 167 species, of which 6 are common. Thus, similarity of the spider fauna of the family Thomisidae at genera level is near 37%, while at species level it is 3.7%.” Even if “to account for the endemic elements of the Hawaiian fauna would require only about one successful establishment every 20,000 years or more”(Gressitt 1971a, b), it means that some of the groups of land animals do not disperse by air or floating logs at all or meet with considerable difficulties in establishing themselves on the new place. These “conservative” groups are of special interest for the zoogeographer, as they reflect the ancient state of the fauna of the particular area. On the list of such “chosen” groups, the Arachnida stay very high, especially some of them. A thorough review of the passive dispersal of Arachnida is due to Szymkowiak et al. (2007). In the nineteenth and twentieth centuries, voluntary or involuntary transport by Man became a major factor in the dispersal of plants and mammals. Again the number of species of Arachnida, using ships or aircraft, is rather limited, and establishment of Man-carried individuals is seldom successful. There are differences from order to order. As Wallwork (1979) points out, it is generally recognized as a principle of zoogeography that random dispersal by wind, water, birds, or Man results in disharmonic faunas. Other sources: Berland (1934), Bristowe (1930), Cowrie and Holland (2006), Darlington (1938), Duffey (1998), Gressit and Yoshimoto (1963), Schatz (1991), Udvardy (1981) 74 5 Factors Determining the Distribution of Arachnida Opiliones Studying the phalangids of the Antilles, Šilhavý (1973) indicates that these animals are “from the phylogenetical point of view a very old order with small ecological valence, conforming with difficulty to changing life conditions and being very sensitive to the shortage of water and not very mobile. Phalangids cannot move by “ballooning” as some spiders can and their possibility of migration among the islands has been minimal even if we admit the possibility of their passive transfer by hurricanes (either as living specimens or as eggs).” This opinion of the leading specialist in Opiliones confirms our view about the limited role of passive dispersal in this group. One very discussed case is the presence in New Caledonia of endemic, relic, and very old (> 200 Ma) family of cave Opiliones of Cyphophthalmi (Troglosironidae, 13 spp.). They are close not to the nearest families (Pettalidae, Sironidae, Stylocellidae) but to the far away Neogoveidae, living in the Americas. Sharma and Giribet (2012) discussed the possible explanation of this relation. Cyphophthalmi are poor dispensers, absent in the oceanic islands, and they certainly cannot overcome marine water expanses. Solifugae Among the 13 000 ship-trapped specimens of insects and arachnids, Holzapfel and Harrell (1968) mention 275 specimens of spiders (Araneae), and among them one specimen of “Solpugida” is probably a mistake. The easternmost localities of Solifugae are in Vietnam (Dinorhax rostrumpsittaci Simon, fam. Melanoblossiidae). They don’t live in apparently suitable Australia and are not found east of Wallace’s Line. The “sun spiders” don’t seem to be very successful candidates for “rafting,” let alone “ballooning.” Good example is their total absence in Madagascar, despite the many species living in Southern Africa. Pseudoscorpiones The tiny false scorpions, living under the bark or in the soil and plant debris, should be transported easily by floating logs, by phoresy, or by human activities. However, apart from the several cosmopolitic species and some store and manure-inhabiting species, there is rather limited number of pseudoscorpions really carried to faraway places. Even with the intense Europe-North America traffic, such examples are rare (Muchmore 1969, 1972). Pseudoscorpions sometime travel attached to birds or mammals (phoretic) (Martens 1975). Seabirds have carried them to the subantarctic of very isolated islands like St. Helena, Ascension, or Tristan-da-Kunha. If we compare two relatively close and well-known faunae like those of Australia and New Zealand (58 and 25 local genera, respectively), we shall see that they have only ten genera and no species in common. Amblypygi Sometime people write amazing things. To quote Besch (1969), “The Charontidae, an Indo-pacific group, are represented in South America only by the genus Charinus, which except for its occurrence in the Galapagos Islands, is found only in the islands of the Indian and Pacific Oceans. It is, therefore, considered to have come from these islands to Galapagos Archipelago.” The author of this highly 5.3 Passive Dispersal and Phoresy of Arachnida: Ballooning, Rafting, and Continental… 75 improbable hypothesis evidently has overlooked the publication of Goodnight and Goodnight (1946), describing Charinus Gertschi from Guiana and Suriname. The subsequent founds of Charinus in Brazil (eight spp.), Venezuela, Peru, and many other countries rather confirm the second alternative of the Goodnights that the discovery of Charinus in Guiana “indicates our lack of knowledge of the neotropical arachnid fauna” (p.327). Actually, the genus Charinus Simon (by the way now in the Charinidae family) is the most widespread genus from all Amblypygi, with 54 tiny species, limited mostly by the climate. Thelyphonida (Uropygi) Another order missing in Australia, New Caledonia, and New Zealand but occurring in South Asia, Papuan area, and the Philippines. The finding of Mastigoproctus (American genus) with one specimen in Hong Kong is, for us, a case of Man’s transport. However, there is another much bigger puzzle with the description of the amblypygid Phrynus exsul Harvey, 2002, from a cave on Flores (Indonesia), the first and only representative of the American family Phrynidae and genus Phrynus. A relict like the Fiji iguanas? There was a discussion about the Hypoctonus species found in West Africa (no other Uropygi in or near Africa!) – whether a relict or a Man’s transport from East Asia, where the other Hypoctonus live (Cooke and Shadab 1973). Finally, from the species from West Africa was described a new genus Etienneus Heurtault, 1984 (another case of “fake disjunction”). The West African uropygids were declared autochthonous and relict by Huff and Prendini 2009 (but why they didn’t spread further in Africa?). Schizomida Despite the statement of Besch (1969) that “they are distinguished by the ease with which they are distributed into new areas by means of Man’s activities” (p. 730), very few of these tiny arachnids are actually being reported or supposed to be transported by Man. Some are known from European greenhouses (Zomus bagnallii Jackson, 1908 in Kew). Others have reached Hawaii or some other Pacific islands (Marianas, Fiji, Marshall Islands, Samoa, New Caledonia). We agree with Rowland (1972) that “the low agility and inability of Schizomida to disperse through suboptimal areas may account for their geographical distribution. They lack any secondary means of travel, such as phoresy, ballooning, resistant eggs, parasitism or other means employed by arachnids” (p.153). Scorpiones According to Vachon (1972) “Isometrus maculatus est, d’ailleurs, le seul scorpion cosmopolite et son extension due, vraisemblablement, à l’homme”. Similar observations made Berland (1932): “…cette espèce, frequentant les habitations humaines et penetrant dans les navires, s’est repandue dans toute la région chaude ou on la trouve parfois dans les îles les plus isolées, comme l’île de Paques, par exemple.” Araneae There are many articles describing the relatively active dispersal of spiders, called “ballooning” (Braendegaard 1937; Brignoli 1983; Szymkowiak et al. 2007 and others). Brignoli (1983) says, “I would like to note that “dispersal” is a 76 5 Factors Determining the Distribution of Arachnida rather poorly chosen term, not only because the distinction against “dispersion” is possible only in English, but also because it is used for only one kind of change of the range, more exactly only for the increase of it. But, as we all know, there are also decreases of the ranges, which also involve movements of the species.” Acari Wallwork (1979) insists that “It is very probably true that short-range overseas dispersal of Oribatids does occur, but the long-range dispersal over inhospitable oceans which would be required to produce the global patterns that we can observe today is unlikely.” For this author about 30% of the Oribatei Inferiores have a cosmopolitan distribution, compared with only 15% of the Superiores. Oribatids are an ancient group, and old genera have existed on the supercontinents and have been carried away by the continent’s products of the breakup of Gondwana and Laurasia. 5.4 5.4.1 Ecological Factors Climatic and Other Barriers and Bridges En somme, si nous laissons de coté l’Hologenèse qui est du domaine de la Génétique et la Dérive des continents qui ne s’appuie sur rien de solide, il ne nous reste plus que les relations intercontinentales des géographies anciennes. Nous considerons comme la cause essentielle des repartitions actuelles les variations du climat au cours des temps géologiques et singulièrement pendant le Quaternaire. R. Furon (1953a, b) The ecological factors (mostly temperature and humidity) of the territories are determining the present-day distribution. In the past the distribution of some arachnids has been different but always reflecting the thermos and hygropreferendum of the animals. Now we can outline several thermophilous groups with sharp limitation in their distribution: Solifugae, Amblypygi, Uropygi, Schizomida, Ricinulei, Holothyrida, and some lower-ranking groups like most Cyphophthalmi. Also the scorpions have limitations – from the 111 spp. in the USA, only 1 species reaches the extreme South of Canada. Another temperature limitation is observed in the high mountains. The presentday climatic zones of the Earth, so important for the distribution of animals, have been classified by Russian-German climatologist W. Köppen first in 1884 and then modified by himself, by Geiger, and other specialists (McKnight and Hess 2000). Some isotherms clearly are limits of the distribution, mostly of the thermophilous groups. The paleoclimates are also important to explain some distributions, mostly the relictual (Elouard 1984). Other factors are dryness/rainfall, length of the “moderate” seasons, etc. Some of the limitations are not so evident. Why there are no Solifugae in Madagascar – only 419 km far from the richest area in the world what concerns the Solifugae? This group has not reached Australia with its dry lands and deserts, maybe because of the vast areas of rain forest and other unsuitable habitats for Solifugae. Why many 5.4 Ecological Factors 77 ancient orders are lacking from the ancient land New Zealand? The way of dispersal and of reaching new territories by Arachnida are already known, but very few have profited by “rafts” to reach Australia from New Guinea and vice versa. There are many factors determining the ability of an arachnid to reach a new territory, to survive on it and even to start further speciation. Important factors are also time, glaciation, saturation of the new land with competitors, winds, etc. Some groups are much more plastic and ubiquitous than others. Such are spiders, many groups of mites, and to some extent Opiliones. Interesting case of development in complete isolation are some caves like Movile in Romania and Ayalon in Israel. In Movile have been found blind spiders (and other arthropods) of families usually not entering caves (Clubionidae, Theridiidae, Hahniidae). In Ayalon dead scorpions of a new and very strange family Akravidae have been found, along with a new genus of pseudoscorpions (Ayalonia). Time of speciation is also an important factor in the distribution of arachnids. Some are like “living fossils.” Others, like the arachnids in the Hawaiian lava tubes, have evolved in these tubes for a few thousand years in real troglobites. The physical barriers are of different types – sea, high mountains, large rivers, rain forest, deserts, and many others. They are not existing forever – the past distribution of many groups proves that the paleogeography is determining quite different paleozoogeography. Except of some mites, Arachnida do not live in sea water and have little resistance to it. All disjunctions, ubiquitous or pantropical species or genera remain from the former situation of the territories or may be explained by dispersal via the three W (wind, wings, waves) and more recently by the human transport (unlike some other invasive animals, the arachnids are transported involuntary). In the deserts prevail scorpions and solpugids, some other orders (spiders, mites) are barely represented, and some others (Ricinulei, Uropygi, Holothyrida) are completely missing. In the high mountains, on the contrary, spiders are prevailing; there are many mites, pseudoscorpions, and Opiliones; and the other orders are either missing or represented by low number of species (Beron 2008b). The grassland is without Amblypygi and other arachnids needing rock shelters and caves to hide, but spiders and mites are plentiful. The Holarctic broadleaf forest and even more the taiga are already outside the termopreferendum of the “southern” orders, and spiders, mites, pseudoscorpions, and Opiliones are masters of these biomes. Even more this is true for the orophytic areas. Humidity is important for most arachnids, and very few (scorpions, Solifugae, some mites) avoid areas with high humidity. 5.4.2 Real and “Fake” Disjunctions When you have a case of species of an American genus (and family!) found in a cave of Flores (Indonesia), as in the case of Phrynus exsul (Harvey, 2002), you may think of at least six options: 78 5 Factors Determining the Distribution of Arachnida 1. The species was wrongly identified. This applies usually for very old unchecked identifications, which need reassessment. 2. It goes here for an animal, transported somehow (usually by Man) to a faraway place. 3. To relict locality. 4. To lack of enough research in the area in between – the “gaps” might be in our knowledge. 5. To the fact that this animal is different from the “congeners” in the main area and should be given another name. 6. Mistake in the label. The case of the new Phrynus is really puzzling, as it has been described by a very qualified researcher, found repeatedly; the distance is so big that we cannot presume lack of collecting in between. Here are some cases of real and “fake” disjunctions. One of the classical groups to study disjunctions is the amphibians. Examples are the family Proteidae (Proteus, Necturus), the Cryptobranchidae (one extant species of Cryptobranchus in North America, two spp. of Andrias in China and Japan, and one fossil in Europe), and the New Zealand frogs (four spp. of Leiopelma) with the other two members of Leiopelmatidae of genus Ascaphus in North America. A good example we were teaching in the University were the newts of the genus Hydromantes, found in France, Italy, Sardinia, and California. Among the reptiles long-standing example was the turtle “Podocnemis” madagascariensis in Madagascar, considered member to a South American genus. However: 1. Some researchers created for the North American frogs a special family Ascaphidae. Still, they are very close with Leiopelma within the suborder Archaeobatrachia, and the problems of the origin of New Zealand frogs remain open. 2. The status of the only European member of the American family Plethodontidae (the lungless salamanders) is still a controversy. The three American species remain in the genus Hydromantes; the seven species from southern France, Italy, and Sardinia are given either generic or subgeneric status as Speleomantes and Atylodes. Still, a problem remains how they occurred in so distant regions. In 2005 in South Korea was found a new lungless salamander (Karsenia Min et al.), which indicates that the plethodontids were widespread once in the Holarctis. So, we see that such sensations occur with Amphibia, let alone the invertebrates! 3. The aberrant “Podocnemis” in Madagascar is no more a Podocnemis but is called Erymnochelys madagascariensis. Together with other Neotropical elements, it remains the only Madagascan member of the South American family Podocnemididae. 4. The enigmatic West African whip scorpions (an order which is otherwise not represented in or near Africa) have been longtime considered a zoogeographical puzzle. In Senegal, Guinea-Bisau, Guinea, and Gambia, uropygids were described as members of the Southeast Asian genus Hypoctonus (H. africanus Henschel, 1899). At first it was presumed that they have been transported by 5.4 Ecological Factors 79 Man from Southeast Asia (Cooke and Shadab 1973), but finally they received a separate generic status Etienneus (Heurtault, 1984) and were declared Gondwana relicts by Huff and Prendini (2009). Some mistakes are due to mislabeling or incorrect reading of old labels. One Mediterranean scorpion has been published by Roewer from “Samoa,” but the inquiry revealed the real locality – island Samos in Greece! In the materials published from the collection (Roewer), there are several such “zoogeographical puzzles” (Helversen and Martens 1972). 5.4.3 Old World-New World Disjunctions Amblypygi The genus Trichodamon Mello-Leitão, 1935 (Brazil, two spp.), is the only member of the Old World family Phrynichidae in the Americas (tropical and Southern Africa, Algeria, Morocco, Yemen, Saudi Arabia, Oman, Mauritius, Seychelles, Zanzibar, Sao Tome, Madagascar, Sri Lanka, Brazil, 32 spp.). The finding of an Old World species of Phrynus in Indonesia has been discussed above. Thelyphonida (Uropygi) The genus Thelyphonellus Pocock, 1894 (Brazil, Guyana, Venezuela) and Ravilops Viquez et Armas, 2005 (Dominican Republic), are the only American representatives of the subfamily Hypoctoninae (Burma, South China, Malaysia, West Africa, India, Sri Lanka). The Old World Mastigoproctus transoceanicus Lazell, 2000 from Hong Kong is, for this author, a case of transport by Man. Scorpiones One of the most enigmatic disjunctions is the case of the eyeless European scorpion Belisarius xambeui Simon, 1879 (France and Spain), placed in Chactidae (Brotheinae) by Soleglad and Fet (2003) and in the new family Troglotayosicidae (known from a cave in Ecuador) by Lourenço (1998), as base of a new subfamily Belisariinae. In both cases is presumed a South American connection – a striking case of New World – Europe disjunction. Such relations are presumed in the recently described from Israel cave scorpion Akrav and in other groups of animals (Isopoda terrestria). Another classical case of disjunction is the occurrence of genus Heteronebo Pocock on the islands Abd-El-Kuri (Yemen). Later (Francke 1978) some Heteronebo have been found in the Caribbean. However, it was proved that the species described by Pocock (Heteronebo granti and H. forbesii) do live on Abd-el-Kuri islands. Lourenço (1998) tries to explain this peculiar distribution with the “generalized tracks” of Croizat (1958). Similar explanation was proposed by Francke and Soleglad (1981) for the disjunction Caraboctonidae – Iuridea. Opiliones There are several interesting cases, analyzed (mostly in Japanese) by Suzuki (1972, 1977). Interesting case is the family Caddidae (in the traditional 80 5 Factors Determining the Distribution of Arachnida sense) – Caddella Hirst (South Africa), Hesperopilio Shear (Australia and Chile ), Caddo Banks (Japan, North America, fossil in Europe), Acropsopilio Silvestri (Japan, N. America, S. America, Australia, New Zealand), and Austropsopilio Forster (Australia) (= Tasmanopilio Hickman, Tasmania). According to Groh and Giribet (2015), Acropsopilionidae is no more a subfamily of Caddidae but a separate family in Caddoidea and, together with Caddidae, is transferred in Dyspnoi. Hesperopilio Shear is referred to Phalangioidea without indication of family. These changes the genera enumerated are still remarkable as connection between South America and Notogea. One genus (Jarmilana) of the Afrotropical family Pyramidopidae has been described from a cave in Belize. Americovibone Hunt et Cokendolpher is member of another interesting family of Eupnoi – Neopilionidae. One species of this genus lives in Chile, and recently another species has been described from New Zealand (Americovibone remota Taylor, 2016). The case with the genus Ceratomontia Roewer, 1915, with 22 spp., 18 known from South Africa and 4 from South America (Argentina, Uruguay, S. Brazil), was tested cladistically by Cruz Mendes and Kury (2008), who concluded that “The consensus hypothesis did not recover a monophyletic Ceratomontia, instead, the South African species constitute a clade with the Austromontia and Monomontia species. The Ceratomontia species from South America form a clade that is sistergroup of the clade formed by the South African genera of the ingroup. The result shows that Ceratomontia is involved in a “Gondwanan relationship” but also includes other genera, suggesting that it is not a monophyletic group.” Pseudoscorpiones Recently has been described from Spain and Portugal one remarkable new genus of cave Pseudoscorpions of the South American and Antillean family Bochicidae – Titanobochica Zaragoza et Reboleira, 2010 – and the genus Troglobisium Beier, 1939, from Spain was transferred to the same family. Both were declared relict (Reboleira et al. 2010). Araneae There are many cases like the genus Caloctenus Keyserling – one species in Ethiopia (Caloctenus abyssinicus Strand, 1917) and the other four in South America. Do they really belong to one genus? Similar is the case with genus Vulsor Simon – seven spp. in Madagascar and the Comoro Islands one (Vulsor occidentalis Mello-Leitão, 1922) in Brazil. Acari A good example is Trichosmaris papuana Beron, 2002 (Smarididae), described by us from the center of New Guinea. The other members of the genus are known from Guatemala and the USA. 5.4 Ecological Factors 5.4.4 81 West African-Indomalayan Disjunction One of the classic cases of disjunctions is that between the animals of the tropical rain forest of West and Central Africa and the wet forested areas of Southeast Asia, separated by steppes, deserts, and dry mountains in between. Examples are apes, peacocks, Tragulidae, Lorisidae, and many others. Are there such disjunctions among Arachnida? 5.4.5 Strange (Old?) Disjunctions This is found in many groups of Arachnida. Good example is the family Travuniidae, found (mainly in caves) in South Europe, North America, and (Yuria Susuki, inc. sedis) Japan. Another example from Opiliones – the family Sironidae (again: South Europe, North America and the relict genus Suzukielus Juberthie in Japan!). 5.4.6 Disjunctions in the Southern End of the World (New Zealand, Australia, South Africa, Patagonia) They are among the most interesting in the zoogeography and are explained by the former continuity of these lands, the trans-Antarctic road and similar. Good examples are found among the Opiliones and the spiders (outlined further). Such disjunctions have been detected also among other groups of animals and have provoked some authors (Lopatin 1980) to speak about a greater Notogea, comprising also Patagonia. Eskov and Golovatch (1986), analyzing the origin of the trans-Pacific disjunctions, conclude that the “southern Pacific semicircle” have preserved a considerable number of relicts extinct elsewhere…the fact is that patterns like South America, New Zealand, Australia, as well as the other types of circum-Pacific disjunctions, are nothing but particular cases of the reduction of a formerly single bipolar distribution. These authors insist also that “…the biogeographical significance of recent relicts is limited unless supported by paleontology.” 5.4.7 Boreomontane and Arctic-Alpine Disjunctions Holdhaus (1954) analyzed in details the distribution of some animals in the far north of the Holarctic and, after a wide gap, in the mountains like Alps (boreoalpine or arctic-alpine distribution) or some other mountains. This type of distribution is usually attributed to the consequences of the ice age in Eurasia and North America. Unlike Insecta, there are relatively few examples of boreomontane 82 5 Factors Determining the Distribution of Arachnida disjunctions in groups like Arachnida, Myriapoda, or Isopoda (Buresch and Arndt 1926; Beron 1969a, 2008b). Among Bulgarian animals as such are considered the mites Laelaps clethrionomydis and Hirstionyssus gudauricus and the spider Meioneta gulosa. A difference should be made between the notions “arctic-alpine” and “boreomontane” distribution. The animals with arctic-alpine distribution occur in the Arctic and Subarctic tundra and high in the mountains of Middle or South Europe. We speak about boreomontane distribution when animals inhabit the northern coniferous forests of Europe and North America and the relevant zones in the mountains of Europe, Caucasus, and Central Asia. The term “boreo-alpine” is less accurate, merging both above described and is better to avoid its use. Araneae Many studies are due to Thaler (1976), Thaler and Knoflach (2002), Relys (2000), and others for the spiders of the Alps. Relys (2000) registered 13 arctic-alpine species in the Eastern Alps, the lowest point being at 1618 spp. – Meioneta gulosa (L. Koch) – but most species have been detected over 2100 m. At this altitude more than 10% of all spider specimens in a community usually belong to arctic-alpine species. The 14 boreomontane species were located at altitudes above 1900 m on the timberline ecotone. According to Relys (2000), “despite high species richness neither boreomontane nor arctic-alpine spiders, nor both groups together, can make up such a significant part of subalpine spider communities as do alpine-endemic species (e.g., Pardosa oreophila Simon).” According to Bosmans and De Keer (1987), 14 of the spider species of the Pyrenees are arctic-alpine elements, 11 of them belonging to Linyphiidae, 1 to Araneidae, 1 to Lycosidae, and 1 to Gnaphosidae. 5.4.8 Transpacific Disjunctions Good examples are the Opiliones found in Japan and in North America. One (doubtful) explanation is the Beringian bridge. This bridge is essential for the “Northern Semicircle Eurasia – North America.” The “Southern Semicircle” (New Zealand, Australia, Antarctica, Cape area, Patagonia) also provoqued many explanations, Paleantarctic and others. Details with many examples (mostly insects) could be found in the article of Eskov and Golovatch (1986). The conclusion in this article is that the “southern Pacific semicircle” has preserved a considerable number of relicts extinct elsewhere. And also “…the fact is that patterns like South America – New Zealand – Australia, as well as the other types of circum-Pacific disjunctions, are nothing but particular cases of the reduction of a formerly single bipolar distribution.” 5.5 Endemics and Relicts in Arachnida 5.5 5.5.1 83 Endemics and Relicts in Arachnida Endemism Here the analyses are based on endemism to the level of family/subfamily and genus/subgenus. What is really endemism? Endemics are taxa which are known only from one area (geographical unit or, more narrow, cave, or mountain). In many cases they are considered endemic as long as they are not found elsewhere (especially the newly described taxa). For convenience, we use the name of countries to mark the endemic taxa. In use are the terms “Bulgarian endemic,” or “Balkan endemic.” The political borders, which are not natural and not permanent, do not make too much sense. If we take our country, Bulgaria, we see that most Bulgarian mountains are on the borders with the neighboring countries. An endemic species described from one part of the mountain almost surely will be found in the other parts. Exceptions are very narrow endemics like the cave animals (the Opiliones Paralola buresi from four caves in Stara Planina and nowhere else). So, the names of countries make sense in the case of islands (Malta, New Guinea – the whole island, Madagascar, Mauritius, New Caledonia, New Zealand – even there we find differences between the northern and the southern islands). Islands (even big) are good examples for endemism (Sardinia, Tasmania, Mallorca). We speak about endemics for Cuba and Jamaica but also about Antillean endemics. To call “endemic” taxon living in “Russia” or “the USA” does not make sense – is it Baltic shore or Sakhalin? Is it Florida or Oregon? Some countries like Brazil, DR Congo, or Canada are very big but uniformed in nature. So, sometimes we have to mark endemics only in a very broad sense (African, Indian, Australian). There is also a notion of “subendemic” to indicate the main areal and some smaller penetrations. We also have to distinguish between paleo- and neoendemics. Other sources: Brignoli (1979), Gromov (1998), Komposch (2011), Lazzeroni (1969), Lourenço (2007), Lourenço and Đình Sắc Pham (2010), Thaler (1976), Prendini et al. (2006). Some Endemic Genera and Families in Different Orders Palpigradi Allokoenenia Silvestri, 1913 – Guinea Triadokoenenia Condé, 1991 – Madagascar Solifugae Galeodumus Roewer, 1960 – Afghanistan Gluviema Caporiacco, 1937 – Somalia Paragaleodiscus Birula, 1941 – Yemen 84 5 Factors Determining the Distribution of Arachnida Roeweriscus Birula, 1937 – Iran Zombis Simon, 1882 – Israel Barrus Simon, 1880 – Egypt Prosolpuga Roewer, 1934 – Namibia Solpugista Roewer, 1934 – Namibia Solpugisticella Turk, 1960 – Kenya Solpuguna Roewer, 1933 – Namibia, South Africa, and Botswana Toreus Purcell, 1903 – South Africa Subfam. Melanoblossiinae – Southern Africa Daesiella Hewitt – Namibia Lawrencega Roewer, 1933 – Namibia, South Africa Melanoblossia Purcell, 1903 – South Africa Microblossia Roewer, 1941 – Namibia Unguiblossia Roewer, 1941 – Namibia Cordobulgida Mello-Leitão, 1938 – Argentina Gaucha Mello-Leitão, 1924 – Chile Gauchella Mello-Leitão, 1937 – Bolivia Metacleobis Roewer, 1934 – Brazil Mummucina Roewer, 1934 – Chile Mummuciona Roewer, 1934 – Venezuela Mummucipes Roewer, 1934 – Paraguay Sedna Muma, 1971 – Chile Uspallata Mello-Leitão, 1938 – Argentina, Chile Acanthogylippus Birula, 1934 – Israel Bdellophaga Wharton, 1981 – Namibia Lipophaga Purcell, 1903 – Namibia, Southern Africa Trichotoma Lawrence, 1968 – Namibia Ammotrechesta Roewer, 1934 – Central America Ammotrechinus Roewer, 1934 – Haiti, Jamaica Ammotrechona Roewer, 1934 – Cuba Antillotrecha Armas, 1994 – Dominican Republic, Leeward Islands, and Cuba Campostrecha Mello-Leitão, 1937 – Ecuador Dasycleobis Mello-Leitão, 1940 – Argentina Neocleobis Roewer, 1934 – Galapagos Scorpiones Belisarius Simon, 1879 – French and Spanish Pyrenees Alayotityus Armas, 1973 – Cuba Ananteris Borelli, 1910 – Trinidad y Tobago Tityopsis Armas, 1974 – Cuba Rhopalurus Thorell, 1876 – Cuba and Hispaniola Cazierius Francke, 1978 – Greater Antilles Oiclus Simon, 1880 – Lesser Antilles Didymocentrus Kraepelin, 1905 – Lesser Antilles and Central Cuba Pseudochactas Gromov, 1998 – Uzbekistan, Tajikistan 5.5 Endemics and Relicts in Arachnida Troglokhammouanus Lourenço, 2007 – Laos (cave) Vietbocap Lourenço et Đình Sắc Pham, 2010 – Vietnam, Laos (caves) Lanzatus Kovařik, 2001 – Iran Saharobuthus Lourenço et Duhem, 2009 – Western Sahara Somalibuthus Kovarik, 1998 – Somalia Somalicharmus Kovarík, 1998 – Somalia Amblypygi Musicodamon Fage, 1929 – South Morocco, Algeria Suborder Palaeoamblypygi – endemic suborder Fam. Paracharontidae – endemic family Paracharon Hansen, 1921 – Guinea-Bissau Thelyphonida (Uropygi) Glyptogluteus Rowland, 1973 – Philippines (Panay) Chajnus Speijer, 1936 – Sunda Islands Uroproctus Pocock, 1894 – India (Assam) Mimoscorpius Pocock, 1894 – Philippines Amauromastigon Mello-Leitão, 1931 – Brazil Ravilops Viquez et Armas, 2005 – Dominican Republic Etienneus Heurtault, 1984 – West Africa Schizomida Subfamily Megaschizominae Megaschizomus Lawrence, 1969 – Mozambique, Republic of South Africa Afrozomus Reddell et Cokendolpher, 1995 – Angola, DR Congo Adisomus Cokendolpher et Reddell, 2000 – Brazil Anepsiozomus Harvey, 2001 – Seychelles Artacarus Cook, 1898 – Liberia, Ivory Coast Attenuizomus Harvey, 2000 – North Australia Brignolizomus Harvey, 2000 – Australia Cubazomus Reddell et Cokendolpher, 1995 – Cuba Draculoides Harvey, 1992 – Australia Javazomus Reddell et Cokendolpher, 1995 – Java Julattenius Harvey, 1992 – Australia (Queensland) Kenyazomus Armas, 2014 – Kenya Luisarmasius Reddell et Cokendolpher, 1995 – Cuba, Pinos, Puerto Rico Mahezomus Harvey, 2001 – Seychelles Mayazomus Reddell et Cokendolpher, 1995 – Mexico (Tabasco, Chiapas) Neozomus Reddell et Cokendolpher, 1995 – India (Maharashtra) Notozomus Harvey, 1992 – Australia (Queensland) Oculozomus Reddell et Cokendolpher, 1995 – Sumatra Ovozomus similis (Hirst, 1913) – Seychelles Pacal Reddell et Cokendolpher, 1995 – Mexico Schizomus Cook, 1899 – Sri Lanka Secozomus latipes (Hansen, 1905) – Seychelles 85 86 5 Factors Determining the Distribution of Arachnida Sotanostenochrus Reddell et Cokendolpher, 1991 – Mexico Stewardpeckius Reddell et Cokendolpher, 1995 – Jamaica Tayos Reddell et Cokendolpher, 1995 – Ecuador Trithyreus Kraepelin, 1899 – Burma (= Myanmar) Pseudoscorpiones Neobalkanella Ćurčić, 2013 – Serbia Chiliochthonius Vitali-di-Castri, 1975 – Chile Congochthonius Beier, 1959 – DR Congo Drepanochthonius Beier, 1964 – Chile Francochthonius Vitali-di-Cstri, 1975 – Chile Malcolmochthonius Benedict, 1978 – USA (California, Oregon) Microchthonius Hadži, 1933 – Croatia Neochthonius Chamberlin, 1929 – USA (California) Selachochthonius Chamberlin, 1929 – South Africa, Lesotho Arcanobisium Zaragoza, 2010 – Spain Troglochthonius Beier, 1939 – Croatia, ? Sardinia Anisoditha J.C. Chamberlin et R.V. Chamberlin, 1945 – Paraguay Cryptoditha J.C. Chamberlin et R.V. Chamberlin, 1945 – Brazil Haploditha Caporiacco, 1951 – Venezuela Neoditha Feio, 1945 – Brazil Sororoditha J.C. Chamberlin et R.V. Chamberlin, 1945 – Brazil Verrucaditha J.C. Chamberlin, 1929 – USA Afroroncus Mahnert, 1981 – Kenya Ideoroncus Balzan, 1887 – Brazil, Paraguay Nannoroncus Beier, 1955 – Kenya, Uganda Shravana Chamberlin, 1930 – Thailand Indohya Beier, 1974 – India Pseudogarypus Ellingsen, 1909 – Canada, USA Neopseudogarypus Morris, 1948 – Tasmania Antillobisium Dumitresco et Orghidan, 1977 – Cuba Apohya Muchmore, 1973 – Mexico Bochica J.C. Chamberlin, 1930 – Grenada, Trinidad Leucohya J.C. Chamberlin, 1946 – USA (Texas), Mexico Paravachonium Beier, 1956 – Mexico Troglobochica Muchmore, 1984 – Jamaica Titanobochica Zaragoza et Reboleira, 2010 – Portugal Troglohya Beier, 1956 – Mexico Vachonium J.C. Chamberlin, 1947 – Mexico, Belize Arcanobisium Zaragoza, 2010 – Spain Titanobochica Zaragoza et Reboleira, 2010 – Portugal Beierobisium Vitali-di-Castri, 1970 – Falkland Islands Gymnobisium Beier, 1931 – South Africa, Lesotho Vachonobisium Vitali-di-Castri, 1963 – Chile Ammogarypus Beier, 1962 – Namibia, South Africa 5.5 Endemics and Relicts in Arachnida Eremogarypus Beier, 1955 – Namibia Meiogarypus Beier, 1955 – Namibia Neogarypus Vachon, 1937 – Zimbabwe Paragarypus Vachon, 1937 – Madagascar Paramenthus Beier, 1963 – Israel Hemisolinus Beier, 1977 – St. Helena Alabamocreagris Ćurčić, 1984 – USA (Alabama) Americocreagris Ćurčić, 1982 – USA (Oregon, Washington) Chinacreagris Ćurčić, 1983 – China Fissilicreagris Ćurčić, 1984 – USA (California) Globocreagris Ćurčić, 1984 – USA (California) Insulocreagris Ćurčić, 1987 – Croatia (Vis), Herzegovina Lissocreagris Ćurčić, 1981 – USA Minicreagris Ćurčić, 1981 – USA Nepalobisium Beier, 1974 – Nepal Novobisium Muchmore, 1967 – USA Occitanobisium Heurtault, 1977 – France Pedalocreagris Ćurčić, 1985 – Far East of Russia Paedobisium Beier, 1939 – Romania, Spain Pararoncus J.C. Chamberlin, 1938 – Japan Protoneobisium Ćurčić, 1988 – Croatia Roncobisium Vachon, 1967 – France Saetigerocreagris Ćurčić, 1984 – USA Simonobisium Heurtault, 1974 – France, Italy Tartarocreagris Ćurčić, 1984 – USA (Texas) Trisetobisium Ćurčić, 1982 – USA Tuberocreagris Ćurčić, 1978 – USA Afrowithius J.C. Chamberlin, 1931 – South Africa Aisthetowithius Beier, 1967 – Kenya, Tanzania Cryptowithius Beier, 1967 – Kenya Cyrtowithius Beier, 1955 – Namibia, South Africa Hyperwithius Beier, 1951 – Vietnam Paragoniochernes Beier, 1932 – South Africa Parallowithius Beier, 1955 – Namibia, South Africa Plesiowithius Vachon, 1954 – Mauretania Pogonowithius Beier, 1979 – DR Congo Protowithius Beier, 1955 – Juan Fernandez Islands Pseudatemnus Beier, 1947 – South Africa Scotowithius Beier, 1977 – St. Helena Sphallowithius Beier, 1977 – St. Helena Thaumatowithius Beier, 1940 – Reunion, Mauritius Trichotowithius Beier, 1944 – Ethiopia, Kenya Tropidowithius Beier, 1955 – Peru Amaurochelifer Beier, 1951 – Vietnam 87 88 5 Factors Determining the Distribution of Arachnida Ancistrochelifer Beier, 1951 – Laos, Cambodia, Vietnam Aperittochelifer Beier, 1955 – South Africa Aporochelifer Beier, 1953 – Indonesia (Flores) Aspurochelifer Benedict et Malcolm, 1979 – USA Canarichelifer Beier, 1965 – Canary Islands, Ilhas Selvagens Chamberlinarius Heurtault, 1983 – Ivory Coast Cubachelifer Hoff, 1946 – Cuba, Dominican Republic Ectoceras Stecker, 1875 – India Florichelifer Hoff, 1964 – USA (Florida) Haplochelifer J.C. Chamberlin, 1932 – USA Hygrochelifer Murthy et Ananthakrishnan, 1977 – India Idiochelifer J.C. Chamberlin, 1932 – USA Juxtachelifer Hoff, 1956 – USA (New Mexico) Kashimachelifer Morikawa, 1957 – Japan Levichelifer Hoff, 1946 – Mexico, USA Litochelifer Beier, 1948 – South Africa Lophodactylus J.C. Chamberlin, 1932 – Brazil Macrochelifer Vachon, 1940 – China Mexichelifer Muchmore, 1973 – Mexico Nannocheliferoides Beier, 1974 – India Pachychelifer Beier, 1962 – Georgia Paisochelifer Hoff, 1946 – USA Papuchelifer Beier, 1965 – Indonesia (Papua), Papua New Guinea Phorochelifer Hoff, 1956 – USA Pilochelifer Beier, 1935 – Mauritius, Reunion Pseudorhacochelifer Beier, 1976 – Madeira, Canary Islands Pugnochelifer Hoff, 1964 – USA Rhopalochelifer Beier, 1964 – South Africa Sinochelifer Beier, 1967 – China Stenochelifer Beier, 1967 – India Telechelifer J.C. Chamberlin, 1949 – ? Sri Lanka Tetrachelifer Beier, 1967 – Vietnam Xenochelifer J.C. Chamberlin, 1949 – USA (California) Scotowithius Beier, 1977 – St. Helena Sphallowithius Beier, 1977 – St. Helena Athleticatemnus Beier, 1979 – DR Congo Brazilatemnus Muchmore, 1975 – Brazil Caecatemnus Mahnert, 1985 – Brazil Mesatemnus Beier et Turk, 1952 – Cyprus Synatemnus Beier, 1944 – Tanzania Acanthicochernes Beier, 1964 – Papua New Guinea, Solomon Islands Acuminochernes Hoff, 1949 – USA Adelphochernes Beier, 1937 – Philippines (Mindanao, Mindoro) Ancalochernes Beier, 1932 – Mexico Atherochernes Beier, 1954 – Venezuela 5.5 Endemics and Relicts in Arachnida Cacoxylus Beier, 1965 – Solomon Islands (Guadalcanal), Papua New Guinea Calidiochernes Beier, 1954 – Venezuela Calymmachernes Beier, 1954 – Australia (W. Australia) Caribochernes Beier, 1976 – Dominican Republic Chiridiochernes Muchmore, 1972 – Indonesia (Sulawesi) Chrysochernes Hoff, 1956 – USA (New Mexico) Cocinachernes Hentschel et Muchmore, 1989 – Mexico Conicochernes Beier, 1948 – Australia Coprochernes Beier, 1976 – Costa Rica Corosoma Karsch, 1879 – Brazil Cyclochernes Beier, 1970 – Solomon Islands (Guadalcanal) Dasychernes J.C. Chamberlin, 1929 – Colombia, Panama Diplothrixochernes Beier, 1962 – Argentina Epichernes Muchmore, 1982 – Mexico Eumecochernes Beier, 1932 – Hawaii Gelachernes Beier, 1940 – Solomon Islands, Papua New Guinea Gobichernes Krumpál et Kiefer, 1982 – Mongolia Hexachernes Beier, 1953 – Guatemala Marachernes Harvey, 1992 – Australia Petterchernes Heurtault, 1986 – Brazil Selachochernes Mahnert, 2011 – Juan Fernandez Islands Illinichernes Hoff, 1949 – USA Maorichthonius Chamberlin, 1925 – New Zealand Sathrochthoniella Beier, 1967 – New Zealand Tyrannochthoniella Beier, 1966 – New Zealand Heterochernes Beier, 1966 – New Zealand Hemisolinus Beier, 1977 – Saint Helena Antillobisium Dumitresco et Orghidan, 1977 – Cuba Caribochernes Beier, 1976 – Dominican Republic Cubachelifer Hoff, 1946 – Cuba, Dominican Republic Mexichthonius Muchmore, 1975 – Mexico (Campeche) Apohya Muchmore, 1973 – Mexico Vachonium J.C. Chamberlin, 1947 – Mexico, Belize Mexichelifer Muchmore, 1973 – Mexico Ancalochernes Beier, 1932 – Mexico Epichernes Muchmore, 1982 – Mexico Papuchelifer Beier, 1965 – Indonesia (Papua), Papua New Guinea Selachochthonius Chamberlin, 1929 – South Africa, Lesotho Gymnobisium Beier, 1931 – South Africa, Lesotho Afrowithius J.C. Chamberlin, 1931 – South Africa Ammogarypus Beier, 1962 – Namibia, South Africa Eremogarypus Beier, 1955 – Namibia Meiogarypus Beier, 1955 – Namibia Cyrtowithius Beier, 1955 – Namibia, South Africa Paragoniochernes Beier, 1932 – South Africa 89 90 5 Factors Determining the Distribution of Arachnida Parallowithius Beier, 1955 – Namibia, South Africa Pseudatemnus Beier, 1947 – South Africa Aperittochelifer Beier, 1955 – South Africa Litochelifer Beier, 1948 – South Africa Rhopalochelifer Beier, 1964 – South Africa Cryptoditha J.C. Chamberlin et R.V. Chamberlin, 1945 – Brazil Haploditha Caporiacco, 1951 – Venezuela Neoditha Feio, 1945 – Brazil Sororoditha J.C. Chamberlin et R.V. Chamberlin, 1945 – Brazil Ideoroncus Balzan, 1887 – Brazil, Paraguay Tropidowithius Beier, 1955 – Peru Lophodactylus J.C. Chamberlin, 1932 – Brazil Brazilatemnus Muchmore, 1975 – Brazil Caecatemnus Mahnert, 1985 – Brazil Atherochernes Beier, 1954 – Venezuela Calidiochernes Beier, 1954 – Venezuela Atherochernes Beier, 1954 – Venezuela Corosoma Karsch, 1879 – Brazil Dasychernes J.C. Chamberlin, 1929 – Colombia, Panama Aporochelifer Beier, 1953 – Indonesia (Flores) Chiridiochernes Muchmore, 1972 – Indonesia (Sulawesi) Ectoceras Stecker, 1875 – India Hygrochelifer Murthy et Ananthakrishnan, 1977 – India Paramenthus Beier, 1963 – Israel Congochthonius Beier, 1959 – DR Congo Afroroncus Mahnert, 1981 – Kenya Nannoroncus Beier, 1955 – Kenya, Uganda Aisthetowithius Beier, 1967 – Kenya, Tanzania Cryptowithius Beier, 1967 – Kenya Pogonowithius Beier, 1979 – DR Congo Trichotowithius Beier, 1944 – Ethiopia, Kenya Chamberlinarius Heurtault, 1983 – Ivory Coast Synatemnus Beier, 1944 – Tanzania Congochthonius Beier, 1959 – DR Congo Afroroncus Mahnert, 1981 – Kenya Nannoroncus Beier, 1955 – Kenya, Uganda Aisthetowithius Beier, 1967 – Kenya, Tanzania Cryptowithius Beier, 1967 – Kenya Pogonowithius Beier, 1979 – DR Congo Trichotowithius Beier, 1944 – Ethiopia, Kenya Chamberlinarius Heurtault, 1983 – Ivory Coast Athleticatemnus Beier, 1979 – DR Congo Synatemnus Beier, 1944 – Tanzania Chiliochthonius Vitali-di-Castri, 1975 – Chile Drepanochthonius Beier, 1964 – Chile 5.5 Endemics and Relicts in Arachnida Francochthonius Vitali-di-Castri, 1975 – Chile Vachonobisium Vitali-di-Castri, 1963 – Chile Diplothrixochernes Beier, 1962 – Argentina Australochelifer Beier, 1975 – Australia Conicochernes Beier, 1948 – Australia Pararoncus J.C. Chamberlin, 1938 – Japan Kashimachelifer Morikawa, 1957 – Japan Opiliones Fam. Troglosironidae – New Caledonia Troglosiro Juberthie, 1979 – New Caledonia Pettalus Thorell, 1878 – Sri Lanka Rakaia Hirst, 1925 – New Zealand Aoraki Boyer et Giribet, 2007 – New Zealand Neopurcellia Forster, 1948 – New Zealand – South Island Purcellia Hansen et Soerensen, 1904 – Western South Africa Parapurcellia Rosas Costa, 1950 – Eastern South Africa Austropurcellia Shear, 1980 – Australia (Queensland) Manangotria Shear et Gruber, 1996 – Madagascar Karripurcellia Giribet, 2003 – Western Australia Speleosiro Lawrence, 1931 – South Africa Chileogovea Roewer, 1961 – Chile Fam. Ogoveidae – equatorial Africa Ogovea Hansen et Soerensen, 1914 – Bioko, Cameroon, Congo Huitaka Shear, 1979 – Colombia Shearogovea Giribet, 2011 – Mexico Meghalaya Giribet, Sharma et Bastawade, 2007 Fangensis Rambla, 1994 – Thailand Metasiro Juberthie, 1960 – Florida, Georgia, S. Carolina Odontosiro Juberthie, 1961 – Portugal Iberosiro de Bivort, 2004 – Portugal Paramiopsalis Juberthie, 1962 – Portugal, Spain Suzukielus Juberthie, 1970 – Japan Marwe Shear, 1985 – Kenya ¿Tranteeva Kratochvil, 1958 – Bulgaria ? Ankaratra Shear et Gruber, 1996 – Madagascar (one sp., inc. sedis) Caddella Hirst, 1925 – South Africa Hesperopilio Shear, 1996 – Western Australia Austropsopilio Forster, 1955 – Australia (New South Wales) Tasmanopilio Hickman, 1957 – Tasmania Yuria Suzuki, 1964 – Japan Buemarinoa Roewer, 1956 – Sardinia Fam. Synthetonychiidae – New Zealand (14 spp.) Synthetonychia Forster, 1954 – New Zealand (14 spp.) Fam. Escadabiidae – Brazil 91 92 5 Factors Determining the Distribution of Arachnida Biantoncopus Martens et Schwendinger, 1998 – Philippines (Leyte) Caenoncopus Martens et Schwendinger, 1998 – Sumatra Palaeoncopus Martens et Schwendinger, 1998 – Sumatra Konfiniotis Roewer, 1940 – Crete Acuclavella Shear, 1986 – Idaho, etc. Ceratolasma Goodnight et Goodnight, 1942 – Oregon Crosbycus Roewer, 1914 – British Columbia Ausobskya Martens, 1972 – Greece, southern Bulgaria Banksula Roewer, 1949 – California (10 spp.) Bishopella Roewer, 1927 – southeastern USA Calicina Ubick et Briggs, 1989 – California Crosbyella Roewer, 1927 – southeastern USA Haasus Roewer, 1949 – Israel Lola Kratochvil, 1937 – Dalmatia Maiorerus Rambla, 1993 – Canary Islands Microcina Briggs et Ubick, 1989 – California Paralola Kratochvil, 1958 – Bulgaria Proscotolemon Roewer, 1916 – Japan Sitalcina Banks, 1911 – California Tolus Goodnight et Goodnight, 1942 – Tennessee Trojanella Karaman, 2005 – Serbia Undulus Goodnight et Goodnight, 1942 – Alabama Wespus Goodnight et Goodnight, 1942 – Arkansas Aburista Roewer, 1935 – Ghana Sokodea Roewer, 1935 – Togo Typhloburista Lawrence, 1947 – Ivory Coast Afroassamia Caporiacco, 1940 – Ethiopia Assamiella marginata (Roewer, 1912) – Burma Gudalura Roewer, 1927 – India Micrassamula Martens, 1977 – Nepal Neassamia Roewer, 1935 – Thailand Nepalsia Martens, 1977 – Nepal Nepalsioides Martens, 1977 – Nepal Pechota Roewer, 1935 – Malacca Popassamia Roewer, 1940 – Burma Puria Roewer, 1923 – India (Maharashtra) Tavoybia quadrispina Roewer, 1935 – Malacca Umtaliella Lawrence, 1934 – Zimbabwe Irumua Roewer, 1961 – DR Congo Machadoessa Lawrence, 1951 – Angola Mutadia Kauri, 1985 – DR Congo Numipedia Kauri, 1985 – DR Congo Typhlobunus Roewer, 1915 – Kenya 5.5 Endemics and Relicts in Arachnida Araneae Atrophothele Pocock, 1903 – Socotra Aurecocrypta Raven, 1994 – Australia Barycheloides Raven, 1994 – New Caledonia Barychelus Simon, 1889 – New Caledonia Cyrtogrammomma Pocock, 1895 – Guyana Encyocrypta Simon, 1889 – New Caledonia Fijocrypta Raven, 1994 – Fiji Idiommata Ausserer, 1871 – Australia Moruga Raven, 1994 – Australia Natgeogia Raven, 1994 – New Caledonia Nihoa Raven et Churchill, 1992 – Oceania Orstom Raven, 1994 – New Caledonia Ozicrypta Raven, 1994 – Australia Plagiobothrus Karsch, 1891 – Sri Lanka Questocrypta Raven, 1994 – New Caledonia Seqocrypta Raven, 1994 – Australia Synothele Simon, 1908 – Australia Trittame L. Koch, 1874 – Australia (Queensland) Tungari Raven, 1994 – Australia (Queensland) Zophorame Raven, 1990 – Australia (Queensland) Zophoryctes Simon, 1902 – Madagascar Neodiplothele Mello-Leitão, 1917 – Brazil Paracenobiopelma Feio, 1952 – Brazil Ammonius Thorell, 1899 – Cameroon Eubrachycercus Pocock, 1897 – Somalia Reichlingia Rudloff, 2001 – Belize Troglothele Fage, 1929 – Cuba Fam. Halidae Jocqué, 1994 – Madagascar Hala Jocqué, 1994 – Madagascar Tolma Jocqué, 1994 – Madagascar Teloleptoneta Ribera, 1988 – Portugal Masirana Kishida, 1942 – Japan Falcileptoneta Komatsu, 1970 – Japan Rhyssoleptoneta Tong et Li, 2007 – China Opilioacarida Adenacarus Van der Hammen, 1966 – Yemen Amazonacarus Vasquez, Araújo et Feres, 2014 – Brazil Brasilacarus Vasquez, Araújo et Feres, 2014 – Brazil Indiacarus Das et Bastawade, 2007 – India Phalangiacarus Coineau et Van der Hammen, 1979 – Gabon Siamacarus Leclerc, 1989 – Thailand Vanderhammenacarus Leclerc, 1989 – Thailand 93 94 5 Factors Determining the Distribution of Arachnida Parasitiformes Holothyrida Australothyrus van der Hammen, 1972 – Australia Dicrognathus Gerlach, Lehtinen et Mandl, 2010 – Seychelles Hammenius Lehtinen, 1981 – New Guinea Haplothyrus Lehtinen, 1995 – New Caledonia Holothyrus Gervais, 1842 – Mauritius Indothyrus Lehtinen, 1995 – Sri Lanka Leiothyrus van der Hammen, 1983 – New Guinea Michaelothyrus Gerlach, Lehtinen et Mandl, 2010 – Seychelles Sternothyrus Lehtinen, 1995 – Seychelles Thonius Lehtinen, 1981 – New Guinea Caribothyrus Kontschán et Mahunka, 2004 – Dominican Republic Diplothyrus Lehtinen, 1999 – Brazil, French Guyana Ixodida Fam. Nuttalliellidae – Southern Africa Nuttalliella Bedford, 1931 – Southern Africa – from Tanzania to Namibia 5.5.2 Relicts and Relictness Are There Relicts at All? The Issue of Relicts Types of Relicts The notion of “relict” is widely used by biogeographers, despite the warning of some of them to their colleagues to not allow themselves to be carried away too much (Elenevskiy and Radigina 2002). This notion was used widely by biospeleologists, influenced by the book of Jeannel “Les fossiles vivants des cavernes” (1943). Some researchers strongly oppose this very notion, as does the prominent Italian Arachnologist Brignoli (1979): “Le terme de “relicte” (ou même de “fossile vivant”) si souvent employé pour les troglobies, n’a pour moi aucun sense.” Checking in the dictionaries, we can find several interpretations of “relict.” Merriam-Webster Dictionary: “A surviving species of an otherwise extinct group of organisms; also: a remnant of a formerly widespread species that persists in an isolated area.” The Free Dictionary: “An organism or species of an earlier time surviving in an environment that has undergone considerable change.” Botanists have three concepts to classify relicts: geographic, taxonomic, and lineage relicts. Birstein (1947), confirmed by Vandel (1964): “Les rélicts comme des types animaux (ou végétaux) dont l’évolution est arrêtée ou du moins fort ralentie, et qui ont conservé le facies de leurs lointains ancêtres.” 5.5 Endemics and Relicts in Arachnida 95 In the recent paper of Grandcolas et al. (2014), relicts (geographical or philogenetic) are identified as “a species or a group of species remaining from a large group that is mainly extinct.” We shall not discuss here this problem on the scale of all animals and plants. For us the statement of Brignoli is far too strong and cannot be supported. Of course, relicts and even “living fossils” do exist – classical examples are the two fish species of Latimeria, the two species of the reptile tuatara (Sphenodon), the okapi, and many, many others. Worth mentioning are the marine relicts in the cave waters (the Sphaeromides and the other Cirolanidae). High in the mountains of Papua New Guinea, far from any sea, we found in the caves polychaete worms (Namanereis), Isopoda Anthuridea (Cyathura), and snails of marine origin at almost 3000 m (Selmistomia). There is no doubt that they are relicts. What, if not relict, is the recently discovered in Laos rodent Laonastes, the only living member of a family Diatomyidae, extinct 11 million years ago? Or the only member of the family Microbiotheriidae, also known (at latest) from the Miocene – the small marsupial from Chile Dromiciops gliroides (“monito del monte”)? Here we shall limit our discussion using examples among Arachnida. Amblypygi The species Paracharon caecus Hansen, 1921, in the family Paracharontidae and even the suborder Palaeoamblypygi from Guinea-Bissau is declared “living fossil” by various authors. Scorpiones Among scorpions one of the most striking examples or relictness is the recently described family Pseudochactidae Gromov, 1998. It contains for the moment three genera, all considered relicts: Pseudochactas Gromov in Central Asia, Troglokhammouanus Lourenço (cave in Laos), and Vietbocap Lourenço et Đình Sắc Pham (caves in Vietnam and Laos) (Lourenço and Pham 2012, 2014). It is a very primitive family (Prendini et al. 2006). A relict scorpion of Buthidae family (new genus) was described from the Great Rift Valley in Kenya (Riftobuthus inexpectatus Lourenço, Duhem et Cloudsley -Thompson, 2010). Another case of relict scorpion is Euscorpius oglasae Caporiacco, 1950, on the island of Montecristo. This is a case of paleoendemism, according to Vignoli et al. (2007). Pseudoscorpiones Among the pseudoscorpions, especially the troglobites, there are many which could qualify for the category of relics. Many detailed studies on the pseudoscorpions, termed as relicts, are due to B. Ćurčić and his team (Ćurčić, 1983, 1985, Ćurčić and Dimitriević 1984 and others). Schawaller (1987) recorded a second relict locality of the northern species Syarinus strandi (Ellingsen) in Central Europe (South Germany), after the finding in Austria of this species, known from Norway (Mahnert 1976). There are also other examples of relicts in European Syarinidae. One of the most striking is Hadoblothrus aegaeus Beron, 1985, found by us in caves on Santorin and Iraklia islands. 96 5 Factors Determining the Distribution of Arachnida For us there is no doubt that the family Syarinidae contains some striking cases of relictness. According to Beier (1963), “Syarinidae: Wohl bei keiner anderen Familie ist der Reliktcharakter so ausgeprägt.” Gardini (1982): “Hadoblothrus, monotipico, è di incerta affinità e representa uno dei più singolari relitti della regione pugliese.” It is curious that not a single representative of the family Syarinidae has been found so far in the numerous (and partly well investigated) caves on the mainland of Balkan Peninsula. Hadoblothrus aegaeus has been found in the islands from where not a single cave Neobisiidae has been recorded. The closest Syarinidae to the Balkan Peninsula are Pseudoblothrus Beier (one sp. in France, two species in Crimea, two in Italy, and two in Switzerland, all living in caves, except for P. regalini Inzeghi, 1983), Hadoblothrus gigas from South Italy (two caves), and Syarinus strandi (Austria, Germany, Finland, and Norway; all other five Syarinus species living in North America, outside caves). Another striking example of relict distribution are two genera of the South American family Bochicidae in caves of Spain (Troglobisium Beier, 1939) and south Portugal (Titanobochica Zaragoza et Reboleira, 2010). Opiliones Roewer (1935) declared the few Laniatores in Europe being all relicts. Martens (1972), describing the first non-troglobite Laniatores on Balkan Peninsula (Ausobskya athos from Greece), came to the conclusion that “The European Laniatores should no longer be regarded as tertiary relics, for they are nowadays widely distributed inhabitants of soil litter in areas not covered by ice during the glacious periods.” At least, for genera like Lola in Dalmatia, Paralola in Bulgaria, and Trojanella in Serbia (also in Stara Planina, both very restricted monotypic troglobites), for us, there is no doubt that they represent fauna and environment of the past. Concerning the relicts in the other countries, we must mention the North American cave harvestmen (Briggs 1971, 1974) and the whole family Troglosironidae in New Caledonia (Giribet et al. 2012). In the fam. Pyramidopidae, Meiorerus randoi Rambla from a cave in Fuerteventura and Conomma orientale (Roewer) from Moshi (Tanzania) are considered relictual (Sharma et al. 2011). Thelyphonida (Uropygi) Huff and Prendini (2009) consider the only African genus Etienneus Heurtault in West Africa as Gondwanan relict. Araneae Interesting is the case of the relict family Liphistiidae, sole member of the suborder Mesothelae. This family is found in Japan, China, and Southeast Asia. According to Xu X et al. (2015), these spiders are “living fossils,” and the suborder Mesothelae is an ancient clade sister of all modern spiders. Again according these authors, Liphistiidae genera originate in Asia in the Paleogene (4–24 Ma). This timing is relatively recent, taking into account the old age of the spider divergence (297.6 Ma) between the Mesothelae and the Opisthothelae (Mygalomorphae and Araneomorphae). 5.6 The Northern Limits of Arachnida 97 Acari Oribatids are a very ancient group of arachnids and genera present in the world fauna today have existed unchanged for long periods of geological time. It is postulated that many of these genera predate the breakup of the supercontinent Pangaea and the drifting apart of continental landmasses (Wallwork 1979). The distribution of some genera on the southern landmasses reflects a former continuous pattern which extended across Gondwanaland. Among the cosmopolitan genera prevail the “primitive” ones (Inferiores), compared with the derived ones (Superiores). Wallwork (1979) emphasizes on the zoogeographic principle that random dispersal by wind, water, birds, or Man results in dysharmonic faunas. In contrast, harmonic faunas are those which present the same proportion of specific or generic elements in two or more geographically separated areas. Such harmony implies a previous generic continuity between the faunas in question and, by inference, previous land connections between the now-separated areas in which they live. Different groups of Oribatida show harmonic patterns of distribution either in the Southern Hemisphere (Crotonia or Podacaridae) or in the Northern Hemisphere (Ameronothridae and others). 5.6 The Northern Limits of Arachnida If we consider the distributions beyond the northern polar circle, we find their representatives of only four arachnid units: Acariformes (superorder) (Acaridida, Prostigmata, Oribatida), Parasitiformes (superorder) (Mesostigmata, Ixodida), Opiliones, and Araneae (Cokendolpher 1981). Opiliones From the catalogue of Cokendolpher and Lee (1993) of the Opiliones of North America and Greenland, we find the following limited number of species (parts of these territories are situated south of the NPC): Alaska. 13 spp. of the genera Dendrolasma (Nemastomatidae), Leiobunum, Leptobunus, Leuronychus, Liopilio, Mitopus, Nelima, Parodiellus, Phalangium (Phalangiidae), Sabacon (Sabaconidae) Greenland. Mitopus morio (Phalangiidae) Northwest territories (Canada). Mitopus morio (Phalangiidae) Yukon territory (Canada). Liopilio (Phalangiidae) Opiliones from Eurasia north of the NPC: According to Kauri (1980) and Stol (1993, 2003, 2007), a total of five Opilionids inhabit the Faroe Islands and four inhabit Iceland. North of the NPC are known: Finland. According to Lehtinen (1964), in Finnish Lappland (mostly Nord of 66°N) are known six species of Opilions of the genera Nemastoma (Nemastomatidae) and Mitopus, Oligolophus, Phalangium, Euphalangium Platybunus (Phalangiidae). Similar Opiliones are recorded from Sweden, Norway, and Russia. In conclusion, north of the NPC are known Opiliones from three families (Phalangiidae, Sabaconidae, Nemastomatidae). The widest distributed species is Mitopus morio (Phalangiidae). 98 5 Factors Determining the Distribution of Arachnida What concerns the Pseudoscorpions, Beier (1963) says: “None, except a few introduced synanthropic species, are found at higher latitudes than ca. 65°N.” Scorpiones According to Fet (2010), “The northern limit of natural (nonanthropogenic) scorpion distribution in Europe is in Saratov Province, Russia, at 50°40′54″ N, for Mesobuthus eupeus (Buthidae).” In Western Europe, the northernmost record of scorpions seems to be in Austria (Euscorpius germanus, North Tyrol, Walchsee, 47°39′N; Komposch et al. 2001). Davygora and Rusakov (2001) recorded Mesobuthus eupeus from Orenburg Province, Russia. The finding in Aituar Steppe (Orenburg Natural Reserve) at 51°13′N, 57°75′E is “the northernmost limit of natural scorpion distribution in Asia and the Palearctic” (Fet 2010). According to this author, “only some records of Paruroctonus boreus (Girard 1854) in southern Alberta, Canada, lie slightly further north.” Solifugae In Europe – Spain (Gluvia dorsalis), Sicily (Biton ehrenbergi, B. velox), Macedonia (Galeodes graecus, G. elegans), Greece, Bulgaria (G. graecus), Ukraine (Galeodes araneoides). In Asia – Mongolia (Karschia, Galeodes, Eusimonia – five spp.). In North America – two species reach the southern provinces of Canada: Eremobates docolora Brookhart et Muma and Hemerotrecha denticulata Muma. Palpigradi In Europe the limit is crossing Slovakia, Hungary, Austria, and France. Amblypygi In Asia – on the Greek islands of Rhodes and Kos and in southern Turkey, as well as two spp. of Charinus Simon in Israel and one sp. in Pakistan. In North America – the genera Phrynus, Paraphrynus, and Acanthophrynus (Phrynidae) are known from the states of Arizona, California, Florida, and Texas. Thelyphonida (Uropygi) In Asia – Typopeltis Pocock (? the Far East of Russia). In North America – Mastigoproctus Pocock (USA – Arizona, Florida, New Mexico, Oklahoma, Texas) Schizomida In Asia – India, Taiwan, and Ryukyus. In North America – Hubbardiidae (Hubbardia Cook - Arizona, California, Stenochrus Chamberlin, Agasthoschizomus – USA: Texas). After Reddell and Cokendolpher (1995), two species (Hubbardia idria and H. briggsi) reach in California 36°30′N. Araneae The huge order of Araneae contains many cold-resisting species (up to 6700 m in Himalaya, 203 spp. above 3500 m. and many inhabitants of Arctic and Subarctic areas (Beron 2008b, 2016a, b, c; Larsen and Scharff 2003 (Greenland), Marusik et al. 2005 (Yakutia), Marusik and Koponen 2002; Koponen 1993, 1995, 1996). Even so, the northern end of the world is much less populated by spiders than the more southern areas and especially in the tropics. The next table is modified from Koponen (1996). 5.6 Area Mediterranean Switzerland Russian plain Belgium Great Britain Norway Finland Alaska Iceland Greenland a 99 The Northern Limits of Arachnida Species 4000 900 940 650 620 535 620 339 90 70 Families 50 39 36 35 35 29 27 10 10 Area (km2) – 41,200 2,000,000 30,500 230,700 323,900 338,100 1,723,300 89,700a 341,700a Ice – free area The analysis of Koponen (1996) shows that the family Linyphiidae prevails with 45% of all spiders in Finland, 43% in the Sakhalin-Kuril Islands, 48% in Yakutia, 60% in northeastern Siberia, and 62% in Central Siberia (on close latitude). In the most northern areas, Linyphiidae is characteristic (Koponen 1995) – from 63% of all spiders in Greenland and 75% among the nine families in Faroe, remains the only family with five spp. on Yan Mayen. Koponen (1984) announce that 216 species of spiders are known from Inari Lapland (68°20′–70°05′N in Finland). The harsh climate and lack of forests on the Arctic and Subarctic islands are the main reason for the low number of spider species on them (5 on Jan Mayen, 14 on Svalbard, 67 on the Faroe, 70 from Greenland, 90 from Iceland). The estimate of Koponen (1995) is that on the ice-free area of these islands is 450,000 km2, and they are populated by ca.150 spp. of spiders. The similarity between the spiders of Iceland and Scandinavia is 93%, but the fauna of Greenland is clearly Nearctic. Braendegård (1960) identified a collection of 103 spiders (nine species, eight native), brought from Peary Land (ca. 82°N). These species were Dictyna borealis Cambridge. Acari The general paper on Arctic mites (Trägårdh 1904) appeared very early, since this time some more publications contributed to the better understanding of the peculiar fauna surviving north of the polar circle. According to Makarova (2015), “The number of mite species known in Greenland increased from 158 to 290 (59 species were identified only to the generic level). They belong to the orders Parasitiformes (Mesostigmata 63 species, Ixodida 1) and Acariformes (Prostigmata 94, Endeostigmata 5, Oribatida 110, Astigmata 17). The free-living mite fauna in general is characterized by a complete taxonomic structure and a full range of ecological diversity. The Greenland fauna of oribatid mites mainly consists of widespread polyzonal species (70%), while specialized cryobiont (arctic and arctomontane) species are few (4%); the Nearctic species prevail over the European invaders with a ratio of 1.5:1. 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Bulletin de la Société neuchâteloise des Sciences naturelles 116(1):141–152 Koponen S (1995) Spider fauna (Araneae) of the arctic-subarctic Atlantic islands. Boletim Museu Munilipal Funchal 4:373–377 Koponen S (1996) Diversity and similarity of northern spider faunas. Acta Zool Fenn 201:3–5 Larsen S, Scharff N (2003) The spiders of Greenland – checklist (Arachnida: Araneae). Ent Meddr 71(1):53–61 [Lazukov, GI] (1973) Time and place of the formation of the cold resisting flora and fauna. Vestnik Moskovskogo Universiteta, Geographiya 3:31–47 Lehtinen P (1964) The Phalangids and Pseudoscorpionids of finnish lapland. Rep Kevo Subarctic Res Station 1:279–287 Chapter 6 Review of the Classification of the Orders of Arachnida Abstract Review of all orders with maps of their geographical distribution. 6.1 Palpigradi Order Palpigradi – 102 spp. Fam. Eukoeneniidae – Europe; North, South, and tropical Africa; Madagascar; Indonesia; Philippines; Mauritius; Reunion; Sri Lanka; India; Seychelles; South America; Mexico; Lebanon; New Guinea; Saudi Arabia; Thailand (ca. 95 spp.) Eukoenenia Börner, 1901 – Europe, Morocco, Algeria, Indonesia, Singapore, Madagascar, Angola, Kenya, Gabon, India, Sri Lanka, Chile, Paraguay, Brazil, Lebanon, Malta, New Guinea, Mexico, etc. (cosmopolitan, ca. 82 spp.) Allokoenenia Silvestri, 1913 – Guinea (one sp.) Koeneniodes Silvestri, 1913 – Madagascar, Guinea, Thailand, Philippines, Indonesia, New Guinea, Mauritius, Reunion, Seychelles, Sri Lanka (eight spp.) Leptokoenenia Condé, 1965 – Congo, Brazil, Saudi Arabia (four spp.) Fam. Prokoeneniidae – Chile, USA, Southeast Asia, Madagascar (seven spp.) Prokoenenia Börner, 1901 – Thailand, Indonesia (Java, Sulawesi), Madagascar, Chile, USA (six spp.) Triadokoenenia Condé, 1991 – Madagascar (one sp.) © Springer International Publishing AG, part of Springer Nature 2018 P. Beron, Zoogeography of Arachnida, Monographiae Biologicae 94, https://doi.org/10.1007/978-3-319-74418-6_6 135 136 6.2 6 Review of the Classification of the Orders of Arachnida Schizomida (According to Manzanilla et al. 2008 – ca. 260 spp.) Fam. Protoschizomidae – Mexico, USA (16 spp.) Agastoschizomus Rowland, 1971 – Mexico (eight spp.) Protoschizomus Rowland, 1975 – Mexico (eight spp.), ? Texas Fam. Hubbardiidae – USA, Mexico, South and Central America, Caribbean, tropical Africa, South Africa, Australia, Philippines, Burma, Malaysia, Java, Sumatra, Thailand, Sri Lanka, Ryukyu Island, Taiwan, New Guinea, Seychelles, Madagascar, Hawaii, greenhouses in Europe Subfam. Megaschizominae – Mozambique, Republic of South Africa (two spp.) Megaschizomus Lawrence, 1969 – Mozambique, Republic of South Africa (two spp.) Subfam. Hubbardiinae – South and Central America, Caribbean, tropical Africa, Australia, Burma, Malaysia, Java, Sumatra, Philippines, Seychelles, Madagascar (121 sp.) Adisomus Cokendolpher et Reddell, 2000 – Brazil (one sp.) Afrozomus Reddell et Cokendolpher, 1995 – Angola, DR Congo (one sp.) Anepsiozomus Harvey, 2001 – Seychelles, Socotra (two spp.) Antillostenochrus Armas et Teruel, 2002 – Cuba, Dominican Republic, Puerto Rico (10 spp.) Apozomus Harvey, 1992 – Australia, Borneo, New Guinea, Ryukyu Island, Taiwan (17 spp.) Artacarus Cook, 1898 – Liberia, Ivory Coast (one sp.) Attenuizomus Harvey, 2000 – Australia (NT, four spp.) Bamazomus Harvey, 1992 – Seychelles, Madagascar, Thailand, Hong Kong, Japan (Ryukyu Island), West Malaysia, Papua New Guinea, Australia, Hawaii (11 spp.) Belicenozomus Armas et Viques, 2010 – Belize and Honduras (two spp.) Brignolizomus Harvey, 2000 – Australia (three spp.) Bucinozomus Armas et Rehfeldt, 2015 – greenhouse in Germany (one sp.) Burmezomus Bastawade, 2004 – Burma (one sp.) Calima Moreno-González et Villarreal, 2012 – Colombia (two spp.) 6.2 Schizomida 137 Caribezomus Armas, 2011 – Jamaica (one sp.) Clavizomus Reddell et Cokendolpher, 1995 – Java, West Malaysia, Singapore (one sp.) Cokendolpherius Armas, 2011 – Cuba (one sp.) Colombiazomus Armas et Delgado-Santa, 2012 – Colombia (two spp.) Cubazomus Reddell et Cokendolpher, 1995 – Cuba (three spp.) Cubacanthozomus Teruel, 2007 – Cuba (one sp.) Draculoides Harvey, 1992 – W. Australia (six spp.) Enigmazomus Harvey, 2006 –Somalia, Arabian Peninsula (two spp.) Gravelyzomus Kulkarni, 2012 – India (one sp.) Guanazomus Teruel et Armas, 2002 – Cuba (one sp.) Hansenochrus Reddell et Cokendolpher, 1995 – Venezuela, Central America, Lesser Antilles (14 spp.) Heterocubazomus Teruel, 2007 – Cuba (one sp.) Heteronochrus Armas et Viquez, 2010 – Guatemala (one sp.) Hubbardia Cook, 1899 – USA (Arizona, California) (nine spp.) Javazomus Reddell et Cokendolpher, 1995 – Java (one sp.) Julattenius Harvey, 1992 – Australia (Queensland)(two sp.) Kenyazomus Armas, 2014 – Kenya (one sp.) Lawrencezomus Armas, 2014 – Cameroon, Liberia (two spp.) Luisarmasius Reddell et Cokendolpher, 1995 – Puerto Rico, Cuba (Pinos) (two spp.) Mahezomus Harvey, 2001 – Seychelles (one sp.) Mayazomus Reddell et Cokendolpher, 1995 – Mexico (seven sp.) Notozomus Harvey, 2000 – Australia (Queensland, 17 spp.) Orientzomus Cokendolpher et Tsurusaki, 1994 – Philippines (Luzon), Japan, Bonin Island (two spp.) Ovozomus Harvey, 2001 – Seychelles, Ceylon, India, Mayotte, Cook Island, Reunion (two spp.) Pacal Reddell et Cokendolpher, 1995 – Mexico (three spp.) Paradraculoides Harvey, Berry, Edward et Humphreys, 2008 – W. Australia (four spp.) Piaroa Manzanilla, Giupponi et Tourinho, 2008 – Venezuela, Colombia, Costa Rica (three spp.) Reddellzomus Armas, 2002 – Cuba (one sp.) Rowlandius Reddell et Cokendolpher, 1995 – Venezuela, Central America, Caribbean, Brazil (55 spp.) Secozomus Harvey, 2001 – Seychelles (one sp.) 138 6 Review of the Classification of the Orders of Arachnida Fig. 6.1 Hubbardia pentapeltis ♀ (Photo and courtesy Marshal Hedin) Sotanostenochrus Reddell et Cokendolpher, 1991 – Mexico (San Luis Potosí, Tamaulipas)(two spp.) Stenochrus Chamberlin, 1922 – USA (Texas), Guatemala, Mexico, Cuba, Puerto Rico, Dominican Republic, Jamaica, South America (21 spp.) (S. portoricensis Chamberlin, 1922 is more widely distributed, imported in Europe) Stenoschizomus González-Sponga, 1997 – Venezuela (one sp.) Stewartpeckius Reddell et Cokendolpher, 1995 – Jamaica (one sp.) Surazomus Reddell et Cokendolpher, 1995 – Costa Rica, South America (six spp.) Tayos Reddell et Cokendolpher, 1995 – Ecuador (one sp.) Trithyreus Kraepelin, 1899 – Burma (Myanmar) (one sp.) Troglocubazomus Teruel, 2003 – Cuba (one sp.) Wuayuuzomus Armas et Colmenares, 2006 – Venezuela (one sp.) Zomus Reddell et Cokendolpher, 1995 – Malaysia (incl. Sarawak), Singapore, Seychelles, Fiji, Samoa, Rodrigues, England (Kew Garden) (one sp.) (Fig. 6.1) 6.3 Thelyphonida (Uropygi) Cooke and Shadab (1973), Gravely (1916), Harvey (2003, 2013d), Haupt (2009b), Huff and Prendini (2009), Mello-Leitão (1931), Rowland (1973a, b, 2002), Speijer (1933, 1936), Viquez and Armas (2005), Werner (1935) 6.3 Thelyphonida (Uropygi) 139 The first comprehensive papers on the group were the revisions of Pocock (1894) and Kraepelin (1897). Important contributions were made also by Thorell, Gravely, Mello-Leitão, Werner, Speijer, Rowland, Haupt, and others. Rowland and Cooke (1973) list 85 species in the order. Haupt (2009a) synonymized Abaliella Strand, Minbosius Speijer, and Tetrabalius Thorell with Thelyphonus Latreille. Krehenwinkel et al. (2009) described the new genus Thelyphonoides from Panay (Philippines). According to the calculations of Harvey (2002b, 2003, 2013d) and the additions since this time, there are ca. 110 species in the order Thelyphonida (Uropygi), belonging to 15 genera and one recent family (Blick and Harvey 2011). They are distributed as follows: Fam. Thelyphonidae – Southeast Asia, Indonesia, Philippines, Samoa, New Guinea, Fiji, Solomon Island, Vanuatu, India, Sri Lanka, China, North, South, and Central America, Caribbean Islands, West Africa (ca.110 spp.) Subfam. Thelyphoninae – Southeast Asia, Philippines, Samoa, New Guinea, Fiji, Solomon Island, India (49 spp.) Ginosigma Speijer, 1936 – Sunda Islands, Thailand, Laos, Cambodia, Vietnam (two spp.) Glyptogluteus Rowland, 1973 – Philippines (Panay)(one sp.) Thelyphonoides Krehenwinkel et al., 2009 – Philippines (Panay)(one sp.) Thelyphonus Latreille, 1802 (Chajnus Speijer, 1936; Abaliella Strand, 1928; Minbosius Speijer, 1936; Tetrabalius Thorell, 1889; fide Haupt 2009a) – Indonesia, Singapore, Philippines, Vanuatu (New Hebrides), Fiji, Rennell, Solomon Islands, New Guinea, New Britain, Burma, Sri Lanka, India, Thailand, W. Samoa, Borneo, Moluccas, Cambodia, Vietnam (45 spp.) Subfam. Hypoctoninae – Burma (Myanmar), South China, Malaysia, West Africa, India, Sri Lanka, Brazil, Guyana, Venezuela, Dominican Republic (27 spp.) Etienneus Heurtault, 1984 – Western Africa (one sp.) Hypoctonus Thorell, 1889 – Burma (Myanmar), South China, Malaysia, Thailand, Bangladesh, Java, India (19 spp.) Labochirus Pocock, 1894 – India, Sri Lanka (four spp.) Thelyphonellus Pocock, 1894 – Brazil, Guyana, Venezuela (two spp.) Ravilops Viquez et Armas, 2005 – Dominican Republic (one sp.) Subfam. Mastigoproctinae (Uroproctinae) – Brazil, Cuba, Colombia, El Salvador, Costa Rica, Honduras, Belize, Venezuela, Mexico, Guatemala, Haiti, Martinique, USA, Hainan, Philippines (23 spp.) 140 6 Review of the Classification of the Orders of Arachnida Uroproctus Pocock, 1894 – India (Assam), Cambodia (one sp.) Mimoscorpius Pocock, 1894 – Guatemala (one sp.) Mayacentrum Viquez et Armas, 2006 – Guatemala, El Salvador, Honduras, Belize (three spp.) Mastigoproctus Pocock, 1894 (Amauromastigon Mello-Leitão, 1931 – Brazil) – Cuba, Brazil, Colombia, Venezuela, USA, Mexico, Haiti, Martinique (17 spp.) Valeriophonus Viquez et Armas, 2005 – Costa Rica (one sp.) Subfam. Typopeltinae – China, Taiwan, Hainan, Japan, Thailand, Vietnam, Laos (11 spp.) Typopeltis Pocock, 1894 (Teltus Speijer, 1936) – China, Taiwan, Hainan, Japan, Thailand, Laos, Vietnam; doubtful finding in the Far East of Russia (11 confirmed spp.) 6.4 Amblypygi These big and conspicuous, largely cavernicolous, dwellers of the warmer places have been subject to many articles (and many synonyms) by earlier researchers, starting with Linnaeus, Lamarck, Herbst, and Gerstaecker, and also the researchers of nineteenth and early twentieth centuries (Gervais, C.L. Koch, L. Koch, Bilimek, Karsch, Pocock, Butler, Kraepelin, Simon, Gravely, Thorell, Hansen, and others). Many contributions have been made in the last 90 years by Franganillo, MelloLeitão, Fage, Lawrence, Dunn, Whittick, Caporiacco, Moreno, Ravelo, Kritscher, Rowland, Armas, González-Sponga, Delle Cave, Harvey, Rahmadi, and other authors. The papers of Mullinex and Quintero and especially the revisions of Weygoldt are the basis of the modern understanding of the order and counting (Harvey 2003, 136 spp.; Blick and Harvey 2011; Miranda and Giupponi 2011; Harvey 2013c, actualized 2017, 184 spp.; Miranda et al. 2018, 220 sp.) of 18 genera and five families. Suborder Euamblypygi Fam. Charinidae – Greece (Rhodes, Kos), Turkey, Israel, Egypt, Cuba, Jamaica, Galapagos Islands, US Virgin Islands, tropical South America (Guiana, Brazil, Suriname, Venezuela, Peru), Australia, Andaman Islands, India, W. Samoa, Vanuatu, Oman, Indonesia (Java, Borneo), Singapore, Malaysia, Burma, Vietnam, Laos, Cambodia, Papua New Guinea (incl. New Britain), Seychelles, Madagascar, New Caledonia, Solomon Islands, tropical Africa (Somalia, Tanzania, Guinea, Equatorial Guinea, Sao Tome, Principe, Somalia, Socotra, Pakistan) (90 spp.) 6.4 Amblypygi 141 Catageus Thorell, 1889 (= Stygophrynus Kraepelin, 1896)– Burma, Solomon Islands, Thailand, Laos, Cambodia, Vietnam, Indonesia, Malaysia (nine sp.) (nine sp.) Weygoldtia Miranda, Giupponi, Prendini et Scharff, 2018 - one sp. W. davidovi (Fage, 1946) - Laos, Cambodia, Vietnam Charinus Simon, 1892 (Charinides Gravely, 1911; Enantiosarax Mello-Leitão, 1931; Oligacanthophrynus Caporiacco, 1947; Lindosiella Kritscher, 1959; Speleophrynus Ravelo, 1975; Tricharinus Quintero, 1986) – Greece (Rhodes, Kos), Turkey, Israel, Cuba, Jamaica, Guiana, Brazil, Suriname, Venezuela, Peru, Galapagos Islands, US Virgin Islands, Australia, Oman, Andaman Islands, India, W. Samoa, Vanuatu, Indonesia (Java, Borneo), Singapore, Malaysia, Papua New Guinea, Seychelles, Madagascar, New Caledonia, Solomon Islands, Tanzania, Guinea, Equatorial Guinea, Sao Tome, Principe, Somalia, Egypt, Socotra, Pakistan (74 spp.) Sarax Simon, 1892 (Phrynichosarax Gravely, 1915) – Solomon Islands, Malaysia, Singapore, Philippines (Luzon), Indonesia (Java, Kalimantan), India, Andaman Islands, Papua New Guinea (incl. New Britain and New Ireland), Vietnam, Laos, Cambodia, Borneo (17 spp.) Fam. Charontidae – Australia (incl. Christmas Island), Indonesia (Java, Maluku, Sumbawa), Malaysia, Burma, Thailand, Vietnam, Palau, Papua New Guinea, Philippines, Singapore, Solomon Islands (nine spp.) Charon Karsch, 1879 – Australia (Northern Territory, Queensland, Christmas Island), Indonesia (Java, Maluku, Sumbawa), Malaysia (incl. Borneo), Palau, Papua New Guinea, Philippines, Singapore, Solomon Islands (nine spp.) Fam. Phrynichidae – tropical and Southern Africa, Algeria, Morocco, Yemen, Saudi Arabia, Oman, Mauritius, Seychelles, Zanzibar, Sao Tome, Madagascar, India, Sri Lanka, Brazil (33 spp.) Subfam. Damoninae – tropical and Southern Africa, Algeria, Morocco, Yemen (12 spp.) Damon C.L. Koch, 1850 (Titanodamon Pocock, 1894; Nanodamon Pocock, 1894) – South Africa, Malawi, Mozambique, Zambia, Zimbabwe, Ethiopia, Kenya, Somalia, Tanzania, Yemen, Angola, Namibia, Cameroon, Congo, DR Congo, Equatorial Guinea, Gabon, Nigeria, Liberia, Ivory Coast, Sao Tome, Mali, Ghana, Benin, Senegal, Sierra Leone, Togo, Botswana, Sudan (10 spp.) Musicodamon Fage, 1939 – Algeria, Morocco (one sp.) 142 6 Review of the Classification of the Orders of Arachnida Phrynichodamon Weygoldt, 1996 – Namibia, South Africa (one sp.) Subfam. Phrynichinae – Brazil, Kenya, Tanzania, Zanzibar, DR Congo, Madagascar, Malawi, Zimbabwe, Angola, Namibia, Eritrea, Somalia, Djibouti, Kenya, Sri Lanka, Yemen, Saudi Arabia, Oman, Socotra, Mauritius, Seychelles (21 spp.) Euphrynichus Weygoldt, 1995 – Kenya, Tanzania (incl. Zanzibar), DR Congo, Madagascar, Malawi, Zimbabwe (two spp.) Phrynichus Karsch, 1879 (Myodalis Simon, 1936) – Eritrea, Somalia, Djibouti, Kenya, India, Sri Lanka, Yemen, Saudi Arabia, Oman, Madagascar, Mauritius, Socotra, Seychelles (16 spp.) Trichodamon Mello-Leitão, 1935 – Brazil (two spp.) Xerophrynus Weygoldt, 1996 – Angola, Namibia (one sp.) Fam. Phrynidae – tropical South America, USA (Arizona, California, Florida), Central America, Bahamas, Cuba, Barbados, St. Vincent, Netherlands Antilles, Grenades, Haiti, Saint Lucia, Mexico, Indonesia (Flores) (62 spp.) Subfam. Heterophryninae – tropical South America (14 spp.) Heterophrynus Pocock, 1894 – Brazil, Peru, Colombia, French Guiana, Guyana, Venezuela, Ecuador, Suriname (14 spp.) Subfam. Phryninae – tropical South America, USA (Arizona, California, Florida, Texas), Mexico, Central America, Bahamas, Cuba, Barbados, St. Vincent, Netherlands Antilles, Grenades, Haiti, Saint Lucia, Indonesia (Flores) (48 spp.) Acanthophrynus Kraepelin, 1899 – Mexico, ? USA (Arizona, California)(one sp.) Paraphrynus Moreno, 1940 – USA (Arizona, Florida), Mexico, Cuba, Guatemala, Colombia, Costa Rica, El Salvador, Nicaragua, Panama, Honduras, Bahamas (16 spp.) Phrynus Lamarck, 1801 (Admetus C.L. Koch; Neophrynus Kraepelin, 1895) – Mexico, Barbados, St. Vincent, Nicaragua, Colombia, Ecuador, Guyana, Cuba, Dominican Republic, Virgin Islands, Jamaica, Martinique, Cayman Islands, Antigua, Barbuda, Bermuda, Bahamas, Guatemala, Honduras, Belize, Costa Rica, USA (Texas, Florida), Trinidad, Panama, Venezuela, Netherlands Antilles, Grenadines, Puerto Rico, Haiti, Saint Lucia, Brazil, Indonesia (Flores) (31 spp.) 6.5 Ricinulei 143 Suborder Palaeoamblypygi Fam. Paracharontidae – Guinea-Bissau (one sp.) Paracharon Hansen, 1921 – Guinea-Bissau (one sp.) 6.5 Ricinulei This small (3 genera, 55 spp., according to Harvey 2003; 64 spp. after Cook 2008; 67 after Tourinho et al. 2010; 58 after Prendini 2011) group of arachnids is sometimes considered representing “living fossils” (Selden 1986). The studies on recent Ricinulei start in 1838 (Guérin-Méneville). Longtime they are considered very rare, thanks to the technique of collecting. From the 66 living valid species, recorded up to 2011, 49 have been described after 1950 by Roewer, Platnick, Shadab, Cooke, Cooreman, González-Sponga, Gertsch, Brignoli, Cor. Gutierrez, Dumitresco and Juvara-Balş, Merrett, Marquez and Conconi, Goodnight and Goodnight, Armas, Legg, Naskrecki Cokendolpher, and other researchers. The increase is mostly to the expense of the American genera Cryptocellus and the newly described Pseudocellus (44 new species). In 60 years only five new species have been added to the six African Ricinoides known before 1950 (Legg, 1976, 1977 from Sierra Leone, 1982 from Ivory Coast, Naskrecki, 2008 from Ghana, Penney et al. from Gambia). Wunderlich (2012) downgraded the suborders Neoricinulei Selden, 1992, and Palaeoricinulei Selden, 1992, to superfamilial rank, the Poliocheroidea Scudder, 1884, resp. Ricinoidoidea Ewing, 1929. Suborder Neoricinulei Selden, 1992 Fam. Ricinoididae (? 66 spp.) Cryptocellus Westwood, 1874 (Heteroricinoides Dumitresco et Juvara-Balş, 1977) – Colombia, Brazil, Guyana, Panama, Venezuela, Suriname, Costa Rica, Honduras, Nicaragua, Ecuador, Peru, Tobago (34 spp.) Pseudocellus Platnick, 1980 – USA (Texas), Mexico, Guatemala, Honduras, El Salvador, Panama, Cuba (21 spp.) Ricinoides Ewing, 1929 – Gambia, Guinea, Guinea-Bissau, Nigeria, Ghana, Cameroon, Sierra Leone, Equatorial Guinea, Benin, Congo, Togo, Ivory Coast (11 spp.) 144 6.6 6 Review of the Classification of the Orders of Arachnida Scorpiones According to Dupré (2010), there are about 1950 valid species of scorpions, belonging to 152 genera and 7–20 families (18 fam. + Akravidae, after Prendini 2011), but the latest number of species (after Jan Ove Rein 2017, The Scorpion files) is 2326. The present arrangement follows generally the systems of Soleglad and Fet (2003) and Lourenço (2000b), but also takes into account the objections of Prendini and Wheeler (2005) and the development of the taxonomy after 2005. The number and contents of higher taxa in the paper of Soleglad and Fet were strongly disputed by Prendini and Wheeler (2005). Still, as these authors point “The suprageneric classification of recent (including extant) scorpions is in a state of flux.” This flux is more obvious than in any other of the arachnid orders. The present author finds this situation quite unfortunate for the zoogeography, which should be based on good supraspecific taxonomy for explaining endemism, distribution, and the whole history of the order. And this is particularly true for scorpions, considered very important biogeographically. Vachon (1968) emphasizes that “Scorpions can play an important part in biogeography. These animals travel slowly…Scorpions thus evolve in situ and are practically immune to variations in micro-climate. Thus they constitute excellent biogeographic test and their presence (or absence) provides a better understanding or interpretation of the population background of a given region.” Fet and Soleglad (2014) reject the criticism of Prendini and Wheeler (2005). As it is not to us to solve here the problem, we shall use the list of families in Prendini (2011), considering Akravidae recent family, without discussing the higher taxa and the subfamilies. Nenilin and Fet (1992) analyzed the zoogeography of the world scorpion fauna on the level of 1985–1986, but since that time many changes occurred in the nomenclature of scorpions and new data were obtained. Numbers of Families in Classifications of Recent Scorpions Proposed During the Past 30 Years (Partly from Prendini and Wheeler 2005) Lamoral (1980) – 7 Stockwell (1989) – 13 Sissom (1990) – 9 Kovařik (1998) – 13 Fet et al. (2000) – 16 Lourenço (2000b) – 20 Soleglad and Fet (2003) – 14 Soleglad et al. (2005) – 13 Prendini and Wheeler (2005) – 18 Prendini (2011) – 18 (+ Akravidae ? extinct) Present paper – 20 6.6 Scorpiones 145 Superfam. Bothriuroidea Fam. Bothriuridae – South America, Namibia, Australia, Indian Himalaya (12 gen., 138 spp.) Superfam. Buthoidea Fam. Buthidae – all continents, tropical and subtropical countries, and in some countries with moderate climate (Spain, France, Cyprus)(70 genera, 939 spp.) Fam. Microcharmidae – Madagascar (one gen., six spp.) (synonymized with Buthidae by Volschenk et al. 2008 ?) Superfam. Chaeriloidea Fam. Chaerilidae – Bangladesh, India, Indonesia, Malaysia, Nepal, Singapore, Sri Lanka, Vietnam (1 gen., 43 spp.) Fam. Pseudochactidae – Uzbekistan, Tajikistan, Afghanistan, caves of Laos, and Vietnam (three gen., five spp.) Superfam. Chactoidea Fam. Chactidae – Central and South America, Mexico (Baja California) (9 genera, 178 spp.) Fam. Euscorpiidae – South Europe, North Africa, Middle East, Central Asia (south), Mexico, Guatemala (4 gen., 25 spp.) Fam. Scorpiopidae – Afghanistan, Southeast Asia, India, Pakistan, Indonesia, Malaysia (5 gen., 48 spp.) Fam. Troglotayosicidae – Pyrenees (France/Spain)(?), Equatorial Amazonas (? two genera, three spp.) Fam. Superstitioniidae – Mexico, Southeast of the USA (four genera) Superfam. Vaejovoidea Fam. Iuridae – North and South America, Turkey, Greece (6 gen., 36 spp.) Fam. Vaejovidae – Mexico, South of the USA (17 gen., 181 spp.) Superfam. Scorpionoidea Fam. Hemiscorpiidae – Middle East (2 gen, 13 spp.) Fam. Heteroscorpionidae – Madagascar (one gen., six spp.) Fam. Hormuridae (Liochelidae)(incl. Hadogenidae) – Australia, Africa, Central and South America, Antilles, Southeast Asia, Tibet, Pacific islands, Madagascar, India, Indian Ocean Islands, Seychelles, Mauritius, Hispaniola, Pacific Cocos Island (9 gen., 69 spp.) Fam. Scorpionidae (incl. Diplocentridae) – North America, Central America, the northern part of South America, Antilles, Middle East, Africa, East, South and Southeast Asia, Indonesia (12 gen., 248 spp.) Fam. Akravidae – Israel (one sp.) Fam. Lisposomidae – South Africa (one genus) Fam. Urodacidae – Australia (2 gen., 21 spp.) 146 6.7 6 Review of the Classification of the Orders of Arachnida Pseudoscorpiones More than 3538 described species worldwide (Harvey 2002, 2013f, actualized). After the catalogue of Harvey (1990) and of his revision of 1992 and reviews of 2002, 2011, and 2013f and the additions of Mahnert (several papers) and others, it is generally accepted that the order includes 2 suborders and 26 recent families with more than 454 genera. Suborder Epiocheirata Fam. Chthoniidae – Cosmopolitic, ca. 650 spp., 28 gen. Fam. Tridenchthoniidae (Dithidae) – South Africa, Australia, New Caledonia, Lord Howe Island, Ecuador, Cuba, Jamaica, Mexico, Costa Rica, Trinidad, Peru, Suriname, Paraguay, Argentina, Brazil, Venezuela, Equatorial Africa, Ethiopia, Caroline Islands, India, Vietnam, Philippines, Seychelles, Indonesia (Moluccas, Sumatra, Java), New Guinea, Japan, Palau Island, Bhutan, Nepal, Laos, Marcus Island, Malaysia, South Africa, Sao Tome, USA (15 gen., 71 spp.) Fam. Pseudotyrannochthoniidae – Australia, Tasmania, Korea, Eastern USA, Japan, Chile, China (5 gen., 49 spp.) Fam. Lechytiidae – Turkey, USA, Mexico, Chile, Peru, Argentina, Paraguay, Galapagos Islands, Jamaica, Dominican Republic, Trinidad, Guadeloupe, Martinique, Congo, D.R. Congo, Kenya, Tanzania, Nepal, Vietnam, India, Juan Fernandez Islands, South Africa, Caroline Islands, Marshall Islands, Marcus Island (1 gen., 23 spp.) Fam. Feaellidae – tropical and Southern Africa, Seychelles, Madagascar, India, Sri Lanka, Australia (1 gen., 12 spp.) Fam. Pseudogarypidae – North America, Tasmania, Baltic Amber from the Oligocene (two genera, seven recent species) Suborder Iocheirata Fam. Ideoroncidae – Argentina, Brazil, Paraguay, Chile, US Virgin Islands, Mexico, USA, Afghanistan, Iran, Malaysia, India, Cambodia, Thailand, Vietnam, Indonesia (Sumatra), Sri Lanka, Kenya, Tanzania, Uganda, Congo, Zimbabwe, Botswana (13 gen., 70 spp.) Fam. Bochicidae (Vachoniidae) – Central America as far north as Texas, Cuba, Grenada, Trinidad, Jamaica, Spain, Portugal (14 gen., 44 spp.) 6.7 Pseudoscorpiones 147 Fam. Hyidae – Philippine Islands, Indonesia (Java, Sumatra), Sri Lanka, India, Australia (2 gen., 14 spp.) Fam. Gymnobisiidae – Southern South America, South Africa (4 gen., 11 spp.) Fam. Neobisiidae – Europe, Corsica, Sicily, Sardinia, Balearic Islands, Canary Islands, Malta, Georgia, Armenia, Azerbaijan, Azores, Iran, Turkey, Turkmenistan, Kyrgyzstan, Far East of Russia, Tibet, Nepal, India, Philippines, Mongolia, Afghanistan, Vietnam, Thailand, Japan, Korea, Burma, China, Taiwan, Canada, USA, DR Congo, Kenya, Rwanda, Tanzania, Zimbabwe, Algeria, Morocco, Cyprus (33 genera, 595 spp.) Fam. Syarinidae – South America, Antillean Islands, Mexico, USA, Canada, Italy, Greece, Norway, Finland, Austria, Germany, France, Switzerland, Crimea, Algeria, Balearic Islands, Canary Islands, Lebanon, Malta, Morocco, Portugal, Sardinia, Sicily, Spain, Bhutan, India, Sri Lanka, Nepal, New Caledonia, Tuvalu, Philippines, Solomon Island, New Zealand, New Guinea, tropical Africa, South Africa, Seychelles, Galapagos Island (18 genera, 111 spp.) Fam. Parahyidae – Caroline Islands, Singapore (one genus, one sp.) Fam. Garypidae (Synsphyronidae) – Armenia, Algeria, Balearic Island, Crete, Egypt, Canary Island, Corsica, France, Greece, Ilhas Selvagens, Israel, Italy, Malta, Libya, Sardinia, Spain, Portugal, Tunisia, ex-Yugoslavia, Bonaire, Aruba, Mustique Island, Mexico, USA (California, Florida), India, Seychelles, South Africa, Japan, Indonesia (Krakatau, Nicobar Island), Maldives Island, Sri Lanka, Kenya, Somalia, Yemen, Namibia, South Africa, Zimbabwe, Madagascar, Marshall Islands, Ascension Island, Australia (incl. Tasmania), New Zealand (10 genera, 80 spp.) Fam. Geogarypidae – tropical Africa, South Africa, Aldabra, Seychelles, Indonesia, Malaysia, India, Sri Lanka, Bhutan, China, Taiwan, Argentine, Chile, Brazil, Paraguay, Guyana, Grenada, Juan Fernandez Island, Canary Island, Algeria, Morocco, Australia, Russia, Kazakhstan, Kyrgyzstan, Turkmenistan, Thailand, Caroline Island, Mariana Island, Marshall Island, Solomon Island, Tuvalu, Marquesas Island, Marcus Island, Europe, Turkey, Malta, Sardinia, Sicily, Madeira, Hawaii, Israel, Iran (3 gen., 60 spp.) Fam. Larcidae – USA, Spain, Italy, Austria, Denmark, Poland, Romania, Sweden (2 genera, 15 spp.) Fam. Cheiridiidae – Chile, Guadeloupe, Cuba, Aruba, Curacao, Namibia, tropical Africa, South Africa, Lesotho, Russia, Estonia, Finland, Azerbaijan, China, Taiwan, Korea, Nepal, Philippines, Japan, Hawaii, Papua New Guinea, Salomon Islands, Mariana Islands, Australia, New Zealand; for Ch. museorum Leach, Europe, India, Mozambique, DR Congo, Russia, Latvia, Kyrgyzstan, Turkey, South Africa, USA (7 gen., 73 spp.) 148 6 Review of the Classification of the Orders of Arachnida Fam. Pseudochiridiidae – Madagascar, Aldabra, tropical Africa, India, Andaman Island, Burma, Indonesia, Malaysia, Nepal, Philippines, New Guinea, Brazil (2 gen., 12 spp.). Fam. Olpiidae – Australia, New Zealand, Norfolk Island, Aldabra Islands, Madagascar, Algeria, Seychelles, USA (Florida), Lesser Antilles, Jamaica, South America, Turkey, Indonesia, India, Burma, Turkmenistan, Tajikistan, Uzbekistan, Japan, Cambodia, Vietnam, Cyprus, Yemen, Iran, Iraq, Jordan, Israel, Lebanon, Corsica, France, Italy, Sardinia, Madeira Islands, Greece, Croatia, Papua New Guinea, New Caledonia, Salomon Islands, Tuvalu, Samoa, Caroline Islands, Mariana Islands, Marshall Islands, tropical Africa, Namibia, South Africa, Cabo Verde, Egypt, Djibouti (36 gen., 268 spp.) Fam. Garypinidae – Peru, Chile, USA, Mexico, South America, Jamaica, Galapagos Islands, Solomon Islands, Hawaii, Australia, New Guinea, Israel, Syria, Turkey, Yemen, Morocco, Cyprus, Crete, Greece, Corfu, Italy, Sicily, Croatia, Hawaii, South Africa, Yemen, Iran, Indonesia, Thailand, Cambodia, Vietnam, and others (21 gen., 76 spp.) Fam. Menthidae – Mexico, USA (California), Argentina, Chile, Israel, Socotra, Australia (5 gen., 12 spp.) Fam. Sternophoridae – Ethiopia, Australia, Papua New Guinea, India, Sri Lanka, Laos, Cambodia, Vietnam, El Salvador, Dominican Republic, USA (Florida), Mexico, Cayman Islands (3 gen., 20 spp.) Fam. Withiidae – South Africa, tropical Africa, Seychelles, Cabo Verde, Sao Tome, St. Helena, New Zealand, Lord Howe Island, Juan Fernandez Islands, USA (Texas), South America, Mexico, Cuba, Virgin Islands, Vietnam, Indonesia, Christmas Island, India, Cambodia, Malaysia, Thailand, Philippines, Solomon Islands, Saudi Arabia, Pakistan, Iran, Nepal, China, Taiwan, Mongolia, Japan, Yemen, Israel, Morocco, Tunisia, Libya, USA, Mariana Islands, Marshall Islands, Papua New Guinea, Greece, Bulgaria, Russia, France, Corsica, Italy, Sardinia, Portugal, Spain, Switzerland, Turkey, Georgia, ex-Yugoslavia; only for Withius piger, Algeria, Australia, Austria, Balearic Island, Canary Islands, Chad, Chile, China, Cuba, Denmark, Egypt, France, Ghana, Great Britain, Hungary, India, Italy, Libya, Malta, Mexico, Morocco, Netherlands (38 gen., 158 spp.) Fam. Cheliferidae – South Africa, tropical Africa, North Africa, Socotra, Japan, Europe, Malta, Madeira, Canary Islands, Australia, New Zealand, Papua New Guinea, USA, Mexico, Guatemala, Virgin Islands, Dominican Republic, Cuba, Brazil, Ecuador, Mauritius, Reunion, Indochina, India, China, Mongolia, Indonesia, Afghanistan, Pakistan, Kyrgyzstan, Kazakhstan, Tajikistan, Uzbekistan, Iran, Iraq, Saudi Arabia, Israel, Jordan, Yemen (58 gen., 273 spp.) 6.8 Opiliones 149 Fam. Atemnidae (Miratemnidae) – Southeast Asia, Central Asia, Mongolia, Japan, Philippines, Bhutan, Nepal, India, Nicobar Islands, Solomon Islands, Sri Lanka, Indonesia, China, North Africa, tropical Africa, Yemen, Sao Tome, Comoro Islands, Seychelles, Madagascar, Europe, Cyprus, Canary Islands, Balearic Islands, Sardinia, Sicily, Puerto Rico, Brazil, Ecuador, Guyana, Cuba, Dominican Republic, St. Vincent, Costa Rica, US Virgin Islands, Brazil, Argentina, Peru, Colombia, Panama, Venezuela, Haiti, Paraguay, St. Vincent, St. Paul Island, Ilhas Selvagens, USA (Arizona), New Caledonia, Lord Howe Islands, Caroline Islands, Mariana Islands, Christmas Island, Marquesas Islands, Marshall Islands, Papua New Guinea (21 gen., 178 spp.) Fam. Chernetidae (Myrmochernetidae) – Europe; South America; Mexico; USA; Central America, Antilles; San Ambrosio Island; New Guinea; Solomon Islands; Mongolia; China, Japan; North, Central, and South Africa; New Zealand; Australia; etc. (117 gen., 663 spp.) 6.8 Opiliones The large order Opiliones (46 families, 1649 extant genera, and more than 6000 species, according to Harvey 2002 and Pinto-da-Rocha and Giribet 2007; 6534, acc. to Kury 2013, 6653 acc. to Kury, April 2017) was traditionally subdivided into three suborders: Cyphophthalmi, Laniatores (mostly tropical), and Palpatores (mostly Holarctic). Recently, the “Palpatores” are firmly considered as two suborders – Eupnoi and Dyspnoi (Giribet and Kury, Pinto-da-Rocha and Giribet 2007; Kury 2011, 2013). The knowledge on the Opiliones was first summarized by Roewer (1923), who described 2260 valid species and greatly enlarged the knowledge on Opiliones by many publications since that time. Unfortunately, there is a strong suspicion that some localities of Roewer’s material have been confused (von Helversen and Martens (1972), von Helversen (1968)) and should be considered very carefully when zoogeographical conclusions are made. It is always wiser to use the data of the modern authors with reliable localities. Classification of Opiliones After Kury (2013, April, 2017). The Number of Species in the Families Is Not Updated Cyphophthalmi (6 fam., ca. 36 genera, 208 spp.). The immense progress in the study of Cyphophthalmi can be assessed, comparing the present day picture and the biogeographical and taxonomical outline of the suborder of Juberthie and Massoud (1979). Their paper starts with the passage: “Les Opilions du sous-ordre des 150 6 Review of the Classification of the Orders of Arachnida Cyphophthalmes représentent une petite famille de 15 genres et 56 espèces….réparties sur l’ensemble du globe.” Fam. Stylocellidae – Southeast Asia (from India to New Guinea) (6 gen., 36 spp.) Fam. Ogoveidae – Western Equatorial Africa (one gen., three spp.) Fam. Neogoveidae – South America (Colombia, Brazil, Ecuador, Guyana, French Guiana, Peru, Suriname, Trinidad and Tobago, Venezuela), USA (Florida, Georgia, South Carolina), West Africa (Bioko, Gabon, Sierra Leone) (9 gen., 22 spp.) (+many undescr.) Map 6.1 Distribution of Pettalidae (▼) and Neogoveidae (■) in South America and the southern part of North America Superfamily Sironoidea Fam. Pettalidae – New Zealand, Australia, Chile, South Africa, Sri Lanka (9 gen., 61 spp.) Fam. Sironidae – South Europe, Corsica, Carpathians, Asia Minor, USA (Western and Eastern shore), Japan, ? Kenya (7 gen., 52 spp.) (Fig. 6.2) 6.8 Opiliones Fig. 6.2 Fam. Troglosironidae – New Caledonia (1 gen., 13 spp.) Troglosiro longifossa Sharma et Giribet (courtesy of G. Giribet) Map 6.2 Distribution of Sironidae (♦) in Europe and Western Asia 151 152 6 Review of the Classification of the Orders of Arachnida Map 6.3 Distribution of Neogoveidae (●) and Sironidae (♦) in North America Map 6.4 Distribution of Stylocellidae (►), Pettalidae (▼), and Sironidae (■) in Southeast Asia 6.8 Opiliones 153 Map 6.5 Distribution of Pettalidae (▼– Australia, New Zealand) and Troglosironidae (■ – New Caledonia) Suborder Eupnoi (4 fam., ca. 1810 spp.) Fam. Phalangiidae – Europe; North and South America; Asia Minor; China; Central Asia; North, Central, and South Africa; Canary Islands; Madeira; Borneo (56 gen., 486 spp.). Hesperopilio Shear (Australia, Chile) is nested in Phalangioidea without familial attachment Fam. Sclerosomatidae – Europe; Asia; Africa; South, Central, and North America; Antilles; Australia; New Guinea; Solomon Island (153 gen., ca. 1300 spp.) Fam. Neopilionidae (incl. Monoscutidae) – South Africa, Chile, Argentina, Brazil, Australia, Tasmania, New Zealand, Auckland, Snares, Campbell Islands (20 gen, 64 spp.) Fam. Protolophidae – the Western United States (eight spp.) Suborder Dyspnoi (9 fam., 387 spp.) Fam. Caddidae – South Africa, Eastern North America, Japan (two gen., nine rec. spp.) Fam. Acropsopilionidae – Australia, North America, South America, Japan, New Zealand (3–4 gen., 14 spp., incl. Hesperopilio Shear) Fam. Ischyropsalididae (incl. Ceratolasmatidae) – Europe (incl. Abkhazia, Crimea), Algeria, Tajikistan, USA (Oregon, Washington, California, Idaho), Mexico, Honduras (3 gen., 45 spp.) (Fig. 6.3) 154 6 Review of the Classification of the Orders of Arachnida Fig. 6.3 Ischyropsalis hellwigii (Panzer) (courtesy of Ula N., from A. Kury) Fam. Sabaconidae – Japan, France, Spain, Alaska, Nepal, China (Sichuan), Siberia, USA (Alabama, Kentucky, Oregon, California) (1 gen., 35 spp.) Fam. Taracidae – USA, Canada, Far East of Russia (3 gen., 15 spp.) Fam. Dicranolasmatidae – Mediterranean, Asia Minor, Hungary, Russia (Krasnodar), Iraq, Western North Africa (1 gen., 16 spp.) Fam. Trogulidae – Western and Southern Europe, Western North Africa, the Levant, Caucasus, Northern Iran (4 gen., 56 spp.), introduced in the USA (Fig. 6.4) Fam. Nemastomatidae – USA (Washington, California), Mexico, Coronado Island, Thailand, Japan, Bulgaria, Abkhazia, Crimea, Europe, Algeria, Tajikistan, Asia Minor, Crete (21 gen., 173 spp.) (Fig. 6.5) Fam. Nipponopsalididae – Japan, Korea (one gen., three spp.) Suborder Laniatores According to the Catalogue of Kury (2003), the suborder Laniatores counts 1317 genera and 3901 spp., including 746 genera and 2372 spp. in the New World. Kury (2013) defines the number of Laniatores at 4183 spp., Kury (2016) at 4212 spp., and Kury (status April 2017) at 4248 spp. The Laniatores in Europe are very few – the 6.8 Opiliones 155 Fig. 6.4 Trogulus nepaeformis (Scopoli) (courtesy Ivo Antušek, from A. Kury) Fig. 6.5 Nemastoma bimaculatum (Fabricius) (courtesy of Jonathan Michaelson, from A. Kury) 156 6 Review of the Classification of the Orders of Arachnida families Travuniidae, Cladonychiidae, and Phalangodidae (Roewer 1935; Rambla 1978; Kury and Mendes 2007, and others). Fam. Nippononychidae – Japan, Korea (four genera, 10 spp.) Fam. Paranonychidae – USA, Canada, Japan, Korea (7 genera, 26 spp.) Fam. Travuniidae – Serbia, Croatia, S. France, Sardinia, USA (Washington), Japan (13 gen., 41 spp.) Fam. Synthetonychiidae – New Zealand (1 genus, 14 spp.) Fam. Triaenonychidae – North and South America, Australia, Tasmania, New Zealand, New Caledonia, Japan, Korea, Madagascar, South Africa (107 gen., 477 spp.) Fam. Assamiidae – Africa, Southern Asia; subfam. Dampetrinae to Indonesia, New Guinea, and Australia (267 gen., 483 spp.) Fam. Pyramidopidae – West Africa, Tanzania, Canary Island, Belize (15 gen., 47 spp.) Fam. Epedanidae (incl. Dibuninae, Acrobuninae, Sarasiniciinae, Sarasinellinae) – Philippines, New Guinea, Thailand, Sulawesi, Borneo, Sumatra, Japan, Malaysia, India, Taiwan, China, Vietnam (70 gen., 174 spp.) Fam. Petrobunidae – Southeast Asia (two gen., five spp.) Fam. Podoctidae (incl. Erecananinae and Ibaloniinae) – Africa, Madagascar, Reunion, Japan, Palau, Taiwan, Mauritius, Seychelles, Indonesia, Malaysia, New Guinea, Cuba, India, Philippines, Vanuatu (54 gen., 132 spp.) Fam. Sandokanidae [Oncopodidae, praeoccup.] – Philippines, Sumatra, Borneo, Burma, Nepal, India, Malaysia, Singapore, Thailand (6 gen., 72 spp.) Fam. Tithaeidae – Southeast Asia (6 gen., 39 spp.) Fam. Agoristenidae – Antilles, South America (27 gen., 76 spp.) Fam. Cosmetidae – From Argentina to Southern USA (125 gen., 719 spp.) (Fig. 6.6) Fam. Cranaidae – South America, Trinidad, Panama, Costa Rica (75 gen., 165 spp.) Fam. Gonyleptidae – From Patagonia to Costa Rica, Guatemala, and Falklands (272 gen., 829 spp.) (Fig. 6.7) Fam. Manaosbiidae – South America, Panama, Costa Rica, Trinidad, Windward Islands (27 gen., 47 spp.) Fam. Stygnidae – Northern South America, Lesser Antilles (30 gen., 104 spp.) Fam. Stygnopsidae (Isaeinae, Troglostygnopsinae) – Mexico, Guatemala, Belize, El Salvador, USA (Texas) (9 gen., 40 spp.) Fam. Phalangodidae – USA, Japan, Mediterranean (21 gen., 116 spp.) 6.8 Opiliones 157 Fig. 6.6 Sibambea sp. (Cosmetidae, Ecuador) (courtesy of A. Anker, from A. Kury) Fig. 6.7 Bourguyia bocaina Yamaguti et Pinto-da-Rocha (courtesy J. Wiley & Sons, from A. Kury) 158 6 Review of the Classification of the Orders of Arachnida Fig. 6.8 Biantes sp. (Biantidae, Singapore) (flickr, from A. Kury) Fam. Biantidae – Madagascar, Comoros, tropical Africa, Seychelles, Nepal, India, Burma, Bioko, South Africa, Sumatra, Malaya, Socotra, West Indies, South America (33 gen., 134 spp.) (Fig. 6.8) Fam. Escadabiidae – Brazil (six gen., eight spp.) Fam. Kimulidae (Minuidae) – South America (Brazil, Venezuela, Peru), Antilles (Cuba, Puerto Rico, Dominican Republic)(10 gen., 36 spp.) Fam. Samoidae – Cuba, Venezuela, Trinidad, Jamaica, Mexico, Haiti, Guatemala, Fiji, Seychelles, Australia (New South Wales), Samoa, Leeward Islands, Costa Rica, Cameroon, Panama, Mozambique, Indonesia (24 gen., 47 spp.) Fam. Stygnommatidae – Mexico, Belize, Ecuador, Jamaica, Puerto Rico, Colombia, Costa Rica, Panama, Venezuela, ? Malaya (1 gen., 33 spp.) (Fig. 6.9) Fam. Fissiphalliidae – Colombia (one gen., seven spp.) Fam. Guasiniidae – Venezuela, Brazil (two gen., three spp.) Fam. Icaleptidae – Colombia, Ecuador (two gen., two spp.) Fam. Zalmoxidae (Stygnoleptinae) – Mexico, Costa Rica, El Salvador, Brazil, Guyana, French Guiana, Bolivia, Peru, Venezuela, Argentina, Cuba, Panama, Jamaica, Belize, Trinidad, Ecuador, New Guinea, Solomon Islands, Fiji, Mauritius, Madagascar, Seychelles, Bismarck Archipelago, Philippines, Java, 6.9 Solifugae (Solpugida, Solifugi) 159 Fig. 6.9 Zygobunus sp. (Stygnommatidae, Costa Rica) (courtesy A.P. González, from A. Kury) Sulawesi, Moluccas, Australia, Marianas, New Caledonia, Marshall Islands, Caroline Islands, Polynesia (66 gen., 209 spp.) Fam. Cladonychiidae – Europe (Holoscotolemon Roewer), USA (17 spp.) Fam. Briggsidae (Pentanychidae, praeoccup.) – USA (Oregon, Washington)(six spp.) 6.9 Solifugae (Solpugida, Solifugi) Map 6.6 Distribution. of Solifugae (courtesy of W. Savary) 160 6 Review of the Classification of the Orders of Arachnida Fig. 6.10 The order counts 1113 spp. from 142 genera and 12 families (Prendini 2011, Blick and Kury 2011) Fam. Galeodidae – Africa (North Africa, Somalia), Asia (Turkey, Iran, Israel, Afghanistan, Central Asia, India, Yemen, Pakistan, Cyprus), Europe (Bulgaria, Greece, Macedonia, Ukraine, Russia) (9 genera, 201 species) (Fig. 6.10) 6.9 Solifugae (Solpugida, Solifugi) Map 6.7 Distribution of fam. Galeodidae 161 162 6 Review of the Classification of the Orders of Arachnida Fam. Karschiidae – Southeast Europe (Greece), Asia (Central Asia, Arabian Peninsula, Mongolia, Turkey, Iran), North Africa (4 gen., 40 spp.) Fam. Daesiidae – Europe (Spain, Greece), Central Asia, the Middle East, Africa, and South America (29 gen., 191 spp.) Map 6.8 Distribution of fam. Daesiidae 6.9 Solifugae (Solpugida, Solifugi) 163 Fam. Solpugidae – Africa, Asia (Iraq, Turkey, Saudi Arabia, etc.) (17 gen, 196 spp.) Map 6.9 Distribution of fam. Solpugidae 164 6 Review of the Classification of the Orders of Arachnida Fam. Ceromidae – Southern Africa (3 gen., 20 spp.) Map 6.10 Distribution of fam. Ceromidae 6.9 Solifugae (Solpugida, Solifugi) 165 Fam. Melanoblossiidae – Southeast Asia, Southern Africa (6 gen., 16 [31] spp.) Fam. Mummuciidae – South America (10 gen., 20 spp.) Map 6.11 Distribution of fam. Mummuciidae 166 6 Review of the Classification of the Orders of Arachnida Fam. Gylippidae – Central Asia, Turkey, the Near East, and Southern Africa (five genera, 27 species) Fam. Eremobatidae – USA, Mexico (8 genera, 192 species) Map 6.12 Distribution of fam. Gylippidae 6.9 Solifugae (Solpugida, Solifugi) Map 6.13 Distribution of fam. Eremobatidae 167 168 6 Review of the Classification of the Orders of Arachnida Fam. Ammotrechidae – USA, Mexico, Central and South America, West Indies (21 genera, 88 species) Map 6.14 Distribution of fam. Ammotrechidae 6.9 Solifugae (Solpugida, Solifugi) Fam. Hexisopodidae – Southern Africa (2 genera, 23 species) Map 6.15 Distribution of fam. Hexisopodidae 169 170 6 Review of the Classification of the Orders of Arachnida Fam. Rhagodidae – Northeastern Africa, Southwestern Asia, the Near East (27 genera, 99 species) Map 6.16 Distribution of fam. Rhagodidae 6.10 Araneae The order counts 42055 spp. (Platnick 2011) or 46464 spp. (World Spider Catalog, February 2017) or 47052 (end November 2017) or 47553 (WSC June 2018). The families of spiders used to be 114, but the efforts of some molecular specialists they became 117 (WCS, June 2018). The figures for the number of spider species change very rapidly for the different families. Maps 6.17 – Distributions of spiders Suborder Mesothelae – Southeast Asia, China, Japan (97 species) Fam. Liphistiidae – Southeast Asia, China, Japan (97 species) 6.10 Araneae Map from Xu et al. (2015) Map 6.17a Distribution of fam. Liphistiidae 171 172 6 Review of the Classification of the Orders of Arachnida Suborder Orthothelae Infraorder Mygalomorphae Fam. Actinopodidae – South America, Panama, Australia (48 spp.) Map 6.17b Distribution of Actinopodidae Fam. Antrodiaetidae – USA (33 spp.), Japan (two spp.) (35 spp.) Fam. Atypidae – Europe, Asia, Africa, USA, Mexico (52 spp.) Fam. Barychelidae – Australia, New Caledonia, Madagascar, Seychelles, Mauritius, tropical Africa, Southeast Asia, New Guinea, Sri Lanka, Socotra, Oceania, Fiji, South America, India, Mexico, Belize, Caribbean (296 spp.) Subfam. Barychelinae – Socotra, Australia, New Caledonia, Africa, Madagascar, Seychelles, Mauritius, South America Subfam. Sasoninae – Brazil, Venezuela, Australasia Subfam. Trichopelmatinae – Caribbean, Central to South America Fam. Ctenizidae – Mediterranean (Greece, Turkey,) Japan, Taiwan, China, Thailand, Central Asia, Southern Africa, USA, Mexico, Guatemala, Costa Rica, Australian region, New Guinea (135 spp.) Subfam. Ctenizinae – USA to Guatemala and Costa Rica, Thailand, China, Taiwan, Europe, Central Asia, Australian region, South Africa Subfam. Pachylomerinae – Australian region, America, Mediterranean, Japan, Taiwan, Costa Rica, USA Fam. Cyrtaucheniidae – Africa, Mediterranean, Australia, Central and South America, USA, Mexico, Turkmenistan, Tajikistan, Afghanistan, Thailand (107 spp.) 6.10 Araneae 173 Subfam. Aporoptychinae – Africa, Australia, Central and South America Subfam. Cyrtaucheniinae – Mediterranean Fam. Dipluridae – South America, Cuba, Africa, South Africa, Australia, Oceania, Southern USA to Costa Rica, Taiwan, Thailand, Central Asia, New Caledonia, Caribbean, India, Madagascar (188 spp.) Subfam. Diplurinae – South America, Cuba Subfam. Euagrinae – Africa, South Africa, Australia, Southern USA to Costa Rica, Taiwan, Central Asia, New Caledonia Subfam. Ischnothelinae – Mexico to Argentina, Peru, Caribbean, India, Africa, Madagascar Subfam. Masteriinae – Caribbean, Central and South America, Oceania, Australia Fam. Euctenizidae – USA, Mexico, Guadeloupe (76 spp., up to 2012 subfam. of Cyrtaucheniidae) Fam. Hexathelidae – New Zealand, Australia, Tasmania, Chile (Mediothele), North and Central Africa, Europe, Asia (Macrothele) (113 spp.) Subfam. Hexathelinae – Australia, New Zealand, Chile, Argentina Subfam. Macrothelinae – Africa, Europe, Asia, New Zealand Map 6.17c Distribution of fam. Hexathelidae Fam. Idiopidae – Australia, New Zealand, Central and South America, Madagascar, India, Sri Lanka, Seychelles, Southern Africa, Africa, South Asia, Middle East, Thailand, Canary Islands (325 spp.) Subfam. Arbanitinae – Australia, New Zealand Subfam. Genysinae – Central and South America, Madagascar, India, Sri Lanka, Seychelles 174 6 Review of the Classification of the Orders of Arachnida Subfam. Idiopinae – Southern Africa, India, Sri Lanka, South America, Africa, South Asia, Middle East, Canary Islands Fam. Mecicobothriidae – USA, Mexico, Brazil, Argentina, Uruguay (nine spp.) Map 6.17d Distribution of fam. Mecicobothriidae 6.10 Araneae 175 Fam. Microstigmatidae – Panama, Brazil, Argentina, Uruguay, Colombia, Venezuela, Ecuador, South Africa (17 spp.) Subfam. Micromygalinae – Panama (one spp.) Subfam. Microstigmatinae – Brazil, Argentina, Uruguay, Colombia, Venezuela, Ecuador, South Africa (15 spp.) Map 6.17e Distribution of fam. Microstigmatidae Fam. Migidae – Madagascar, tropical Africa, Chile, Argentina, Australia, New Zealand (97 spp.) Subfam. Calathotarsinae – Chile, Argentina, Australia Subfam. Miginae – Chile, New Zealand, Australia, Africa Subfam. Paramiginae – Madagascar, Africa 176 6 Review of the Classification of the Orders of Arachnida Fam. Nemesiidae – South America, USA, Mexico, Burma, India, China, Europe (Majorca, Spain, Portugal), Russia, Australia, New Zealand, Madagascar, South Africa, Ryukyu Islands, Southeast Asia (404 spp.) Fam. Paratropididae – St. Vincent, Panama, Peru, Brazil, Venezuela (11 spp.) Subfam. Glabropelmatinae – Panama, Venezuela Subfam. Paratropidinae – St. Vincent, Peru, Brazil, Venezuela Fam. Theraphosidae (incl. Metinae) – South and Central America, Caribbean, Australia, South and Southeast Asia, New Guinea, Africa, USA, Mexico, Asia Minor, Europe (962 spp.) Suborder Araneomorphae Fam. Agelenidae – Mediterranean, Europe to Central Asia, USA to Chile, New Zealand, Socotra, Himalaya, Cuba, Bahamas, Cyprus, China; Tegenaria, worldwide (1277 spp.) Fam. Amaurobiidae – Cosmopolitic (286 spp.) Fam. Ammoxenidae – Australia, Tasmania, Botswana, South Africa, Namibia, Zimbabwe (18 spp.) Fam. Amphinectidae – Western South America, Australia, Tasmania, New Zealand, introduced in the USA (159 spp.) Map 6.17f Distribution of fam. Amphinectidae 6.10 Araneae 177 Fam. Anapidae (incl. Micropholcommatinae and Holarchaeidae) – Central to South America, USA, Mexico, New Caledonia, Australia, Tasmania, New Zealand, New Guinea, Europe, China, Taiwan, Nepal, Korea, Japan, South Africa (223 spp.) Map 6.17g Distribution of subfam. Micropholcommatinae Fam. Anyphaenidae – Australia, New Zealand, India, Polynesia, Europe, Turkey, Kurile Island, China, USA, Canada, Mexico, Algeria, S. Africa, Central and South America, Caribbean, Galapagos (558 spp.) Fam. Araneidae (incl. Nephilinae – pantropical, 75 spp.) – worldwide (3122 spp.) Fam. Archaeidae – Australia, South Africa, Madagascar (71 spp.) Fam. Arkyidae – Australia (37 spp.) Fam. Austrochilidae – Chile, Argentina, Tasmania (10 spp.) Subfamily Austrochilinae Zapfe, 1955 – Chile, Argentina (nine spp.) Subfamily Hickmaniinae 1967 (as family Hickmaniidae) (Tasmania, cave) Fam. Caponiidae – USA, Mexico, Caribbean, Central and South America, South Africa (110 spp.) Fam. Chummidae – South Africa (two spp.) 178 6 Review of the Classification of the Orders of Arachnida Fam. Cithaeronidae – East Africa, India, Ivory Coast, Ethiopia, Greece, Libya to Malaysia, Australia, and Brazil (eight spp.) Fam. Clubionidae – Cosmopolitic (615 spp.) Fam. Corinnidae – Worldwide (778 spp.) Fam. Ctenidae – Cosmopolitic (514 spp.) Fam. Cyatholipidae – Africa, Madagascar, New Zealand, Australia, and one species (Pokennips dentipes) in Jamaica (58 spp.) Fam. Cybaeidae (incl. Argyronetidae) – America, Europe, Japan, Korea, China, Turkmenistan, Tajikistan, Uzbekistan, Sumatra, Venezuela, Colombia; Argyroneta, Palearctic (188 spp.) Fam. Cycloctenidae – New Zealand, Australia, Indonesia (Java) (36 spp.) Fam. Deinopidae – All continents, except Europe and Antarctica (61 spp.) Fam. Desidae – Australia, Tasmania, New Zealand, New Caledonia, Chile, USA, Oceania, Korea, Japan, India, Africa, Galapagos, Paraguay (175 spp.) Fam. Dictynidae – Europe, China, Mongolia, India, Algeria, Kazakhstan, Canary Islands, New Zealand, Australia, Sri Lanka, South Africa, USA, Canada, Mexico, Kyrgyzstan, West Indies, Hawaii, South America, St. Helena, Andaman Islands, Galapagos Island, New Caledonia, etc. (616 spp.) Fam. Diguetidae – USA, Mexico, Argentina, Brazil, Peru, Chile (15 spp.) Fam. Drymusidae – South Africa, Cuba, Brazil, Argentina, Costa Rica, St. Vincent, Hispaniola (16 spp.) Fam. Dysderidae – Southern Europe, Sardinia, Corsica, Crete, Syria, Azerbaijan, Turkmenistan, Israel, Iran, Algeria, Turkey; Dysdera, worldwide (542 spp.) Subfam. Dysderinae – Europe, Crete, Syria, Azerbaijan, Turkmenistan, Israel, Algeria, Turkey; Dysdera, worldwide Subfam. Rhodinae – Mediterranean Fam. Eresidae – Africa, Madagascar, Brazil, Eurasia (98 spp.) Subfam. Eresinae – Africa, Madagascar, Brazil, Eurasia Subfam. Penestominae – South Africa Fam. Eutichuridae – Holarctic, Africa, South America to Costa Rica, USA, Mexico, India, Lord Howe Island, Madagascar, Reunion, Samoa, Cape Verde, Thailand, Indonesia, St. Helena (344 spp.). Formerly (up to 2014) subfamily of Miturgidae, fam. status by Ramirez (2014) Fam. Filistatidae –Africa, Madagascar, USA, Mexico, Guatemala, Cuba, Chile, Brazil, Argentina, Colombia, Galapagos, Australia, New Guinea, Mediterranean to India, Iran, Israel, Tajikistan, Uzbekistan, etc. (152 spp.) Subfam. Filistatinae – Mediterranean to India, Iran, Israel, Tajikistan, Uzbekistan, Mexico Subfam. Prithinae – Africa, Madagascar, USA, Guatemala, Cuba, Chile, Brazil, Argentina, Colombia, Galapagos, New Guinea, etc. Fam. Gallieniellidae – Madagascar, Comoro Island, Kenya, South Africa, Australia, Argentina (55 spp.) 6.10 Araneae 179 Subfam. Gallienellinae – Madagascar, Comoro Island, Kenya, South Africa Subfam. Meedoinae – Australia (five gen.), Argentina (one gen.) Fam. Gnaphosidae – Worldwide (2200 spp.) Fam. Gradungulidae – New Zealand, Australia (A.C.T., Queensland, New South Wales, Victoria) (16 spp.) Fam. Hahniidae – Oceania, Australia, Tasmania, New Zealand, India, Philippines, Sumatra, Sri Lanka, Europe, Kazakhstan, Kyrgyzstan, Canada to Venezuela, Brazil, Mexico, USA, Alaska, etc. (252 spp.) Fam. Hersiliidae – Africa, Australasia, USA, Caribbean, South America, India, Sri Lanka, Spain, Portugal, Algeria, Central Asia, Australia, Borneo (181 spp.) Fam. Homalonychidae – USA, Mexico, ?India (? three spp.) Fam. Huttoniidae –New Zealand (one sp.) Fam. Hypochilidae – China, USA (12 spp.) Fam. Lamponidae – Australia, New Zealand, New Guinea, New Caledonia (genus Centrocalia) (192 spp.) Subfam. Centrothelinae – Australia, New Caledonia Subfam. Lamponinae – Australia, New Zealand, New Guinea Subfam. Pseudolamponinae – Australia Fam. Leptonetidae – USA, Mexico, Panama, Mediterranean, Portugal, Algeria, Japan, China (300 spp.) Fam. Linyphiidae – Worldwide (4547 spp.) Fam. Liocranidae – Cosmopolitic (272 spp.) Fam. Lycosidae – Cosmopolitic (2415 spp.) Fam. Malkaridae – Australia, Chile, Argentina; incl. Pararchaeidae from Australia, Tasmania, and New Zealand, 34 spp. (46 spp.) Map 6.17h Distribution of fam. Malkaridae 180 6 Review of the Classification of the Orders of Arachnida Fam. Mecysmaucheniidae – Chile, Argentina, Falkland Island, Juan Fernandez Islands, New Zealand (25 spp.) Subfam. Mecysmaucheniinae Simon, 1895 – Chile, Argentina, Falkland Island, Juan Fernandez Islands, New Zealand (22 spp.) Subfam. Zearchaeinae Forster et Platnick, 1984 – Chile, Argentina, New Zealand (three spp.) Fam. Megadictynidae – New Zealand (two spp., formerly in Nicodamidae, family status reestablished by Dimitrov et al. 2017) Map 6.17i Distribution of fam. Mecysmaucheniidae Fam. Mimetidae – Palearctic, Africa, Principe, Central and South America, USA, Asia, Australia; Ero and Mimetus, worldwide (152 spp.) Subfam. Gelaninae – Central and South America (18 spp.) Subfam. Melaenosiinae – Principe, India (two spp.) Subfam. Mimetinae – Worldwide Fam. Miturgidae – Thailand, Indonesia, South America to Costa Rica, Holarctic, Africa, Australia, St. Helena, New Zealand, New Guinea, Sicily, Israel, Pakistan (157 spp.) Subfam. Miturginae – Australia, New Guinea, Sicily, Israel, Africa, South America, USA to Argentina 6.10 Araneae 181 Subfam. Systariinae – USA to Mexico, Costa Rica, Brazil, Pakistan, Southeast Asia Subfam. Diaprograptinae – Timor, Australia, New Zealand Subfam. Zorinae – Central and South America, Australia, Tasmania, New Zealand, Israel, Guatemala; Zora, Palearctic (73 spp.) Fam. Mysmenidae – Europe, USA, Mexico, Cuba, Jamaica, Central and South America, tropical Africa, Algeria, South Africa, Galapagos, China, Nepal, India, Sri Lanka, New Guinea, Samoa, Hawaii, Philippines, Seychelles, Fiji, New Caledonia (137 spp.) Fam. Nesticidae – Europe, North America, Mexico, Japan, Jamaica, Cuba, Tibet, China, Laos, Philippines, Sri Lanka, Indonesia, Madagascar, Russia, Hawaii, Fiji, Nepal, New Guinea, Seychelles, South America, Turkey (278 spp.) Fam. Nicodamidae – Australia, New Guinea (27 spp.) Fam. Ochyroceratidae – South Asia, China, Seychelles, Angola, Cuba, Mexico to Peru, Pacific Islands, New Guinea (191 spp.) Fam. Oecobiidae – Mediterranean, Africa, South Africa, Asia, USA; Oecobius is cosmopolitan (111 spp.) Fam. Oonopidae – Central to South America, Mexico, USA to Argentina, Cuba, Antilles, Yemen, Seychelles, Bhutan, Nepal, China, Africa, Philippines, Southeast Asia, Australia, Tasmania, New Zealand, New Caledonia, Hawaii, Socotra, South Africa, Angola, East Africa, Europe, Sierra Leone, Congo, Algeria, Egypt, St. Helena, Sri Lanka (1777 spp.) Subfam. Gamasomorphinae – Cuba, Yemen, Seychelles, Bhutan, Nepal, China, Central to South America, Africa, Philippines, Southeast Asia, Australia, New Zealand, Mexico, USA to Argentina, Hawaii, Socotra, Seychelles, Antilles Subfam. Oonopinae – South Africa, Angola, East Africa, Europe, Panama to Argentina, Seychelles, Hawaii, Sierra Leone, Congo, New Caledonia, Mexico, Cuba, Tasmania, Algeria, Egypt, St. Helena, Sri Lanka Fam. Orsolobidae – South Africa, Australia, New Zealand, South Africa, Chile, Australia, Tasmania, Brazil, Chile, Argentina, Falkland Islands (188 spp.) Fam. Oxyopidae – Cosmopolitic (457 spp.) Fam. Palpimanidae – Africa, Socotra, South America, Cuba, South Asia, Uzbekistan, Mediterranean, Seychelles, Sri Lanka (144 spp.) Subfam. Chediminae – Africa, Socotra, South Asia, Seychelles, Sri Lanka Subfam. Otiothopinae – South America, Cuba, Africa Subfam. Palpimaninae – Mediterranean, Africa, South America, India, Uzbekistan 182 6 Review of the Classification of the Orders of Arachnida Fam. Penestomidae – South Africa, Lesotho (nine spp.) Fam. Periegopidae – Australia (Queensland), New Zealand (three spp.) Fam. Philodromidae – Cosmopolitic (539 spp.) Subfam. Pedinopisthinae – Hawaii Subfam. Philodrominae – Cosmopolitic Subfam. Pselloninae – India, New Guinea Fam. Pholcidae – Worldwide (c. 1611 spp.) Fam. Phrurolithidae – China, India, Indochina, Lord Howe Island, USA, Mexico, Canada, Europe, Madeira, Japan, North Africa, Madagascar, Seychelles, Central Asia, Russia, Taiwan, Philippines, tropical Africa (218 spp.) Fam. Physoglenidae – (72 spp.) Fam. Phyxelididae –Madagascar, Kenya, South Africa, Cyprus, Turkey, Sumatra, Borneo (64 spp.) Fam. Pimoidae – Europe, North America, Russia, China, Japan, Himalaya (40 spp.) Fam. Pisauridae – Cosmopolitic (335 spp.) Fam. Plectreuridae – USA, Mexico, Cuba (31 species) Fam. Prodidomidae – Africa, South America, Cuba, Australia, USA, Mexico, Panama, Galapagos, Bahamas, Japan, Spain, Algeria, Canary Islands (309 spp.) Fam. Psechridae – China, India, Nepal, Sri Lanka, South East Asia, Nicobar Islands, Philippines, Australia (61 spp.) Fam. Salticidae – Worldwide (5989 spp.) Fam. Scytodidae – Worldwide, except of the Far North (239 spp.) Fam. Segestriidae – America, Africa, Asia, Australia, New Zealand (124 spp.) Fam. Selenopidae – America, Asia, Africa, Mediterranean, Madagascar, Reunion, Comoro Islands (257 spp.) Fam. Senoculidae – South and Central America, Mexico (31 spp.) Fam. Sicariidae – Worldwide, except of the Far North (149 spp.) Fam. Sparassidae – Worldwide, except of the Far North (1215 spp.) Fam. Stenochilidae – Thailand, Malaysia, Singapore, Burma, Philippines, Bali, New Guinea, Fiji, Borneo, Vietnam, India (13 spp.) Fam. Stiphidiidae – Australia, Tasmania, New Zealand, New Guinea, Madagascar, Mauritius, New Caledonia (135 spp.) Fam. Symphytognathidae – in the tropics of Central and South America and the Australian region (with Oceania), with three species (Anapistula benoiti, A. caecula, Symphytognatha imbulunga) found in Africa and one (Anapistula ishikawai) in Japan. Anapistula jerai occurs in Southeast Asia (71 spp.) Fam. Synaphridae – Canary Island, Southern Europe, Egypt, Turkmenistan, Spain, three spp. on Madagascar (13 spp.) Fam. Synotaxidae – Australia, Tasmania, New Zealand, South America (11 spp.) Subfam. Pahorinae – endemic to New Zealand Subfam. Synotaxinae – Australia, South America (Chile, Brazil, Colombia, etc.) 6.10 Araneae 183 Map 6.17j Distribution of fam. Synotaxidae Fam. Telemidae – Sumatra, Malaysia, China, Vietnam, Japan, France, Spain, Tanzania, Cameroon, South Africa, Seychelles, New Caledonia, New Guinea, Guatemala, USA (Oregon, Alaska,) (69 spp.) Fam. Tetrablemmidae – Southeast Asia, Africa, Borneo, Central America, Brazil, Colombia, Panama, India, Indonesia (Sulawesi), Samoa, Madagascar, Sri Lanka, Caroline Islands, Seychelles (166 spp.) Fam. Tetragnathidae – Cosmopolitic (994 spp.) Fam. Theridiidae – Cosmopolitic (2400 spp.) Fam. Theridiosomatidae – Cosmopolitic (111 spp.) Fam. Thomisidae – Worldwide (2154 spp.) Fam. Titanoecidae – India, Sri Lanka to China, New Guinea, Marquesas Islands, Europe, Mexico, South America; Titanoeca, Holarctic (54 spp.) Fam. Trachelidae – Mexico, etc. (208 spp.) Fam. Trechaleidae – South and Central America, Mexico, USA, Japan (Shinobius Yaginuma) (120 spp.) Fam. Trochanteriidae – Australia, Chile, Sulawesi, Christmas Island, China, India, Korea, Japan, Africa, Madagascar, South America, New Caledonia (153 spp.) Fam. Trogloraptoridae – Caves in Oregon and California (one sp.) Fam. Udubidae – tropical Africa, Madagascar (2 end. genera) (12 spp.) Fam. Uloboridae – USA, Central and South America, Philippines, Samoa, Fiji, Vanuatu, New Guinea, Russia, Central Asia to Japan, Mediterranean to Azerbaijan, Oceania, Australia, New Zealand; Uloborus is spread worldwide, and Zosis is pantropical (283 spp.) Fam. Viridasiidae – Madagascar, Brazil (nine spp., until 2015 part of Ctenidae) 184 6 Review of the Classification of the Orders of Arachnida Fam. Zodariidae – South America, Mexico, Central America, Australia, New Zealand, New Guinea, Southeast Asia, Europe (1126 spp.) Fam. Zoropsidae (incl. Tengellidae and part of Zorocratidae) – Australia, New Zealand, Sri Lanka, China, Korea, Japan, Cyprus, Mediterranean, North and South Africa, Madagascar (178 spp.) Map 6.17k Distribution of subfam. Tengellidae Acari Traditionally, this numerous group has been considered as containing three orders: Opilioacarida (41 rec. spp., Beron 2014 and suppl.), Acariformes, and Parasitiformes (both containing more than 48 200 described species, Harvey 2002, actually more than 52300 spp. by 2017). The newest general paper on Acari (2009. Manual of Acarology, Third Edition) provides another subdivision. Following the recent subdivision of Parasitiformes in four orders (Lindquist et al. 2009), Holothyrida, Ixodida, and Mesostigmata are treated as separate orders. We prefer to treat here Opilioacarida as an order. How Many Are the Acari? For most groups of Acari, our knowledge is far from the existing taxa in the nature. A very instructive table has been drown by Walter and Proctor (1998). According to it (we are listing only the species, with minor corrections), less than one tenth of the minimal number of living Acari have been recorded and described so far. 6.11 185 Opilioacarida Group Opilioacarida Holothyrida Ixodida Mesostigmata Endeostigmata Oribatida Astigmata Trombidiformes Species 41 29 880 12,512 120 11,000 4500 17,050 Minimum (guess) 85 160 1000 97,520 1200 23,000 90,000 327,250 Maximum (guess) 170 320 1200 200,500 2400 110,000 180,000 637,500 Of course, the calculations of our colleagues are approximate, the real figures for the extant Acari species will be different, but even this approximation shows how far we are from the real picture of the number of species, let alone of the distribution of mites. For most countries there are no checklists and very few taxonomists describe Acari. We can see clearly that when in one country appear acarologists, the list of mites becomes much longer. Good example for this is the flood of new information from countries and regions like Turkey, Iran, Brazil, New Zealand, South Africa, Crimea, and others. If someday mites will be described using DNA, these figures will be different, if you judge from the recent multiplication of number of larger animals like Euscorpius (Scorpiones). 6.11 Opilioacarida The first to draw attention on the existence of such peculiar mites was C. With, describing the order, the only family, and the species Opilioacarus segmentatus (With 1903) and O. italicus (With 1904) – still the only two species known in Europe (southern parts of Italy and Greece). Shortly after him F. Silvestri (1905) discovered opilioacarids also in South America. Redikorzev (1937) found the first and only representative in Central Asia. Chamberlin and Mulaik (1942) not only described the northernmost American species – Neocarus texanus – but also separated it in a new genus, the second in the order. They also separated Opilioacarus hexophthalmus Redikorzev (1937) in a new genus Paracarus. Followed André (1947) and Naudo (1963) with descriptions of new species from tropical Africa and of the fourth genus in the order – Panchaetes. From 1966 to 1979, L. van der Hammen (1966, 1968, 1969, 1971, 1977) wrote important papers (one of them together with I. Coineau), adding to the seven species known prior to them another seven and the new genera Adenacarus, Phalangiacarus, and Salfacarus. Another three papers added new species to the system of Opilioacarida: JuvaraBalş and Baltac (1977) described two new species (the first from caves) from Cuba. More cave species have been described by Leclerc (1989) from Thailand, including 186 6 Review of the Classification of the Orders of Arachnida the new genera Siamacarus and Vanderhammenacarus. Lehtinen (1980) added one more species from Venezuela. Harvey (1996) made a cladogram of opilioacarid genera, suggesting that Paracarus and Siamacarus were the sister group to the other genera. Modern treatment of the order was enterprised by M. Vásquez and H. Klompen (2002, 2004, 2009, 2010). They described 13 more species from the Americas and Madagascar and the new genus Caribeacarus from Central and South America and left the initial genus Opilioacarus only for the two species O. segmentatus and O. italicus from Greece, Italy, and Algeria (more details about the distribution in Brignoli 1967, Beron 1990, and Thaler and Knoflach 2002). Neocarus Chamberlin et Mulaik became the most numerous genus in the order with 15 species. Harvey (2002) listed 9 genera and 20 species in Opilioacariformes, since 4 more genera and 17 new species have been described. Amazonacarus setosus and A. paraensis were described by Vásquez, Araújo, and Feres (2014) from Brazil; also from Brazil Bernardi et al. (2011, 2012, 2013) described more Opilioacarida. Das and Bastawade (2007) described Indiacarus pratyushi, new genus and species from India. Finally, two fossil species (Paracarus pristinus Dunlop, Wunderlich et Poinar, Jr. 2004 and ?Opilioacarus) have been described from Northern Europe (Baltic amber) and the order was mentioned (without more details) from Australia (Walter and Proctor 1998). There is still uncertainty concerning the rank and the place in the system of these strange Opiliones-like mites, inhabiting warmer countries. In the most recent treatment of Kranz et al. (2008), Walter and Harvey use the name Opilioacarida as an order. We are following this practice (Beron 2014). Ordo Opilioacarida (Opilioacariformes) – 13 gen., 41 rec. spp. Fam. Opilioacaridae Adenacarus Van der Hammen, 1966 – Yemen (one sp.) Amazonacarus Vásquez, Araújo et Feres, 2014 – Brazil (two spp.) Brasilacarus Vásquez, Araújo et Feres, 2015 – Brazil (one sp.) Caribeacarus Vásquez et Klompen, 2009 – Cuba, Dominican Republic, Panama (four spp.) Indiacarus Das et Bastawade, 2007 – India (one sp.) Neocarus Chamberlin et Mulaik, 1942 – USA, Mexico, Nicaragua, Venezuela, Cuba (14 spp.) Opilioacarus With, 1902 – Circum-Mediterranean (two spp. + one ? fossil, Baltic amber) Panchaetes Naudo, 1963 – Angola, Ivory Coast (two spp.) Paracarus Chamberlin et Mulaik, 1942 – One sp. (Kazakhstan) and one from Kaliningrad (fossil!) Phalangiacarus Coineau et Van der Hammen, 1979 – Gabon (one sp.) 6.11 Opilioacarida 187 Salfacarus Van der Hammen, 1977 – South Africa, Madagascar, Tanzania (nine spp.) Siamacarus Leclerc, 1989 – Thailand (two spp.) Vanderhammenacarus Leclerc, 1989 – Thailand (one sp.) Map 6.18 Opilioacarida in the Old World Adenacarus – Δ Indiacarus – ◄ Opilioacarus – ▲ Panchaetes – Paracarus – ♦ Phalangiacarus – ▼ Salfacarus – ■ Siamacarus – ► Vanderhammenacarus – 188 6 Review of the Classification of the Orders of Arachnida Map 6.19 Opilioacarida in the Americas Amazonacarus – ■ Brasilacarus – ▲ Caribeacarus – ▼ Neocarus – ◄ 6.12 Trombidiformes 189 Superorder Acariformes The mites of superorder Acariformes are not the favorite subject of zoogeographic analysis for various reasons. Part of the families includes only permanent parasites on other animals, which depend on the distribution of hosts and have no special zoogeography (mange mites, Myobiidae, Listrophoroidea, endoparasitic acari). Other families consist of adult and nymphal stages which are soil inhabiting predators and parasitic larvae (Erythraeidae, Trombiculidae, Trombidiidae s.l.). Many families consist entirely of free living forms (Anystidae, Caeculidae, Teneriffiidae, Bdellidae, numerous Oribatida). They could provide some information of zoogeographic interest (Atyeo 1960; Beron 2016; Colloff and Halliday 1998; Kepka 1968; Robaux 1969; Strandtmann 1974; Travé 1976; Wallwork 1969, 1973, 1979; Hammer and Wallwork 1979; Vu Kuang Manh 2015). As we published a world catalogue of Erythraeoidea and Calyptostomatoidea (Beron 2008a), we shall try to analyze some peculiarities of the distribution of these mites. We shall analyze also the distribution of some free living mites. 6.12 Trombidiformes Prostigmata Several families are analyzed as examples. Erythraeoidea The world catalogue of Erythraeoidea (Beron 2008a) and the supplements (Beron 2017) show that, by October 2017, there are 918 spp. of the families Smarididae (10 genera) and Erythraeidae (57 genera). Some particularities in the distribution of the genera could be the knowledge of these (and many other) mites is very incomplete and depends largely on the geography of specialists (Southcott and others). Also, with Smarididae and Erythraeidae, there are two classifications (of adults and larvae), and they could change with the rearings of larvae. For the time being, some genera (Smaris, Hirstiosoma) seem distributed worldwide (more or less cosmopolitic). Other genera seem more confined to one area (Sphaerotarsus, Surasmaris). Interesting was to find the American genus Trichosmaris in Papua New Guinea, on top of Mt. Fugilil, 3100 m. No mistake could be supposed as the mite was found and described by the present author and it is a native species found in the center of the island, far from the sea or human settlements. Fam. Smarididae Calorema Southcott, 1962 – Panama, Mexico Fessonia von Heyden, 1826 – India, Australia, Burma, Mexico, USA, Nigeria, Europe, Guatemala Kraussiana Southcott, 1961 – South Africa, Malaysia, Caroline Islands Neophanolophus China, 1976 – Malaysia, India Pilosoma Southcott, 1961 – France (= Fessonia) 190 6 Review of the Classification of the Orders of Arachnida Smaris Latreille, 1796 – Australia, Mexico, Panama, Chile, Europe, Cyprus, ? South Africa Surasmaris Southcott, 1995 – Costa Rica Clavismaris Southcott, 1963 – Costa Rica, Texas Clipeosoma Southcott, 1948 – Malaysia, Europe, New Guinea Hirstiosoma Womersley, 1934 – Europe, Mexico, South America, Australia, Tasmania, New Zealand Sphaerotarsus Womersley, 1936 – Australia Trichosmaris Southcott, 1963 – New Guinea, Guatemala, Mexico, USA Fam. Erythraeidae Augustsonella Southcott, 1961 – Mexico, USA Claverythraeus Trägårdh, 1937 – China Collemboerythraeus Noei, Saboori et Hakimitabar, 2017 – Iran Curteria Southcott, 1961 – Austria, Switzerland, Greece, Germany, Holland, Poland, Canary Island, Algeria, Eatoniana Cambridge, 1898 (= Abalakeus Southcott (1994), China, Niger, Iran, Spain) – Algeria, Egypt, Greece, Jersey, Spain, Switzerland, Armenia, Transcaspia, Tunisia, South Africa, ?Argentina Erythraeus Latreille, 1806 – Europe, Iceland, Israel, Iran, Turkey, Japan, China, Taiwan, India, Cambodia, Galapagos, Bolivia, Peru, Egypt, Israel, South Africa, Tanzania, Sudan, Ethiopia, USA, Australia Erythraxus Southcott, 1961 – Mexico Erythrellus Southcott, 1946 – Australia Erythrites Southcott, 1961 – Australia, New Zealand Erythroides Southcott, 1946 – Australia Forania Southcott, 1961 – France, Helladerythraeus Beron, 1988 – Greece Kakamasia Lawrence, 1944 – South Africa Lasioerythraeus Welbourn et Young, 1987 – Dominican Republic, USA, Bolivia Makolia Saboori, Khaustov et Hakimitabar, 2009 – Crimea Kamertonia Gabrys, 1999 – Poland Neosmaris Hirst, 1926 – New Zealand Opserythraeus Fain, 1996 – Rwanda Paraphanolophus Smiley, 1968 – Mexico, Belize Podosmaridia Trägårdh, 1937 – China Rainbowia Southcott, 1961 – Australia Ramsayella Zhang, 2000 – New Zealand Taranakia Southcott, 1988 – New Zealand Tepotztlana Hoffmann et Mendes, 1973 – Mexico Leptus Latreille, 1796 – Cosmopolitic Caeculisoma Berlese, 1888 – Congo, Nigeria, China, Bhutan, Sumatra, Australia, South Africa, New Guinea, New Ireland, Marquesas Islands, New Zealand, Brazil, Argentina, Paraguay Callidosoma Womersley, 1936 – USA, Panama, Guatemala, Brazil, Argentina, Australia, Taiwan, New Guinea, New Zealand, Dominican Republic 6.12 Trombidiformes 191 Carastrum Southcott, 1988 – Australia, Indonesia (Bali) Dambullaeus Haitlinger, 2001 – Sri Lanka Harpagella Southcott, 1996 – Australia Momorangia Southcott, 1972 – New Zealand, Kenya, Brazil Abrolophus Berlese, 1891 – Europe, Madagascar, China, Vietnam, Iran, Indonesia, East Africa, USA, Ghana, Ethiopia, South Africa, India, Canary Island, New Guinea, Australia, New Zealand, Japan, ?Argentina = Hauptmannia Oudemans, 1910 Andrevella Southcott, 1961 – France, Iran Cecidopus Karsch, 1879 – South Africa, Nigeria, Sri Lanka, Pakistan, India, Congo Charletonia Oudemans, 1910 – Madagascar, tropical Africa, South Africa, Egypt, Europe, Iran, India, Sri Lanka, Indonesia, Malaya, Vietnam, Cambodia, China, Japan, Australia, Tasmania, Canada, USA, Brazil, Canary Island Grandjeanella Southcott, 1961 – Balearic Island, South Europe, Iran, Australia, Iguatonia Haitlinger, 2004 – Brazil Neoabrolophus Khot, 1965 – Tanzania, India, Malaysia Pussardia Southcott, 1961 – Australia Rudaemannia Haitlinger, 2000 – Europe, Turkey (= Balaustium) Phanolophus Andre, 1927 – Algeria, Europe, Cyprus, East Africa Balaustium von Heyden, 1826 – Cosmopolitic Bursaustium Haitlinger, 2000 – Turkey, Iran Fozustium Haitlinger, 2005 – Brazil Italustium Haitlinger, 2000 – Italy Lomeustium Haitlinger, 2006 – West Africa Madinahustium Kamran et Alatawi, 2016 – Saudi Arabia Moldoustium Haitlinger, 2007 – Moldova, Ukraine Microsmarialla Khot, 1963 – India Microsmaris Hirst, 1926 – Australia, New Zealand, India Mypongia Southcott, 1961 – Australia Neobalaustium Willmann, 1951 – Romania, Bulgaria Pollux Southcott, 1961 – Australia, India Wartookia Southcott, 1961 – Australia Myrmicotrombium Womersley, 1934 – Australia, Greece (subg. Graecotrombium) The known distribution is rather incomplete. When in one country there are specialists (Australia, Poland, Iran), the knowledge on the fauna is much closer to the actual situation. Conclusion: In the Erythraeoidea, there are no families or subfamilies endemic to a particular continent or zoogeographical region. Endemism is developed only on genus and species level. Calyptostomatoidea Fam. Calyptostomatidae Calyptostoma Cambridge, 1875 – Europe, Asia, Africa, New Guinea, Australia These unmistakable mites are obviously missing in the Americas. 192 6 Review of the Classification of the Orders of Arachnida Superfam. Anystoidea Family Teneriffiidae Austroteneriffia Womersley, 1935 – Australia, Tajikistan, Iran, Japan, Yemen (incl. Socotra) Himalteneriffia Schmölzer, 2002 – India (Ladakh) Mesoteneriffia Irk, 1939 – Alps (Austria, Switzerland) Mesoteneriffiola Schmölzer, 1956 – France Neoteneriffiola Hirst, 1924 – Egypt, Namibia, China, USA, Mexico Parateneriffia Thor, 1911 – Paraguay Sinoteneriffia Yin, Bei et Li, 1995 – China Teneriffia Thor, 1911 – Canary Island, Japan, Malaysia, Mexico The family is spread on all continents, endemism only on genus level. Fam. Anystidae Anystis von Heyden, 1826 Autenriethia Oudemans, 1936 Barellea Oudemans, 1936 Mesoanystis Zacharda, 1985 (fossil) Scharfenbergia Oudemans, 1936 Snartia Oudemans, 1936 Tencateia Oudemans, 1936 Walzia Oudemans, 1936 Chaussieria Oudemans, 1937 Erythracarus Berlese, 1903 Erythrocheylus Berlese, 1903 Namadia Meyer et Ueckermann, 1987 Paratarsotomus Kuznetsov, 1983 Siblyia Oudemans, 1936 Tarsolarcus Thor, 1912 Tarsotomus Berlese, 1882 Family Pseudocheylidae Anoplocheylus Berlese, 1910 – Egypt, Iran, Italy, South Africa, Australia, Brazil, Turkey, Crimea Neocheylus Trägårdh, 1906 – Australia, South Africa, Uganda Pseudocheylus Berlese, 1888 – USA, Brazil, Paraguay The family is spread on all continents, endemism only on genus level. Family Stigmocheylidae Stigmocheylus Berlese, 1910 – USA, Italy, Egypt Superfamily Caeculoidea Family Caeculidae Allocaeculus Franz, 1952 – France, Kazakhstan, Russia, Spain, Austria, Croatia, Czech Republic, Macedonia, Namibia, South Africa, Morocco, Algeria, 6.13 Sarcoptiformes 193 Tunisia, Pakistan, Kenya, Nigeria, Chad, Mozambique, Sudan, Senegal, Canary Island Andocaeculus Coineau, 1974 – Argentina Caeculus Dufour, 1832 – USA, Mexico, Puerto Rico, Senegal, Europe Calocaeculus Coineau, 1974 – South Africa Microcaeculus Franz, 1952 – Austria, Germany, Czech Republic, Italy, France, Corsica, Greece, Portugal, Spain, Canary Island, Salvages Island, South Africa, Argentina, Chile, Morocco, Algeria, Namibia, China, Korea, Australia Neocaeculus Coineau, 1967 – Australia, Tasmania, Lord Howe Island, New Zealand, Namibia Procaeculus Jacot, 1936 – Trinidad, Dominican Republic, Hawaiian Island, Poland (fossil), Mexico, USA, Guatemala The family is spread on all continents, endemism only on genus level. Adamystoidea Family Adamystidae Adamystis Cunliffe, 1957 – USA, Iran, Pakistan, Tajikistan, Uzbekistan, Thailand, South Africa, France, Ukraine Bovidromus Coineau, Theron et Fernandez, 2006 – South Africa Rhinodromus Coineau, Theron et Fernandez, 2006 – South Africa Saxidromus Coineau, 1974 – Mexico, Cuba, South Africa, France So far the family is known from Europe, Asia, Mexico, Cuba, and South Africa, but these mites are rarely studied and could be found in other places as well. The same conclusion is valid also for Pomerantziidae and other small families. Pomerantzioidea Family Pomerantziidae Apomerantzia Quing-Hai Fan et Yan Chen, 2005 – USA Pomerantzia Baker, 1949 – USA, China, Philippines 6.13 Sarcoptiformes Acaridida Although the parasitic mites of this suborder follow the distribution of their hosts, it may be useful to mention a few details of the distribution even of permanent parasites as Listrophoroidea or Psoroptoidea. However, it may be noticed that the large family Atopomelidae is almost absent in Europe and very numerous in the tropical countries (Beron 2011). Oribatida The huge group (suborder or part of suborder) of Oribatida is composed by ca. 6600 spp. of ca.1200 genera and more than 200 families. Several papers outline some of the zoogeographical particularities of Oribatida: Wallwork (1969, 1973, 1979), Hammer and Wallwork (1979), Travé (1976a, b), Schatz (1998), Niedbala (2002, 2004), and others’ interest. 194 6.14 6 Review of the Classification of the Orders of Arachnida Holothyrida These big, conspicuous, and poisonous mites were first noticed by Gervais (1842) on Mauritius Island. He described Holothyrus coccinella, raised as family by Thorell (1882), who started the long series of descriptions of New Guinean Holothyrida (followed by Canestrini 1897). Another early Arachnologist, Butler (1876), recorded these strange mites from Rodrigues. Thon (1905, 1906) and Warburton (1913) described three more species from the Seychelles. The series of early descriptions includes also holothyrids from New Caledonia (Berlese 1923) and New Zealand (Womersley 1935). So far, all findings came from islands. Finally, Thon (1909) raised Holothyrida in a suborder. Van der Hammen (1961, 1965, 1968, 1972, 1983) started the modern stage of the study of Holothyrida. He described (1972) the second family in the suborder – Allothyridae – for the two species from Australia and New Zealand, belonging to the new genera Allothyrus van der Hammen, 1961 and Australothyrus der Hammen, 1983. From 1981 to 1999 P. Lehtinen contributed greatly to the study of Holothyrida by describing a new family Neothyridae (1981) from South America with the two new genera Neothyrus and Diplothyrus (Kontschán and Mahunka 2004 added another new genus Carabothyrus from the Dominican Republic). Lehtinen (1981, 1995, 1999) described also the new genera Hammenius, Haplothyrus, Indothyrus, Lindothyrus, and Sternothyrus (from New Caledonia, Lord Howe Island, the Seychelles, New Guinea, and Sri Lanka) and raised Leiothyrus and Thonius to genera, thus making his own zoogeographical analysis of 1991 out of date. He became the author of 9 out of the 15 genera in the family. Gerlach, Lehtinen, and Madl (2010) added three endemic genera from the Seychelles. Holothyrida (Holothyrina, Holothyroidea, Tetrastigmata) have been considered by the modern (after 1949) authors either as suborder of Parasitiformes or as order. It is considered as order by Walter in Kranz and Walter (2009), and we follow this concept here and in our Catalogue (Beron 2014). Order Holothyrida – Three fam., 13 gen., 29 spp. Fam. Allothyridae – Two gen., three spp. Allothyrus van der Hammen, 1961 – Australia, New Zealand (two spp.) Australothyrus van der Hammen, 1983 – Australia (one sp.) Fam. Holothyridae – 8 gen., 20 spp. Dicrognatus Gerlach, Lehtinen et Madl, 2010 – Seychelles (two spp.) Hammenius Lehtinen, 1981 – New Guinea (four spp.) Haplothyrus Lehtinen, 1995 – New Caledonia (two spp.) Holothyrus Gervais, 1842 – Mauritius (two spp.) Indothyrus Lehtinen, 1995 – Sri Lanka (one sp.) Leiothyrus van der Hammen, 1983 – New Guinea (three spp.) Lindothyrus Lehtinen, 1995 – New Caledonia, Lord Howe Island (two sp.) 6.14 Holothyrida Michaelothyrus Gerlach, Lehtinen et Madl, 2010 – Seychelles (one sp.) Sternothyrus Lehtinen, 1995 – Seychelles (one sp.) Thonius Lehtinen, 1981 – New Guinea (four spp.) Fam. Neothyridae – Three gen., three spp. Caribothyrus Kontschán et Mahunka, 2004 – Dominican Republic (one sp.) Diplothyrus Lehtinen, 1999 – Brazil, French Guyana (two spp.) Neothyrus Lehtinen, 1981 – Peru, Venezuela (one sp.) Incertae sedes – Rodrigues (one sp.) Map 6.20 Holothyrida in the Old World (Indian Ocean) Dicrognatus – ■ Holothyrus – ◘ Indothyrus – ♦ Michaelothyrus – ◄ Sternothyrus – ♣ 195 196 6 Map 6.21 Holothyrida in Australasia Allothyrus – X Australothyrus – Ω Hammenius – ● Haplothyrus – Leiothyrus – Lindothyrus – ◇ Thonius – Review of the Classification of the Orders of Arachnida 6.14 Holothyrida 197 Map 6.22 Holothyrida in Americas Caribothyrus – ▲ Diplothyrus – ▼ Neothyrus – ► The strange poisonous Holothyrida are 29 valid species from 15 genera and 3 families (Beron 2014), and we think that the guesstimate of 320 existing species and even the minimum of 160 (Walter and Proctor 2009) are exaggerations. Holothyrids are large and conspicuous creatures, and it is difficult to miss them where they exist. It is true that 17 of these 29 species have been described after 1980 (by P. Lehtinen, L. van der Hammen, J. Kontschán and S. Mahunka, and H. Klompen). A total restructuring has been done by these authors and by J. Gerlach also in the generic classification of the order (from one genus Holothyrus to 15 genera). Now is more 198 6 Review of the Classification of the Orders of Arachnida credible to analyze the most intriguing distribution of this order, more credible even in comparison with the relatively recent zoogeographical analysis of Lehtinen (1991). Many of the holothyrids are endemic genera, and not because they have not been looked for elsewhere, but it seems that on their islands they might be really endemic. The family Neothyridae is entirely American – four species from Dominican Republic (Caribothyrus Kontschán et Mahunka), French Guyana, Brazil (Diplothyrus Lehtinen), and Peru and Venezuela (Neothyrus Lehtinen). From Australia have been described by Van der Hammen a special family (Allothyridae) and two new genera Australothyrus and Allothyrus. The last one has been recorded also from New Zealand (Allothyrus australasiae Womersley, 1935), the same species described from Australia, but such important finding has not been confirmed since. What concerns the Old World is that there live the remaining 22–23 species of 10 genera. The heavy weight of the whole order seems to be in New Guinea with 11 species of the genera Hammenius Lehtinen, Leiothyrus Van der Hammen, and Thonius Lehtinen. They all belong to the family Holothyridae, spread from Mauritius to New Caledonia, and not to the “Australian” Allothyridae. Endemic genus and species for Sri Lanka is Indothyrus greeni Lehtinen, 1995. New Caledonia and Lord Howe Island, being between Australia and New Zealand, share another genus of Holothyridae – Lindothyrus Lehtinen, 1995. Another (monotypic) genus (Haplothyrus Lehtinen, 1995) is endemic to New Caledonia. Particularly interesting is the presence of Holothyridae on the islands close to Africa, but not in Africa itself or Madagascar. On the Seychelles live at least four species of three endemic genera: Dicrognathus Gerlach, Lehtinen et Mandl, 2010; Michaelothyrus Gerlach, Lehtinen et Mandl, 2010; and Sternothyrus Lehtinen, 1995. They are part of the many endemisms of the granitic islands. Finally, Mauritius, the place from where has been described the first Holothyrid as early as in 1842, remains with two species of the first described genus Holothyrus Gervais (plus one uncertain species from Rodrigues). The distribution of the holothyrids over so many islands far from each other and their (apparent) absence from places like Africa, Madagascar, or India remain among the riddles of zoogeography. 6.15 Ixodida Fam. Nuttaliellidae Nuttalliella Bedford, 1931 (one sp.) This family is (the only described species Nuttalliella namaqua Bedford) endemic for a narrow area in Southern Africa, from Tanzania to Namibia. Fam. Ixodidae The family Ixodidae contains ca. 700 spp. in 14 genera (Filippova 1994, Horak et al. 2002, Kolonin 1978 1983 1984, 2009, Guglielmone et al. 2010), parasites on Bibliography 199 vertebrates all over the world. Kolonin (1981, 1983) subdivided the world into superregions and regions after the tick fauna in them. Fam. Argasidae Also parasites of vertebrates (193 spp. in 5 genera). 6.16 Mesostigmata Many groups are mono- or oligoxenous parasites and their distribution follows the distribution of their hosts (Beron 2014). Some examples are as follows: Fam. Spelaeorhynchidae – the only genus Speleorhynchus and seven species are found in South and Central America and the Antilleans. They follow the distribution of the bats of families Phylostomatidae and Mormoopidae. Fam. 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Chopard et A. Villiers). Scorpions, Pseudoscorpions et Solifuges. Mémoires de l’Institut Français d’Afrique Noire. Série A. Sci Natur 10:93–107 Vachon M (1969) L’évolution du concept d’Arachnide. Bull Mus Nat Hist Nat Paris 41(suppl. I):184–186 Bibliography 317 van der Hammen L (1977) A new classification of Chelicerata. Zoologische Mededelingen, Leiden 51:307–319 van der Hammen L (1989) An introduction to comparative arachnology. SPB Academic Publishing, The Hague Vasquez JM (1995) Los aracnidos de Mexico. Parte 1: Ricinulei, Amblypygi, Solifugae, Palpigradi, Schizomida, Uropygi. Dugesiana 2:15–37 Vázquez RI (1996) Palpigragi, Schizomida, Uropygi, Amblypygi, Solifugae y Ricinulei. In: Llorente Bousquets AN, García-Aldrete JE, Víquez C, Longhorn S (2016) Arachnids of Central America, pp 59–82. Online at http://arachnids.myspecies.info Werner F (1925) Skorpione und Solifugen aus dem oestlichen und nordlichen Spanien. 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Arachnida: Scorpiones, Pedipalpi and Solifugae. In: Expedition to South – West Arabia 1937–8, I, 1–8:43–49 Yeates DK, Harvey MS, Austin AD (2003) New estimates for terrestrial arthropod species-richness in Australia. Rec S Aust Mus Monog Ser 7:231–241 Yin CM, Kim JP (1997) The current status and trends in the research of Chinese arachnology. Kor Arachnol 13(2):51–80 Yoshikura M (1975) Comparative embryology and phylogeny of Arachnida. Kumamoto J Sci Biol 12:71–142 Chapter 7 Regional Arachnogeography Abstract The arachnofauna of various parts of the Earth is analyzed and the particularities, endemics, relicts, and the presumed ways of formation of the fauna are outlined. Also the northern limits of the groups in the Holarctic are indicated, and the connections in the geological time are analyzed. Bibliographies Smaller orders – Harvey (2003) Scorpiones – Dupré (1998, 2013a, 1913b), Fet et al. (2000) Pseudoscorpiones – Schawaller (1980), Harvey (1990), Zaragoza (2000) Opiliones – Giribet (2000), Cokendolpher and Lee (1993), Kury (2011), Schönhofer (2013) Palpigradi – Condé (1996), Harvey (2003) Amblypygi – Harvey (2003) Uropygi (Thelyphonida) – Harvey (2003), Rowland and Cooke (1973) Schizomida – Harvey (2003), Reddell and Cokendolpher (1995) Ricinulei – Harvey (2003) Solifugae – Harvey (2003), Gromov and Savary (on line) Opilioacarida – Beron (2014) Araneae – Roewer (1940), Brignoli (1983), Platnick (1989, 1993, 2014) Acariformes – Beron (2008, 2011, 2015a, 2015b, 2016b) Parasitiformes – Beron (2014, 2016a) 7.1 7.1.1 Holarctic Zoogeography and Arachnozoogeography From 16 recent orders and 10 suborders of Arachnida, 15 orders and almost all suborders are recorded in the huge Holarctic Kingdom. Lacking from the arachnofauna of the Holarctic are only Holothyrida. Some others are recorded only on the edge of © Springer International Publishing AG, part of Springer Nature 2018 P. Beron, Zoogeography of Arachnida, Monographiae Biologicae 94, https://doi.org/10.1007/978-3-319-74418-6_7 319 320 7 Regional Arachnogeography the Kingdom – Schizomida and Ricinulei in the Nearctic part, Thelyphonida (Uropygi) in the Far East of Russia (?), and Texas in the USA. Holarctic is usually regarded as Kingdom composed of two regions (Palearctic and Nearctic). In the scheme of Krzhanovskiy (2002), four regions are included in the Holarctic Kingdom (Boreal Region, Region of the Ancient Mediterranean, East Asian Region, Sonoran Region). He does not use the widely accepted terms Palearctic and Nearctic. 7.2 7.2.1 Europe (General) Geography, General Zoogeography, and Paleogeography The European Environment Agency (2002) (EEA) subdivides Europe into six geographical regions: North Europe, West Europe, Middle Europe, South Europe, Southeast Europe, and East Europe. Caucasus is a border area between East Europe and Asia Minor. The Agency subdivides Europe also into nine biogeographical regions: Arctic, Boreal, Continental, Mediterranean, Alpine, Atlantic, Pannonic, Steppic, and Black Sea regions. To them are attached two more regions: Macaronesian (Azores, Madeira, Ilhas Selvagens, Canary Islands) and Anatolian (Asia Minor). Arctic Region Island, Svalbard (Spitzbergen), Jan Mayen, the Far North of Norway and Kola Peninsula, Arkhangelskaya Oblast, Novaya Zemlya, Franz Josef Land in Russia. Tundra, Boreal forest, cold desert. Several endemic spiders and mites, no endemic genera. Boreal Region Most of the Scandinavian Peninsula, the Baltic states, North of Russia (2,900,000 km2, of which 1,900,000 in Russia). Most of the area is forest (58%) and cultivated lands. Continental Region From France to Central Russia. Most is cultivated land (52%), forest (27%), and meadows (14%). Mediterranean Region South of France; the Iberian, Apennine, and Balkan Peninsula; and the Mediterranean Islands. Mountainous area with many caves, rich in endemic Arachnida and also endemic members of orders not known in the northern regions (Solifugae, Opilioacarida, Scorpiones). The area has 36% of cultivated land, 25% of forests, 29% of meadows, etc. 7.2 Europe (General) 321 Alpine Region According to this subdivision, it includes not only the Alps but also the Scandinavian Mountains, the Pyrenees, the Carpathians, the Dinaric Mountains, the “Rhodopes”(certainly including Rila and Pirin), the Ural Mountains, and the Caucasus. From this area 41% are forests, 26% meadows, 8% cultivated land, and 13% with few or no vegetation. In many endemic arachnids, particularly remarkable is the scorpion Belisarius xambeui. Atlantic Region British Islands, Hebrides, Orkney and Shetland Islands, Faroe Islands, Southeast of Norway, northern part of Iberian Peninsula, and the area from the North of France to Denmark. The area consists of 38% cultivated lands, 30% meadows, 13% forests, etc. Pannonic Region Mostly the Hungarian and the other plains of Central Europe (in Serbia, Romania, Slovakia, etc.). Some typical steppe Arachnida. Steppic Region Dobrogea, south Moldova, Ukraine, European Russia, European part of Kazakhstan. From this area 75% is cultivated land, 17% meadows, 2% forest, etc. The Black Sea Region The western, southern, and eastern coast areas of Black Sea and Central Georgia. The meadows are 34%, forest 29%, cultivated land 14%, etc. These areas are defined by the European Commission and the Council of Europe for evaluation and reporting of nature conservation. The real zoogeographical subdivision of Europe is different. Usually it is divided into Eurosiberian and Mediterranean provinces. From the map of Udvardy (1975), we can see that the biogeographical subdivision is much more detailed. Map 7.1 West Palearctic Biogeographical Realm (Udvardy 1975) Provinces (on this map): 3 – West Eurasian Taiga 5 – Icelandian 6 – Subarctic Birchwoods 8 – British Islands 9 – Atlantic 10 – Boronemoral 11 – Middle European Forest 12 – Pannonian 13 – West Anatolian 16 – Iberian Highlands 17 – Mediterranean Sclerophyll 18 – Sahara 19 – Arabian Desert 20 – Anatolian-Iranian Desert 21 – Turanian 24 – Iranian Desert 26 – High Arctic Tundra 27 – Low Arctic Tundra 28 – Atlas Steppe 29 – Pontian Steppe 31 – Scottish Highlands 32 – Central European Highlands 33 – Balkan Highlands 34 – Caucasus-Iranian Highlands 7.2 Europe (General) 7.2.2 323 Arachnogeography Ref.: Aakra and Houge (2000), Agnarsson (1996), Armas and Rehfeldt (2015), Beier (1963), Beron (2011), Blagoev (2002), Blick et al. (2004), Blick and Christian (2004), Blick et al. (2004), Blick and Komposch (2004), Blick et al. (2004), Bosmans (2009), Bosmans and Chatzaki (2005), Bosmans and De Keer (1985), Bosmans and Vanuytven (2001), Bosmans et al. (2013), Bosmans et al. (2013), Buchar et al. (1995), Buchar and Ružička (1995), Cardoso (2000, 2010), Cardoso and Morano (2010), Cawley (2002), Chemini (1995), Christophoryová et al. (2012), Condé (1996), Ćurčić (1974), Ćurčić et al. (2004), Defosse (2004), Deltshev (2005), Deltshev and Blagoev (2001), Deltshev et al. (2003), Drensky (1936), Farzalieva and Esyunin (2000), Feider (1965), Fet (2010), Gaidus et al. (1999), Gencheva and Georgieva (2013), Gardini (1994), Hadži (1973), Klimeš (2000), Komposch (2004), Komposch and Gruber (2005), Koponen (2005), Kronestedt (2001), Kury and Mendes (2007), Le Peru (2007 2011), Loksa (1969), Mahnert (2004), Marinu and Verneau (2002), Martens (1978), Meinertz (1962), Merrett et al. (1985), Mheidze (1964), Milošević (2002), Morano and Cardoso (2009), Nentwig et al. (2013), Nikolić and Polenec (1981), Novak (2004, 2005), Novak et al. (2006), Otto (2014), Ozimec (2000), Pack-Beresford (1926), Pantini et al. (2013), Pantini and Isaia (2015), Pesarini (1994), Petrov (1997), Prieto (2003, 2008), Proszynski and Starega (1971), Rafalski (1960, 1961, 1967), Rafalski and Starega (1997), Rambla (1967), Roberts (1985), Sanu and Szinetar (1994), Sankey and Savory (1974), Scharff and Gudik – Sørensen (2011), Schmölzer (1995), Simon (1875), Staręga (1976a, 1976b, 1978, 2000), Starkoff (1958), Stoch (2003), Stol (1993 2007), Szalay (1968), Šilhavý (1956), Telnovs (2000a, 2000b), Topcu et al. (2005), Tumšs (1963), Vanhercke (1999), Varol (2003), Vilkas (1992), Weiss and Urak (2000), Wiehle (1953), Wijnhoven (2005), Wintarska (2008), Zaragoza (2007). In present-day Europe are lacking some “southern” groups – Ricinulei, Holothyrida, Thelyphonida (Uropygi), Amblypygi, and Schizomida. Some other warm loving groups are represented only on the southernmost Mediterranean part of the continent and with small number of taxa – Scorpiones, Solifugae, Opilioacarida, Opiliones, Cyphophthalmi, and Laniatores. Well represented are the spiders, pseudoscorpions, opilions (Eupnoi and Dyspnoi), and most groups of Acari, Palpigradi. Across Europe pass the lines marking the northern limit of many groups. Palpigradi In Europe (including Madeira) are registered 28 species of Palpigradi, all belonging to one genus – Eukoenenia Börner (fam. Eukoeneniidae). They are recorded (without counting the subspecies) from France (9), Italy (12), Greece (3), Bulgaria (1), Hungary (1), Austria (2), Bosnia and Herzegovina (1), Croatia (2), Malta (2), Portugal (2), Romania (4), Slovakia (1), Slovenia (1), and Spain (4) – 14 countries (Blick and Christian 2004, Condé 1996). Schizomida Not living naturally in Europe, but four species have been found, imported from other continents in hothouses and other localities in Spain, France, 324 7 Regional Arachnogeography Great Britain, Germany, Czech Republic, Slovakia, and Poland (Barranco et al. 2014; Armas and Rehfeldt 2015). These are Schizomus crassicaudatus (O.P.Cambridge), Bucinozomus hortuspalmarum Armas et Rehfeldt, Zomus bagnalii (Jackson), and Stenochrus portoricensis Chamberlin. Solifugae In Europe Solifugae are found only in Spain, Portugal, Sicily, Greece, Bulgaria, Macedonia, and Ukraine – altogether eight species of the genera Galeodes Olivier (Galeodidae), Biton Karsch, Gluvia C.L. Koch (Daesiidae), Barrussus Roewer, and Eusimonia Kraepelin (Karschiidae) (Birula 1912, Blick 2004, Roewer 1934). Map 7.2 Distribution of Solifugae in Europe Galeodes – X Gluvia – ● Barrussus – ○ Biton – ■ 7.2 Europe (General) 325 Scorpiones According to Fet (2010 and supplements), in Europe are registered 23 spp. of scorpions, but since several new species have been described and with more detailed checking of several populations and “subspecies,” this number could increase to 35. The European scorpions belong to the families Buthidae (Buthus Leach, 3 spp.; Mesobuthus Vachon, 3 spp.), Euscorpiidae (Euscorpius Thorell, 15 spp.), Iuridae (Iurus Thorell, 1 sp.), and (?)Troglotayosicidae [or Chactidae] (Belisarius Simon, 1 sp.) (Gantenbein et al. 1999, Kovařik 1999). Map 7.3 Scorpiones in Europe ■ – Euscorpius O – Iurus X – Mesobuthus ♦ – Buthus ! – Belisarius 326 7 Regional Arachnogeography Pseudoscorpiones The book of Beier (1963) listed 9 families of pseudoscorpions in Europe (including Canary Islands, Madeira, and Azores), but now (Lissner 2014) there are 73 genera, belonging to 14 families (Chthoniidae, Neobisiidae, Bochicidae, Larcidae, Syarinidae, Olpiidae, Garypidae, Geogarypidae, Garypinidae, Atemnidae, Cheiridiidae, Cheliferidae, Chernetidae, Withiidae) – nearly half of the Pseudoscorpiones families in the world. According to Lissner (2014), “The European pseudoscorpion fauna counts to 861 species in 73 genera and 16 families.” These numbers include species and genera from Asian parts of Russia and Turkey (perhaps 20 species?). The fauna is much more diverse in the southern parts of Europe. The highest number of species is found in Italy (226) followed by Spain (213, including the Canary Islands). Blick et al. (2004) list 102 pseudoscorpion species for Central Europe. Species number decline to about 20 in southern Scandinavia (Beier 1963). “None, except a few introduced synanthropic species, are found at higher latitudes than ca. 65°N” (Beier 1963). Endemic genera of Pseudoscorpiones in Europe: Fam. Chthoniidae Neobalkanella Ćurčić, 2013 – Serbia (one sp.) Troglochthonius Beier, 1939 – Bosnia and Herzegovina, cave (one sp.); ? Sardinia Fam. Neobisiidae Insulocreagris Ćurčić, 1987 – Croatia (Vis), Herzegovina (two spp.) Protoneobisium Ćurčić, 1988 – Croatia (one sp.) Archaeoroncus Ćurčić et Rada, 2012 – Croatia (one sp.) Ernstmayria Ćurčić et Dimitrijević, 2006 – Crete (one sp.) Fam. Syarinidae Arcanobisium Zaragoza, 2010 – Spain (A. comasi Zaragoza, 2010) Hadoblothrus Beier, 1952 – Italy, Greece (two spp.) Pseudoblothrus Beier, 1931 – Italy, Crimea, France, Switzerland (seven spp.) Microcreagrella Beier, 1961 – Azores, Madeira, Spain, Portugal (one sp.) Fam. Bochicidae Titanobochica Zaragoza et Reboleira, 2010 – Portugal (T. magna Zaragoza et Reboleira, 2010) Troglobisium Beier, 1939 – Spain [T. racovitzai (Ellingsen, 1912)] Cyphophthalmi In Europe have been registered 5 genera and 25 species of 1 family (Sironidae), mainly in Southern Europe, with 1 species Siro carpathicus Rafalski going North to Poland and Slovakia. There are two main centers of speciation: Iberian Peninsula with 5 genera and 5 species and Balkan Peninsula with ?17 spe- 7.2 Europe (General) 327 cies of 1 genus (Cyphophthalmus, incl. Tranteeva) (Roewer 1923, Martens 1978, Murienne and Giribet 2009). Eupnoi Two families contain the bulk of European Opilions: Fam. Phalangiidae – Phalangium L., Dicranopalpus Doleschall, Lacinius Thorell, Mitopus Thorell, Egaenus C.L. Koch, Opilio Herbst, Oligolophus C.L. Koch, Paroligolophus Lohmander, Dasylobus Simon, Graecophalangium Roewer, Metaphalangium Roewer, Ramblinus Starega, Rilaena Šilhavý, Paraplatybunus Dumitrescu, Lophopilio Hadži, Megabunus Meade, Metaplatybunus Roewer, Platybunus C.L. Koch, Zachaeus C.L. Koch, Stankiella Hadži, Rafalskia Starega, and others Fam. Sclerosomatidae – Gyas Simon, Cosmobunus Simon, Micronelima Schenkel, Leiobunum C.L. Koch, Nelima Roewer, Astrobunus Thorell, Homalenotus C.L. Koch, Mastobunus Simon, Metasclerosoma Roewer (nine genera) Dyspnoi The European (Caucasus included) fauna of Dyspnoi consists of five families: Nemastomatidae (Nemastoma C.L. Koch, Paranemastoma Redikorzev, Mitostoma Roewer, Giljarovia Kratochvil et Miller, Carinostoma Kratochvil, Nemaspela Šilhavý, Acromitostoma Roewer, Saccarella Schönhofer, Hadzinia Šilhavý, Pyza Staręga, Vestiferum Martens, Histricostoma Kratochvil, Mediostoma Kratochvil, Centetostoma Kratochvil, Caucnemastoma Martens, Nemastomella Mello-Leitão), Sabaconidae (Sabacon Simon), Ischyropsalididae (Ischyropsalis C.L. Koch), Dicranolasmatidae (Dicranolasma Sørensen), and Trogulidae (Trogulus Latreille, Anarthrotarsus Šilhavý, Anelamocephalus Simon, Calathocratus Simon, Kofiniotis Roewer (24 genera and 185 accepted spp.) (Schönhofer 2009). Laniatores In Europe Laniatores are much fewer than the “Palpatores” – only 40 spp. of 12 genera and the families Cladonychiidae (Holoscotolemon Roewer), Phalangodidae, and Travuniidae. Many genera (Lola Kratochvil, Paralola Kratochvil, Travunia Absolon et Kratochvil, Trojanella Karaman, Dinaria Roewer) and species are known only from caves and are considered generally as relict (Kury and Mendes 2007). However, Martens (1972) denies their relict character and thinks that this is a normal process of speciation. Araneae In Europe (including Madeira and Azorean Islands) until the end of 2011 have been registered 4892 species and subspecies of spiders (van Helsdingen 2012). They include 4491 “true” indigenous spiders, 99 introduced species, and 302 nomina dubia. The category of unique records/endemics contains 2041 species (45.4%). These spiders belong to the families Atypidae, Ctenizidae, Nemesiidae, Filistatidae, Sicariidae, Scytodidae, Leptonetidae, Pholcidae, 328 7 Regional Arachnogeography Dysderidae, Segestriidae, Oonopidae, Palpimanidae, Mimetidae, Eresidae, Oecobiidae, Uloboridae, Nesticidae, Theridiosomatidae, Anapidae, Mysmenidae, Araneidae, Linyphiidae, Tetragnathidae, Theridiidae, Lycosidae, Agelenidae, Pisauridae, Oxyopidae, Zoropsidae, Cybaeidae, Argyronetidae, Desidae, Hahniidae, Amaurobiidae, Dictynidae, Titanoecidae, Anyphaenidae, Clubionidae, Liocranidae, Gnaphosidae, Philodromidae, Thomisidae, Salticidae, Corinnidae, Zodariidae, Prodidomidae, Miturgidae (Zorinae), and Sparassidae. (oshte!). The most numerous families (figures of 2011) are Linyphiidae (1366), Gnaphosidae (480), Salticidae (400), Dysderidae (331), Lycosidae (303), Theridiidae (258), Agelenidae (200), Thomisidae (193), Araneidae (150), Zodariidae (111), and Philodromidae (106). The best represented what concerns the endemics are Linyphiidae (534, 42.4%), Agelenidae (93, 50%), and most of all Dysderidae (227, 70.5%). There are many regional lists of the spiders in European countries (Decae 2010, Deltshev 2000, Deltshev et al. 2003, Koponen 1991, Le Peru 2011, Nentwig et al. 2013, Song et al. 2002, Thorell 1869, and others). Opilioacarida In Europe (the southern parts of Italy and Greece) are known only two spp. of Opilioacarida: Opilioacarus segmentatus With, 1903, and O. italicus With, 1904 (Beron 1990, 2014, Brignoli 1967, With 1903, 1904) 7.3 7.3.1 Balkan Peninsula and Aegaeis Geography, General Zoogeography, and Paleogeography The eastmost large peninsula of Southern Europe meets the Adriatic Sea on the Northwest, Ionian Sea on the southwest, the Mediterranean and Aegean Sea on the South and Southeast, and the Black Sea on the East and Northeast. The highest point of the Balkans is Mount Musalla (2925 m) in Rila mountain in Bulgaria. The total area of the Balkans is 666,700 square. The Peninsula got its name from the Balkan Mountains (Stara planina) that stretch from the east of Bulgaria to the very east of Serbia. Several attempts have been made to subdivide zoogeographically the most complex area in Europe – on the contact between the Eurosiberian and the Mediterranean units, with montanmediterranean centers of speciation, with arctic-alpine elements, with many endemic cave animals, and with influence from the Steppic area and Asia 7.3 Balkan Peninsula and Aegaeis 329 Map 7.4 Balkan Peninsula Minor. The approach of various authors is different, but usually following the phytogeographical subdivision (Asenov 2006, Brunn and Mercier 1971, Dermitzakis and Papanikolaou 1981, Buresch and Popov, 1973, Gruev 2000, Guéorguiev 1979, 1982, Matvejev 1961, 1969). The first attempts to analyze the zoogeography of Bulgaria were due to Drenski (= Drensky) (1936, 1946, 1966). This founder of the zoogeography in Bulgaria was an arachnologist himself, and some of his analyses were based largely on the distribution of spiders, the sun spider, and some other arachnids. In several papers, Entomologist V.B. Guéorguiev analyzed the contemporary data of the distribution of animals in Bulgaria and provided a new subdivision of 330 7 Regional Arachnogeography Bulgarian territory. The same author analyzed the cave fauna of Balkan Peninsula (Guéorguiev 1977), taking into account the evolution of the territory in the geologic past and the influence of elements from Gondwana, the concept of Egeida, etc. His speleozoogeographic map of the peninsula is unique even to our time. Gradually, as elsewhere, the biogeographical approach became predominant. In the second edition of the manual of Gruev and Kuzmanov (1994), we find six “faunistic complexes” in Bulgaria: Northern Holarctic, Siberian, European, Euroasiatic steppic, Southwestasian, and Mediterranean. Outlined are five “biogeographic regions”: North Bulgarian, Middle Bulgarian, South Bulgarian, Pontic, and Mountain regions. Such units should not bear names of only one state, as the “South Bulgarian region” stretches also in Turkey, Greece, and the Republic of Macedonia, and in these countries it would be called differently. When subdividing the peninsula and other territories, we should disregard the political boundaries. According to Gruev (2000), the submediterranean elements in the eastern parts of the submediterranean province “belong to the thermos-mesophilic and the thermosxerophilic ecological types in conditions of temperate-continental, transitionalcontinental and transitional-mediterranean climate.” Here are the zoogeographic maps of Bulgaria of Drensky (1946) and Gueorguiev (1980) and the biogeogeographic map of Gruev (1988): Map 7.5 Zoogeographical map of Bulgaria after Drensky (1946) I – Areas with Eurosiberian elements: 1, Danubian Bulgaria; 2, Western Bulgaria; 3, Stara planina; 4, Central High Mountain part. II – Areas with Mediterranean elements: 5, Aegean area; 6, Upper Maritza plain; 7, Pontian area. III – Areas with Irano-Turanian elements: 8, Dobrudja; 9, Lower Maritza Thrace 7.3 Balkan Peninsula and Aegaeis Map 7.6 Zoogeographical map of Bulgaria after Gueorguiev (1980, 1992) ___ Border between the Eurosiberian and Mediterranean territory ----- Borders between the zoogeographical areas 1. Danubian area 2. Area of Stara planina 3. Rilo-Rhodopean area 4. Thracian area 5. Strandja area 6. Black Sea area 7. Struma-Mesta area 331 332 7 Map 7.7 Biogeographical map of Bulgaria after Nikolov (1977) 1. North Bulgarian Steppe and Forest – Steppe Province 1.1 – Danubian Forest-Steppe area 1.2 – Dobrudja Forest-Steppe area 2. Balkan Middle-European forest province 2.1 – North Bulgarian forest area 2.2 – Western-Middle Bulgarian area 2.3 – Osogovo-Maleshevo area 2.4 – South Bulgarian Mountain area 3. Balkan Submediterranean province 3.1 – Struma-Mesta area 3.2 – Upper Thracian area 3.3 – East Rhodopean-Sakar area 3.4 – Strandja area 3.5 – Black Sea area Regional Arachnogeography 7.3 Balkan Peninsula and Aegaeis 7.3.2 333 Arachnogeography Drenski (1936) discussed the distribution of spiders in Bulgaria. Guéorguiev (1992) wrote a special publication on the zoogeography of Opiliones in Bulgaria. Many papers of Ćurčić and his coauthors analyze the distribution of pseudoscorpions, and in the book of Deltshev et al. (2003) have been published maps of all species of Araneae in Serbia. In another monograph (Ćurčić et al. 2004) were outlined all species of Pseudoscorpiones in Serbia, Montenegro, and the Republic of Macedonia. Petrov (1997) wrote a checklist of pseudoscorpions in Bulgaria, completed by Petrov and Štáhlavský (2007). According to the analysis of Deltshev et al. (2005), in Bulgaria were recorded by 2005 1091 spp. of Arachnida (outside the Acari): 925 spiders, 61 harvestmen, 51 false scorpions, 2 scorpions, 1 Solifugae, and 1 palpigrade. These authors think that there are 50 Bulgarian endemics and 68 Balkan endemics, and 240 species are considered rare. Since 2005 some more species have been published (the scorpions are now at least four). The northern limits of some groups (Solifugae, Opilioacarida) cross the peninsula. Some others (Amblypygi, Thelyphonida, Schizomida, Ricinulei, Holothyrida) are absent from it. Guéorguiev (1992) analyzed zoogeographically the fauna of Opiliones in Bulgaria (the 42 genera and subgenera known by this time) and concluded that the “boreal” species are 20 (47,6%), the “meridionals” are 7 (16,7%), and the endemics are 15 (35,7%). Presently (2017), thanks to the efforts of P. Mitov, in Bulgaria are known already 63 spp., so another analysis would be interesting. Gruev (1995) analyzed the “Mediterranean faunistic complex” in Bulgaria, based only on invertebrates and mostly on Coleoptera, Chrysomelidae. He concluded that Bulgaria belongs biogeographically entirely to the European-West Siberian area of the Palearctic Region and that none of the territories of the country belongs to the Mediterranean. It is due mostly to the southern borders of Bulgaria, running on high mountains and not allowing to the typical biotic societies to be part of the present-day Bulgarian territory. Only some elements penetrate the areas with submediterranean climate along the river valley of Struma and to the less extent of Mesta and Maritza. Speaking of Arachnida, among these elements are Galeodes graecus (Solifugae), Euscorpius solegladi, E. drenskii, E. popovi, Mesobuthus gibbosus (Scorpiones), and some Araneae, Opiliones, and Pseudoscorpiones. They are part of the submediterranean fauna in Bulgaria, outlined (in Bulgarian) by Gruev (2000). According to him, the submediterranean biogeographic province includes (intrazonally) territories from Iberian Peninsula to Caucasus (the middle of the Iberian Peninsula and the Apennines, South of France, Corsica, parts of the former Yugoslavia, Hungary, Austria, Switzerland, Northern and Middle Greece, northern part of Turkey, the south of Crimea, and Northern Caucasus up to the Caspian Sea). They are characterized (Matvejev 1961) by the following parameters: average annual temperature of 9,3–11,7 °C, average monthly temperature of the warmest month 20–23,2 °C, negative winter temperature (average monthly) from 1 to 3 334 7 Regional Arachnogeography months, and rainfall of 450–650 mm with dry summer. Such climate is found up to 800–1000 m, in places even up to 1500 m and higher. Among the Arachnida in the paper of Gruev (2000) are listed only four species of spiders (Dysdera bulgarica, Pardosa pseudostrigilata, P. strigilata, Trachelas maculatus), after Deltshev (1976). Actually, they are more. The Republic of Macedonia is also with high mountains on all its borders and has mostly submediterranean fauna. Albania, Montenegro, Dalmatia, and Slovenia are exposed to Adriatic (Mediterranean) climatic influence, and are very karstic, the limestone creating usually comfort to warm-preferring arachnids. According to Deltshev (2004), the spiders of the “Mediterranean complex” of the Balkan Peninsula are 13.8% of the total spider fauna of the peninsula. Many Balkan endemics are also of Mediterranean origin. “The mountain-Mediterranean species (e.g., Aculepeira talishia, Pardosa incerta) are of particular interest, since they may be regarded as ancient element.” Most of the territory of Greece belongs to the typical Mediterranean subregion, characterized best by the areal of the olive trees (Olea europaea). The fauna of the Balkan part of Greece contains many solpugids, scorpions, spiders, and other arachnids, unknown in the areas north of the country. The peninsula has many high mountains (up to 2925 m), harboring glacial relicts and montanmediterranean arachnids. Palpigradi From the countries of the peninsula (south of Danube) are known seven Palpigradi species and three subspecies (all from genus Eukoenenia Thorell, Eukoeneniidae) (Condé 1974, 1976, 1977, 1979, 1990, 1996, Georgescu and Decu 1994, Hansen 1926, Beron 1978, Orghidan et al. 1982). Bulgaria – Beron (1978) reports three specimens of ?E. austriaca from caves in NW Bulgaria. Greece – have been reported E. juberthiei cytheriaca Condé, 1979 (Kythira Island); E. j. hellenica Condé, 1979 (Attika); E. naxos Condé, 1990 (Iraklia Island) Bosnia and Herzegovina – E. remyi Condé, 1974 (Herzegovina) Slovenia – E. austriaca austriaca (Hansen, 1901) Croatia – E. pretneri Condé, 1977; E. spelaea hauseri Condé, 1974 Northern Dobrudja (Romania) – E. mirabilis (Grassi et Calandruccio, 1885); E. condei Orghidan, Georgescu, et Sârbu, 1982; E. margaretae Orghidan, Georgescu, et Sârbu, 1982 (Georgescu & Decu, 1994) Solifugae The list of European Solifugae of Blick (2004) contains 18 spp., but geographically half of them live on territories outside Europe (Rhodes, Asiatic Turkey, Cyprus, Canaries). On the Balkan Peninsula (continental part, mostly in Greece) are recorded members of three families of Solifugae (Drensky 1931, Kraepelin 1899, Roewer 1934): Fam. Galeodidae (Galeodes Olivier): G. graecus C.L. Koch (Greece, Bulgaria), G. elegans Roewer (Republic of Macedonia), G. hellenicus Roewer (Greece) Fam. Karschiidae – Barrussus furcichelis Roewer, Eusimonia nigrescens Kraepelin (Greece) 7.3 Balkan Peninsula and Aegaeis 335 Fam. Daesiidae – Biton ehrenbergi Karsch (Greece) The northern boundary of Solifugae in eastern Europe runs through SW Bulgaria (Galeodes graecus C.L. Koch, 1842), the Republic of Macedonia (G. elegans Roewer, 1934), and Russia (G. araneoides Pallas, 1772) (Map 35). Scorpiones On the Balkan Peninsula (including Slovenia and the islands of the Aegean, except for the Dodecanese), there is a complicated picture of the number of taxa of the scorpions, changing very often. We can agree on the current list following Fet (2010) and many supplements, Crucitti (1995, 1999b), Calinescu and Calinescu (1930), Æurèiæ (1971, 1972), Daniel (1962), Fet (1985, 2000), Fet and Braunwalder (2000), Fet et al. (2001), Fet and Soleglad (2007), Fet et al. (2014), Francke (1981), Kaltsas et al. (2008), Kinzelbach (1975), Kovařik et al. (2010), Michalis and Dolkeras (1989), Michalis and Kattoulas (1981), Parmakelis et al. (2006), Tropea et al. (2015), Vachon (1948, 1953), Voulalas and Michalis (1977), and Werner (1937). Fam. Euscorpiidae Euscorpius (E.) hadzii Caporiacco, 1950 – SW Bulgaria, Macedonia, Albania, Bosnia and Herzegovina, Croatia, Greece, Serbia, Montenegro E. (E.) koschewnikowi Birula, 1900 – NE Greece E. (E.) “carpathicus” complex – Bulgaria, Albania E. (E.) sicanus (C.L. Koch, 1837) – Greece E. (E.) tergestinus (C.L. Koch, 1837) – Albania, Croatia, Slovenia, Montenegro E. (E.) deltshevi Fet, Graham, Webber, et Blagoev, 2014 – West Bulgaria, East Serbia E. (E.) solegladi Fet, Graham, Webber, et Blagoev, 2014 – Southwest Bulgaria, Northern Greece E. (E.) drenskii Tropea, Fet, Parmakelis, Kotsakiozi, et Stathi, 2015 – Southeast Bulgaria E. (Alpiscorpius) mingrelicus dinaricus Caporiacco, 1950 – Bosnia E. (A.) mingrelicus caporiaccoi Bonacina, 1980 – Bosnia E. (A.) beroni Fet, 2000 – Albania E. (Polytrichobothrius) naupliensis (C.L. Koch, 1837) – Greece E. (P.) italicus (Herbst, 1800) – Greece E. birulai Fet, Soleglad, Parmakelis, Kotsakiozi, et Stathi, 2014 – Greece (Euboea) E. mylonasi Fet, Soleglad, Parmakelis, Kotsakiozi, et Stathi, 2014 – Greece (Euboea) E. popovi Tropea, Fet, Parmakelis, Kotsakiozi, et Stathi, 2015 – Pirin, Slavyanka (SW Bulgaria, N. Greece) Fam. Buthidae Mesobuthus gibbosus (Brullé, 1832) – Albania, Macedonia, Greece, Montenegro, and European Turkey. The findings in European Turkey, the locality in SW Bulgaria (Pirin, 1200–1300 m, Teruel et al., 2004), seems doubtful, although the species may be expected near the border with the Republic of Macedonia. 336 7 Regional Arachnogeography Fam. Iuridae Iurus dufoureius (Brullé, 1832) – was restricted by Kovarik, Soleglad, Fet, and Yağmur (2010) only to Greece (Peloponnese, Kythira, and Crete). Pseudoscorpiones On the Balkan Peninsula (within the limits outlined above) have been recorded pseudoscorpions from 44 genera and 13 families: Chthoniidae, Geogarypidae, Garypidae, Garypinidae, Olpiidae, Larcidae, Neobisiidae, Syarinidae, Atemnidae, Cheiridiidae, Chernetidae, Cheliferidae, and Withiidae. Fam. Chthoniidae – Chthonius C.L. Koch, Neobalkanella Ćurčić, Microchthonius Hadzi, Mundochthonius Chamberlin, Paraliochthonius Beier, Troglochthonius Beier Fam. Geogarypidae – Geogarypus Chamberlin Fam. Garypinidae – Amblyolpium Simon, Garypinus Daday Fam. Garypidae – Garypus L. Koch Fam. Olpiidae – Olpium L. Koch, Calocheiridius Beier et Turk, Cardiolpium Mahnert, Minniza Simon Fam. Larcidae – Larca Chamberlin Fam. Neobisiidae – Acanthocreagris Mahnert, Balkanoroncus Ćurčić, Ernstmayria Ćurčić et Dmitrievic, Insulocreagris Ćurčić, Microbisium Chamberlin, Neobisium Chamberlin, Protoneobisium Ćurčić, Roncus L. Koch, Archaeoroncus Ćurčić et Rada Fam. Syarinidae – Hadoblothrus Beier Fam. Cheiridiidae – Apocheiridium Chamberlin, Cheiridium Menge Fam. Atemnidae – Atemnus Canestrini, Diplotemnus Chamberlin Fam. Chernetidae – Allochernes Beier, Chernes Menge, Dendrochernes Beier, Dinocheirus Chamberlin, Lamprochernes Töm., Lasiochernes Beier, Pselaphochernes Beier Fam. Cheliferidae – Beierochelifer Mahnert, Chelifer Geoffroy, Cheirochelifer Beier, Dactylochelifer Beier, Hysterochelifer Chamberlin, Rhacochelifer Beier Fam. Withiidae – Withius Kew In the countries of Balkan Peninsula have been recorded pseudoscorpions as follows: Albania, 18 spp.; Bulgaria, 59 spp.; Serbia (whole, incl. Kosovo), 70 spp.; Montenegro, 43 spp.(50 spp., according to N. Ćurčić and Dimitriević, 2016); Republic of Macedonia, 45 spp.; Greece, 121 spp.; European Turkey, 5 spp.; Croatia (whole), 112 spp.; Slovenia (whole), 31 spp.; and Bosnia and Herzegovina, 55 spp. And these pseudoscorpions were recorded by several authors: Beier (1929, 1931, 1939, 1963), Beron (1968, 1985), Ćurčić (many papers since 1972), Ćurčić and Dimitriević (1984, 2006), Ćurčić et al. (2004), N. Ćurčić and Dimitriević (2016), Ćurčić et al. (1993), Dumitresco and Orghidan (1964), Georgescu and Capuşe (1994, 1996), Hadži (1940), Harvey (2011), Mahnert (1973–1982), Ozimec (2000), Petrov (1997, 2000), Petrov and Štáhlavský (2007), Redikorzev (1928), and Schmalfuss and Schawaller 7.3 Balkan Peninsula and Aegaeis 337 (1984). For many species has been used the database of Harvey (2013f) but have been extracted the subspecies and the repeated names (e.g., Chelifer cancroides, Ch. cancroides cancroides). Endemic genera for the peninsula are: Fam. Chthoniidae Neobalkanella Ćurčić, 2013 – Serbia (two spp.) Troglochthonius Beier, 1939 – Bosnia and Herzegovina, cave (two spp.) Fam. Neobisiidae Insulocreagris Ćurčić, 1987 – Croatia (Vis Island), Herzegovina (two spp.) Protoneobisium Ćurčić, 1988 – Croatia (one sp.) Archaeoroncus Ćurčić, 2012 – Croatia (one sp.) Ernstmayria Ćurčić et Dimitriević, 2006 – Crete (one sp.) Besides the endemic genera (all considered relics), particularly interesting is the finding of Hadoblothrus aegeus Beron in caves on Santorini Island – clearly relict and the only member of Syarinidae found so far from Balkan Peninsula. The second species of this genus is known from a cave in Southern Italy. Opiliones On the Balkan Peninsula (including the entire territories of Slovenia and Croatia, Northern (Romanian) Dobrudja, Eastern (Turkish) Thrace and the adjacent islands in Aegean and Ionian seas) have been recorded 60–62 genera of Opiliones from 9 families (Sironidae, Travuniidae, Phalangodidae, Phalangiidae, Ischyropsalididae, Nemastomatidae, Trogulidae, Dicranolasmatidae, Sclerosomatidae) (Boyer et al. 2005, Guéorguiev 1992, Hadži 1928, 1973a, 1973b, Kratochvil 1937, 1946, Novak 2005b, Novak et al. 1995, 1996), Rambla 1968, Roewer 1926, 1927b, 1940, 1959, Schönhofer 2009, Schönhofer and Martens 2009, Schönhofer et al. 2009, Ubick and Ozimec 2005). The following are a number of registered taxa of Opiliones in the Balkan countries: Albania (Mitov 2000, with suppl.; Murányi 2013, 2015) – 46 spp. Bosnia and Herzegovina (Novak 2005a) – 54 valid species and 2 subspecies Bulgaria (Beron and Mitov 1996, Juberthie 1991, Kratochvil 1958a, 1958b, Mitov 1994–2011, Roewer 1926, Staręga 1976, Šilhavý 1965) – 63 spp. European Turkey (Gruber, 1969; Bayram et al., 2010) – 14 spp. Continental Greece (Juberthie 1968, Martens 1972, Rambla 1968, Thaler 1996) – the families Sironidae, Trogulidae, Dicranolasmatidae, Nemastomatidae, Phalangiidae, Sclerosomatidae, Phalangodidae, Phalangiidae, Sclerosomatidae (Siro, Phalangium L., Dicranopalpus Doleschall, Lacinius Thorell, Mitopus Thorell, Egaenus C.L. Koch, Opilio Herbst, Oligolophus C.L. Koch, Paroligolophus Lohmander, Dasylobus Simon, Graecophalangium Roewer, Metaphalangium Roewer, Ramblinus Staręga, Rilaena Šilhavý, Lophopilio Hadži, Megabunus Meade, Metaplatybunus Roewer, Platybunus C.L. Koch, Zachaeus C.L. Koch, Stankiella Hadži, Rafalskia Staręga) 338 7 Regional Arachnogeography Crete (Martens 1965, 1966, Gruber 1998, Roewer 1927b, 1940; Šilhavý 1967) – genera Trogulocratus Roewer, Konfiniotis Roewer, Anarthrotarsus Šilhavý, Platybessobius Roewer, Dicranolasma Sørensen, etc., and the families Trogulidae, Dicranolasmatidae, Phalangiidae, Sclerosomatidae Aegean Islands (di Caporiacco 1928, 1948; Ghigi 1929; Gruber 1963, 1978; Martens 1965; Roewer 1924) – 13 spp., mostly from Rhodes Croatia (Novak 2004) – 64 valid species and 1 subspecies Slovenia (Novak, Delakorda, and Novak, 2006) – 64 spp. Republic Macedonia (Mitov 1995) – Graecophalangium drenskii Mitov – endemic Northern Dobrudja (Dumitrescu 1970, 1972; Dumitrescu and Orghidan, 1964; Babalean 1999, 2005) – 15 species; for Southern Dobrudja (in Bulgaria), Mitov (2008) lists 17 species. Cyphophthalmi As Karaman (2008) points out, the Balkan region “with New Zealand and tropical South-East Asia, is one of the richest in cyphophthalmid on the Earth.” From ca.140 spp. in suborder Cyphophthalmi, ca. 20 have been recorded from Balkan Peninsula. Most species belong to Cyphophthalmus Joseph (on the Balkans live 17 species of this genus). Endemic genus and species Tranteeva paradoxa Kratochvil, 1958, is living in the caves of Central Stara planina (Bulgaria). Karaman (2009) thinks that genus Tranteeva should be synonymized with Cyphophthalmus. The situation for the Balkan Peninsula, with an explosive evolution of only one genus of Cyphophthalmi, is very different from the one for the Iberian Peninsula. This territory contains four of the eight genera currently recognized in the family Sironidae, a generic diversity and morphological disparity of Cyphophthalmi not found in any other region of the world so far (Murienne and Giribet, 2009). Although the two European peninsulas have usually been depicted as glacial refugia, we provide evidence that old endemic lineages in these two territories have undergone very different diversifications: one – the Balkan Peninsula – by hosting an old genus with subsequent explosive evolution; and the other – the Iberian Peninsula – by hosting many ancient genera, each with few species. These differences could be related to the very different palaeogeographic histories of the two peninsulas. (Murienne et al. 2009) As elsewhere in Europe, the bulk of Balkan Opiliones are the representatives of “Palpatores.” Eupnoi Fam. Phalangiidae – Amilenus Martens, Dasylobus Simon, Dicranopalpus Doleschall, Egaenus C.L. Koch, Lacinius Thorell, Lophopilio Hadži, Megabunus Meade, Metaphalangium Roewer, Metaplatybunus Roewer, Mitopus Thorell, Odiellus Roewer, Oligolophus C.L. Koch, Opilio Herbst, Phalangium L., Platybunus C.L. Koch, Rilaena Šilhavý, Rafalskia Staręga, Zachaeus C.L. Koch, Graecophalangium Roewer, Metadasylobus Roewer Fam. Sclerosomatidae – Astrobunus Thorell, Gyas Simon, Leiobunum C.L. Koch, Nelima Roewer, Homalenotus C.L. Koch, Mastobunus Simon 7.3 Balkan Peninsula and Aegaeis 339 Dyspnoi Fam. Ischyropsalididae – Ischyropsalis C.L. Koch (two spp., Slovenia) Fam. Dicranolasmatidae – Dicranolasma Sørensen (five spp. from Corfu, Crete, Bulgaria, Montenegro, and Herzegovina) Fam. Nemastomatidae – Paranemastoma Redikorzev (11 spp., Bulgaria, Albania, Herzegovina, Greece) Fam. Trogulidae – Trogulus Latreille (13 spp., Bulgaria, Greece, Corfu, BosniaHerzegovina, Macedonia, Croatia (Mljet, Hvar, Dubrovnik), Montenegro, Serbia, Albania), Anarthrotarsus Šilhavý (one sp., Kerkira, Crete), Calathocratus Simon (six spp., Russia, Crete, Rhodes), Konfiniotis Roewer (one sp., Crete) Laniatores Described from Balkan Peninsula is the family Travuniidae, known from Croatia, Montenegro, Bosnia and Herzegovina, Serbia, Slovenia, Italy, France, Spain, Switzerland, and Sardinia. From the 15 species in this family, 7 live on the peninsula, mostly in caves (Kury and Mendes 2007). From a cave in Serbia (near Bulgarian border) was described the most interesting new genus and species Trojanella serbica Karaman, 2005. The family Phalangodidae is known from Bulgaria with the endemic genus and species Paralola buresi Kratochvil, 1958 (four caves in western Stara Planina). In Greece have been described three spp. of Ausobskya (Martens 1972, Thaler 1996). One new species of Ausobskya was found by us in Belasitsa Mountain (Bulgaria, on the border with Greece), in leaf litter. Endemic genera of Opiliones on the Balkan Peninsula are: [Tranteeva Kratochvil, 1958] – Bulgaria (one sp.) (= Cyphophthalmus?) Paralola Kratochvil, 1958 – Bulgaria (one sp.) Lola Kratochvil, 1937 – Croatia (Hvar) (one sp.) Ausobskya Martens, 1972 – Greece, Bulgaria (four spp.) Trojanella I. Karaman, 2005 – Serbia (one sp.) Travunia Absolon et Kratochvil, 1932 – Bosnia and Herzegovina, Croatia, Montenegro (four spp.) Dinaria Hadži, 1932 – Bosnia and Herzegovina (one sp.) Anarthrotarsus Šilhavý, 1967 – Greece (one sp., Kerkira, Crete) Kofiniotis Roewer, 1940 – Greece (one sp., Crete) Hadzinia Šilhavý, 1966 – SW Bosnia (one sp.) Araneae On the Balkan Peninsula (in the dimensions described above) have been registered over 1409 species of spiders of 337 genera and 48 families: Atypidae, Ctenizidae, Nemesiidae, Filistatidae, Sicariidae, Scytodidae, Leptonetidae, Pholcidae, Dysderidae, Segestriidae, Oonopidae, Palpimanidae, Mimetidae, Eresidae, Oecobiidae, Uloboridae, Nesticidae, Theridiosomatidae, Anapidae, Mysmenidae, Araneidae, Linyphiidae, Tetragnathidae, Theridiidae, Lycosidae, Agelenidae, Pisauridae, Oxyopidae, Zoropsidae, Cybaeidae, Argyronetidae, Desidae, Hahniidae, Amaurobiidae, Dictynidae, Titanoecidae, Anyphaenidae, Clubionidae, Liocranidae, Gnaphosidae, Philodromidae, Thomisidae, Salticidae, Corinnidae, Zodariidae, Prodidomidae, Miturgidae (Zorinae), and Sparassidae 340 7 Regional Arachnogeography (Bayram 2002, Bayram et al. 2014, Delchev 2000, 2004, Deltshev and Blagoev 2001, Deltshev et al. 2005, Schröder et al. 2011, Tatole 2006). Albania (Deltshev et al. 2011) – 335 spp. of 36 families Bulgaria (Deltshev and Blagoev 2001; Deltshev 1996, 1999, 2005, 2011) – > 1000 spp. Bosna and Herzegovina (Komnenov, 2009) European Turkey (Topçu, Demir, and Seyyar 2005; Demircan and Topçu 2016) – 197 spp. Continental Greece (Bristowe 1934, Brignoli, many papers, Metzner 1999, Bosmans and Chatzaki 2005; Chadzaki et al. 2015) – in mid-2015 from the entire territory of Greece are known 1100 spp. of 50 families. Crete (Bosmans et al. 2013) – 430 spp. (incl. 57 endemic), 44 families Croatia – 614 species Republic of Macedonia (Blagoev 2002; Komnenov 2006) – 558 spp. of 36 families Serbia (incl. Kosovo) (Deltshev, Ćurčić and Blagoev 2003; Ćurčić et al. 2007) – 638 spp. of 224 genera and 36 families The first catalogue of spiders of the Balkan Peninsula was the one of Drensky (1936) with 1066 spp. of 35 families. Modern zoogeographical analyses were made by Deltshev (1999, 2000, 2004, 2008) and others. According to Deltshev (1999), the spider fauna of the Balkan Peninsula is represented by 1409 species of 337 genera and 48 families (now they are more). This number is relatively high, compared with the 1001 spp. of the huge Russian Plain and the 925 spp. of Germany. Deltshev is classifying these spiders in 24 zoogeographical categories in 4 main groups: 1. Widely distributed species (533, or 38,1% of all), mainly Palearctic 2. Balkan endemics (379 spp., 26,9%), largely due to the many thousands of caves and the complex orography of the peninsula, with high mountains and many islands 3. The European complex includes 300 species (21,3%) 4. The Mediterranean complex counts 195 spp. (13,8%), but part of the endemics are also of Mediterranean origin. The conclusion of Deltshev is that “…the Balkan Peninsula represents one of the main centres of speciation in Europe.” Again after Deltshev (2004), on the Balkan Peninsula, there are 379 endemic species of spiders, but now, largely thanks to researchers like Deltshev, Lazarov, Komnenov, Chadzaki, Bosmans, and others, this figure is increased (Deltshev, in. lit.). Some endemic genera of spiders on the Balkan Peninsula are: Fam. Linyphiidae Antrohyphantes Dumitrescu, 1971 Fageiella Kratochvil, 1934 Fam. Dysderidae Dysderocrates Deeleman-Reinhold et Deeleman, 1988 (Balkans) 7.3 Balkan Peninsula and Aegaeis 341 Rhodera Deeleman-Reinhold, 1989 (Crete) (one sp.) Folkia Kratochvíl, 1970 (Balkans) (seven spp.) Minotauria Kulczyn’ski, 1903 (Crete) (two spp.) Stalagtia Kratochvíl, 1970 (Balkans, Greece) Parastalita Absolon et Kratochvíl, 1932 (Bosnia-Herzegovina) (one sp.) Stalita Schiödte, 1847 (Balkans) Stalitella Absolon et Kratochvíl, 1932 (Balkans) (one sp.) Fam. Liocranidae Arabelia Bosselaers, 2009 (one sp.), Vankeeria Bosselaers, 2012 (one sp.) – Greece Fam. Agelenidae Hadites Keyserling, 1862 – Croatia (one sp.) Fam. Leptonetidae Barusia Kratochvil, 1978 (Croatia, Greece, Montenegro) Sulcia Kratochvil, 1938 (Balkans, Greece) Fam. Pholcidae Stygopholcus Absolon et Kratochvil, 1932 – Croatia, Bosnia-Herzegovina, Greece, Montenegro Fam. Hahniidae Cryphoecina Deltshev, 1997 – Montenegro (one sp.) According to Deltshev (2000), “The endemic taxa of spiders (Araneae) in the Balkan peninsula are represented by 348 species included in 30 families. Countries with the highest number of recorded endemic species are Greece (115), Croatia (68), Bulgaria (55), Bosnia (41), Crete (46).” More than 160 of these species are from caves. Since 2000 some more taxa have been described. Again from Deltshev (2000), This contribution is the result of a critical data available for the endemic spiders of the Balkan peninsula territory and comprises 348 species from 30 families: Ctenizidae 5, Nemesiidae 4, Pholcidae 9, Leptonetidae 21, Segestriidae 2, Dysderidae 7, Oonopidae 1, Palpimanidae 1, Uloboridae 1, Nesticidae 6, Theridiidae 5, Anapidae 1, Mysmenidae 1, Linyphiidae 109, Tetragnathidae 2, Araneidae 1, Lycosidae 1, Agelenidae 29, Cybaeidae 1, Hahniidae 5, Dictynidae 1, Amaurobiidae 17, Liocranidae 4, Clubionidae 3, Zodariidae 8, Gnaphosidae 18, Miturgidae (Zorinae) 1, Philodromidae 2, Thomisidae 4, Salticidae 12. The established number is high and represents 25% of all spiders of the Balkan peninsula. The most characteristic families are: Linyphiidae s. l. (31.3%), Dysderidae (21%) and Agelenidae (8.3%). The genus Troglohyphantes is the most numerous and can be regarded as a faunistic phenomenon since from all 53 species established in the territory of the Balkans, 52 are endemics, distributed mainly in caves…The high percentage of endemic spiders (25%) suggests an important process of autochthonous speciation. So the Balkan Peninsula can be considered as a main center of speciation in Europe. The bigger number (32) and percentage (74.41%) of Bulgarian endemics and certainly the presence of paleoendemic genus Antrohyphantes show the importance of eastern part of Balkan Peninsula for the speciation in the region. The local elements comprise paleoendemics (mainly in caves) and neoendemics (mainly in highaltitude zones). The endemics belong to two principal faunistic complexes: Mediterranean and European (Deltshev 1996). 342 7 Regional Arachnogeography In his analysis of the biogeographical connections between the spiders of the Balkans and the Middle East, Brignoli (1986) concludes that these territories “… forment une unité biogéographique dont la faune aranéologique est significativement différente de celle de la “Mitteleuropa” lato sensu et, au même temps, est considerablement semblable à celle de la Méditerranée occidentale.” Opilioacarida The order is known only from the southernmost part of the peninsula (Peloponnese and the islands Karpathos and Kassos), with one species – Opilioacarus segmentatus With, 1903. The species is known also from Algeria, Sicily, and Rhodes (Beron 1990 2014, Thaler and Knoflach 2002). These localities mark the northern limit of the modern Opilioacarida in Europe, but as fossils two species have been described from Baltic amber. Acari From Bulgaria are recorded 1673 species of Acari (Acariformes and Parasitiformes) (Beron 2011, actually about 1700 spp.). Acariformes Trombidiformes Prostigmata Fam. Erythraeidae Endemic genera and subgenera: Helladerythraeus (Beron, 1988) – Rhodes and Kythnos Islands Myrmicotrombium (Graecotrombium) mirum Beron, 1990 – Greece Parasitiformes Ticks are of some zoogeographical interest. On Balkan Peninsula have been recorded five spp. of Argasidae. Ixodidae are recorded in various Balkan countries like Bulgaria (39 spp.), Turkey (38), Romania (25), Serbia (25), Bosnia and Herzegovina (20), Montenegro (18), and Croatia (13) (Drenski 1955, Beron 2011, 2014, Feider 1965, Georgieva and Gecheva 2013) 7.4 7.4.1 Apennine Peninsula Geography, General Zoogeography, and Paleogeography The central and the smallest of the three large peninsulas of Southern Europe spans 1000 km from the Po Valley in the north to the central Mediterranean Sea in the south. The backbone of the peninsula consists of the Apennine Mountains, high up to 2912 m (Corno Grande del Gran Sasso). The surface of the peninsula is 131,337 km2, and the climate is mediterranean. The mountains are mostly verdant, although one side of the highest peak, Corno Grande, is partially covered by Calderone glacier, the only glacier in the Apennines. The Apennines were created in the Apennine orogeny beginning in the Early Neogene (about 20 mya, the Middle Miocene) and continuing today (Gueguen et al. 1998, Rook et al. 2006). 7.4 Apennine Peninsula 343 La Greca (1962) outlined several “types of distribution” of elements in the Italian fauna (enumerated further). This author analyzed many other zoogeographical aspects of the insects in Italy (La Greca 1955, 1958, 1966). 7.4.2 Arachnogeography In the peninsular part of Italy are recorded the orders Palpigradi, Scorpiones, Pseudoscorpiones, Opiliones, Araneae, Opilioacarida, and different other Acari. Lacking are the “southern” orders – Ricinulei, Amblypygi, Thelyphonida (Uropygi), Schizomida, Solifugae, Holothyrida – and the suborders Mesothelae and Palaeoamblypygi. Some orders are well represented – pseudoscorpions (226 spp.) and spiders (1620 spp.) – for the entire territory of Italy. Palpigradi So far from continental Italy have been recorded five spp. of Palpigradi, including the discovery of the order by Grassi and Calandruccio (1885). They all belong to Eukoenenia Börner (Condé 1976, Silvestri 1905). Scorpiones On the continental Apennine Peninsula are recorded, after many changes, four species of Scorpiones (Crucitti 1993, Fet 2010). Fam. Euscorpiidae Euscorpius (Polytrichobothrius) italicus (Herbst, 1800) E. (Alpiscorpius) germanus (C.L. Koch, 1837) – populations in the Apennines are relict or introduced (Fet et al., 2004) E. (Euscorpius) sicanus (C.L. Koch, 1837) E. (Tetratrichobothrius) flavicaudis (DeGeer, 1778) Pseudoscorpiones According to the catalogue of Gardini (2000), in peninsular Italy (without the Alpine part and the islands) were recorded until the end of 1998 pseudoscorpions of the families Chthoniidae, Neobisiidae, Syarinidae, Garypidae, Geogarypidae, Cheiridiidae, Olpiidae, Atemnidae, Chernetidae, Cheliferidae, and Withiidae (Beier, many papers, Gardini 2015, Lazzeroni 1970, and other papers). As a whole, from Italy are known 226 spp. (Lissner 2014) (first place in Europe). Opiliones According to Stoch (2003), in the total of Italian territory, there are 11 families of Opiliones with 43 genera and 120 species (Brignoli 1968, Canestrini 1872, Chemini 1990, 1995, 1996, Gruber 1965, 1985, Marcellino 1968, 1971, 1982, 1983, 1984, 1986, Staręga 1976, Šilhavý 1969). Cyphophthalmi On the Peninsula have been registered Siro valleorum Chemini, 1990 (Lombardia). 344 7 Regional Arachnogeography Eupnoi From continental Italy: Fam. Phalangiidae – Phalangium L., Dicranopalpus Doleschall, Lacinius Thorell, Mitopus Thorell, Egaenus C.L. Koch, Opilio Herbst, and others Fam. Sclerosomatidae – Metasclerosoma Roewer, Astrobunus Thorell, Homalenotus C.L. Koch Dyspnoi Fam. Ischyropsalididae – Ischyropsalis C.L. Koch (five spp.) Fam. Trogulidae – Trogulus Latreille (two spp.) Fam. Dicranolasmatidae – Dicranolasma C.L. Koch (one sp.) Fam. Nemastomatidae – Nemastoma C.L. Koch, Saccarella Schönhofer et Martens (two spp.) Laniatores Ptychosoma vitellinum Soerensen, 1873 (Phalangodidae), was reported by Brignoli (1968) from Southern Italy. Araneae The list of Stoch (2003) indicates that in the total of Italian territory, there are 45 families of spiders of 323 genera and 1411 species. According to the catalogue of Pantini and Isaia (2015), there are 54 families with 426 genera, 1620 species, and 26 subspecies. So, there is a considerable difference. Brignoli (1981) analyzed the entire Italian fauna of Araneae, known up to this time. Opilioacarida The two European species of the order Opilioacarida Opilioacarus italicus With, 1904, and O. segmentatus With, 1903, are represented in Italy (Brignoli 1967, With 1904). 7.5 7.5.1 Iberian Peninsula and the Pyrenees Geography, General Zoogeography, and Paleogeography On the westernmost of the three big peninsulas in South Europe are situated two states – Spain and Portugal. Iberian Peninsula has an area of approximately 580,000 km2. On the southern end, near the Moroccan coast of Africa, is situated the British overseas territory of Gibraltar. To the north the Pyrenees (highest point Pico de Aneto, 3404 m) forms the border with France (incl. the tiny state of Andorra) and stretches from the Atlantic Ocean to the Mediterranean Sea over 491 km. In the southern part (Andalusia) raises the highest mountain of the Peninsula Sierra Nevada (highest point Mulhasén, 3478 m). About ¾ of the peninsula is taken by a plateau called Meseta Central. I had the chance to study the highest parts of Sierra Nevada, “...das südlichste Hochgebirge Europas und zugleich das höhste Gebirge der Iberischen Halbinsel” 7.5 Iberian Peninsula and the Pyrenees 345 (Franz 1979). It is a very dry mountain, with precipitations at 1800 m of only 6735 mm, compared to the 1130 mm at the same altitude on Vitosha (Bulgaria). The difference between the treeless Sierra Nevada and the humid, green Pyrenees is striking. The autoroute ends below the very top of Veleta (3428 m), and one is among the more humid upper parts of the mountain, among the southernmost snow fields, glaciers, and mountain lakes in continental Europe (ca. 37° N) (Höllermann 1972). The botanists have discovered here a mixed community of glacial relicts and betic variants of Alpine plants. The interesting studies of Janetschek (1957), Franz (1979), as well as my own observations show that some typically alticolous species of spiders, opilions, pseudoscorpions, and mites reach the top of the mountain (Mulhacen, 3482 m). 7.5.2 Arachnogeography Situated in the westernmost part of Europe and near Africa, rich in caves and with some of the highest mountains of the continent, the Iberian Peninsula and the Pyrenees have varied arachnofauna with many relicts and strange taxa. Such is the enigmatic scorpion Belisarius Simon, a relative of South American scorpions. Other such relicts are the pseudoscorpions of the family Bochicidae – two caveinhabiting genera (Titanobochica Zaragoza et Reboleira and Troglobisium Beier) belong to one clearly American family. Remarkable is the presence of three endemic genera of suborder Cyphophthalmi, forming one of the most important centers of speciation of this suborder of Opilions. In Spain are registered scorpions of genus Buthus Leach, unknown in other parts of Europe. Similar is also the situation with the Solifugae (Daesiidae) – though the peninsula is passing the northern limit of distribution of this order. Iberian endemics are 236 species of spiders. The paper of Bacelar (1928) “Aracnidios Portuguèses” contains lists of 330 spp. of spiders, 13 pseudoscorpions, 1 scorpion, 28 opilions, and only 45 spp. of acari of three families. Now they are many more. Barranco et al. (2014) recorded from Spain the imported schizomid Stenochrus portoricensis Chamberlin. Palpigradi In the peninsula (all from Spain and from the two slopes of the Pyrenees) have been recorded Eukoenenia bouilloni Condé, 1980; E. brolemanni Hansen, 1926; E. draco zariquieyi Condé, 1951; E. hispanica Peyerimhoff, 1908; E. pyrenaella Condé, 1990; E. pyrenaica Hansen, 1926; E. gadorensis Mayoral et Barranco, 2002; and E. mirabilis Grassi et Calandruccio, 1885. From continental Portugal is known E. mirabilis (Condé 1951, 1980, 1990, Hansen 1926, Mayoral and Barranco 2002b, Peyerimhoff 1908). Solifugae In Iberian Peninsula is known Gluvia dorsalis (Latreille, 1817) (Daesiidae) (Spain, Portugal) (Pocock 1903, Werner 1925). 346 7 Regional Arachnogeography Scorpiones In the continental part of Iberian Peninsula (incl. Pyrenees) are recorded the following scorpions (Simon 1879, Werner 1925, Lourenço and Vachon 2004, Fet 2010): Fam. Buthidae Buthus occitanus (Amoreux, 1789) – Spain, Portugal (recorded erroneously also from Greece and other areas) B. ibericus Lourenço et Vachon, 2004 – Spain B. montanus Lourenço et Vachon, 2004 – Spain Fam. Euscorpiidae Euscorpius (Tetratrichobothrius) flavicaudis (DeGeer, 1778) – Western Mediterranean Fam. ? Troglotayosicidae Belisarius xambeui Simon, 1879 – Pyrenees in France and Spain (Catalonia) Pseudoscorpiones In continental Spain, Portugal, and the whole of the Pyrenees are known pseudoscorpions of 45 genera and the families Chthoniidae, Neobisiidae, Syarinidae, Bochicidae, Larcidae, Olpiidae, Geogarypidae, Garypinidae, Garypidae, Cheiridiidae, Atemnidae, Chernetidae, Cheliferidae, and Withiidae (Beier 1939, 1959, 1961, Mahnert 1977, Navás 1925, Nonidez 1917, Vachon 1940, Zaragoza 1986, 2000, full bibliography; 2007, 2010). With 196 spp. (Lissner 2014, actualized 2017 – 213 spp.), Spain (incl. Canary Islands) is the second richest country in Europe. From Portugal have been recorded 62 spp. of 13 families. The new pseudoscorpion genus Lusoblothrus of the family Syarinidae is described from a cave in the Algarve region, southern Portugal, to accommodate L. aenigmaticus sp. nov., whose morphological affinities within the Holarctic syarinid fauna are not clear and resembles the Gondwanan genera. This discovery emphasizes the relevance of the Algarve region as a hotspot for relictual hypogean fauna within the Iberian Peninsula. (Reboleira et al. 2012) Endemic genera for the peninsula are: Titanobochica Zaragoza et Reboleira, 2010 – Portugal (T. magna Zaragoza et Reboleira, 2010) Arcanobisium Zaragoza, 2010 – Spain (A. comasi Zaragoza, 2010) Troglobisium Beier, 1939 – Spain [T. racovitzai (Ellingsen, 1912)] Lusoblothrus Reboleira et al., 2012 – Portugal [L. aenigmaticus Reboleira et al., 2012] Opiliones According to Rambla (1973), by this time the opilionid fauna of the Iberian Peninsula consisted of 138 spp., including 4 Cyphophthalmi, 8 Laniatores, and 125 “Palpatores” (Eupnoi and Dyspnoi). Since this list has been completed by de Bivort and Giribet (2004), now the figures are 5 Cyphophthalmi, 8 Laniatores, 7.5 Iberian Peninsula and the Pyrenees 347 and 125 “Palpatores.” Endemic to the peninsula are three genera. For Cyphophthalmi the Iberian Peninsula is a hot spot (Murienne and Giribet 2009). The recent treatments of the Opiliones of the peninsula and the Baleares are 113 spp. and 79 endemic spp. (Mello-Leitão 1936, Prieto 2003, 2008 and suppl., based on Juberthie 1956, 1961, 1962, Kraus 1961, Prieto 1990a, 1990b, 2004, Rambla 1967, 1977b, Rambla and Fontarnau 1984, 1986, etc.). Cyphophthalmi Fam. Sironidae Odontosiro Juberthie, 1961 – Portugal (one sp.) (endemic genus) Iberosiro de Bivort et Giribet, 2004 – Portugal (one sp.) (endemic genus) Paramiopsalis Juberthie, 1962 – Portugal, Spain (two spp.) (endemic genus) Cyphophthalmus [Siro] Joseph, 1868 – Portugal (two spp.) Parasiro Hansen et Sorensen, 1904 – Spain (two spp.) Eupnoi Fam. Phalangiidae Metaphalangium Roewer, 1911; Odiellus Roewer, 1923; Phalangium L., 1758; Eudasylobus Roewer, 1911; Mitopus Thorell, 1876; Dicranopalpus Doleschall, 1852 Fam. Sclerosomatidae Homalenotus C.L. Koch, 1839; Cosmobunus Simon, 1879; Leiobunum C.L. Koch, 1839; Mastobunus Simon, 1879; Micronelima Schenkel, 1938 Dyspnoi On the peninsula are known 25 spp. (Schönhofer, 2009, 2013). Fam. Ischyropsalididae – Ischyropsalis C.L. Koch, 1839 (eight spp. from Spain, three from Portugal) Fam. Sabaconidae – Sabacon Simon, 1879 (six spp.) Fam. Nemastomatidae – Acromitostoma Roewer, 1951 (two spp., endemic genus) Centetostoma Kratochvil, 1958 (three spp.) Fam. Trogulidae – Trogulus Latreille, 1802 (three spp.) Laniatores On the peninsula: fam. Phalangodidae with eight spp.: Ptychosoma Sørensen, 1873 – P. espanoli (Rambla, 1975), Spain Scotolemon Lucas, 1860 – seven spp. from Spain and Portugal According to Rambla (1974), on the peninsula have been recorded 138 spp. of Opiliones, including 56 endemics. From the remaining 82 spp., 24% are of Mediterranean type and 8% have North African affiliations. Since 1974 several new taxa have been added to this fauna (Iberosiro de Bivort et Giribet, 2004 and others), and many taxonomic changes took place (Prieto 2008) but the general picture did not change too much. 348 7 Regional Arachnogeography Araneae According to Cardoso and Morano (2010), “At present [end 2009], 1335 species are known from the region, of which 236 are Iberian endemics, in 373 genera and 55 families. Portugal presents 768 species and Spain (including Andorra and Gibraltar), 1213 species. Although linyphiids present the highest number of known species (267), dysderids present the highest endemic richness (46 species).” Speleoharpactea Ribero is an endemic genus in Spain. Bosmans and de Keer (1985, 1987) prepared a catalogue of the spiders of the Pyrenees and zoogeographical analysis. Their list contained 860 spp. of 43 families (238 Linyphiidae, 92 Gnaphosidae, 73 Salticidae, 60 Theridiidae, 53 Lycosidae, 44 Thomisidae), or these 6 families include 560 spp. and the remaining 37 families include 300 spp. At or above 2200 m have been found at least 170 spp. and above 3000 m at least 22 spp. of 6 families, including 10 Linyphiidae. The highest altitudes are reached (after these authors) by Savignia superstes (3290 m), Gnaphosa atramentaria (3190 m), and Talavera petrensis (3180 m). However, I have found spiders on top of the highest summit (Aneto, 3704 m). The highest spiders of Sierra Nevada have been studied by Denis (1957). Alderweireldt and Bosmans (2001) completed the araneofauna of Portugal to 649 spp. of 43 families. The linyphiid fauna of Portugal was reviewed by Bosmans et al. (2010). 7.6 Mediterranean Sea and Its Islands Map 7.8 Mediterranean Sea and its islands The sea between Africa, Europe, and Western Asia inherited what was once the western arm of the Tethys Sea. Its surface is ca. 2.5 million km2. The sea is connected with the Atlantic Ocean by Gibraltar Strait (14 km wide) and is sometimes 7.6 Mediterranean Sea and Its Islands 349 considered part of the Ocean (and Marmara and Black seas are considered parts of the Mediterranean). In the Middle Miocene times, the collision between the Arabian microplate and Eurasia led to the separation between the Tethys and the Indian oceans. This process resulted in profound changes in the oceanic circulation patterns, which shifted global climates toward colder conditions. The Hellenic arc, which has a land-locked configuration, underwent a widespread extension for the last 20 Ma. The opening of small oceanic basins of the central Mediterranean follows a trench migration and back-arc opening process that occurred during the last 30 Myr. This phase was characterized by the anticlockwise rotation of the Corsica-Sardinia block, which lasted until the Langhian (ca.16 Ma). Subsequently, a shift of this active extensional deformation led to the opening of the Tyrrhenian basin. The Betic-Rif mountain belts developed during the Mesozoic and Cenozoic times, as Africa and Iberia converged. One of the most important events in geological history of Mediterranean Sea is the so-called Messinian Salinity Crisis (Hsü et al. 1973). There are several reasons why the sea, closed in the Miocene, is drying out during the end of the Messinian phase 5.96 to 5.33 Ma. The sea level drops by several kilometers below the level of the World Ocean. On the sea bottom remain only several hypersaline “pools.” After 5.5 Ma the rivers import more water, and the hypersaline “dead seas” are diluted, until by 5.33 Ma opens the Gibraltar Strait. The huge waterfall bringing Atlantic water into the Mediterranean bed is called since 1972 the Zanclean Deluge. After some researchers, the waterfall, although in gradins, has been higher from the highest present-day waterfall in the world (Angel Falls – 979 m) and more mighty than Iguacu or Niagara. It has been calculated the filling of the sea during 2 years (may be less) with a water volume 1000 times bigger than Amazonia, and the level raises by 10 m a day. The dry bed of the Mediterranean is of crucial importance for the zoogeography. Many different animals living on the former sea bottom found asylum on the present-day islands (Malta, Sicily, and Cyprus). Some researchers believe that all animals have crossed to the islands by swimming or rafting (at least for the elephants, it seems rather unbelievable). Mediterranean Sea has been since millennia a crossroad of different activities and intense navigation. Even now approximately 220,000 merchant vessels of more than 100 tonnes cross the Mediterranean Sea each year – about one third of the world’s total merchant shipping. This is connected also with dispersion of animal species between Mediterranean countries and with pollution of the sea and its shores. Every year between 100,000 and 150,000 t of crude oil are deliberately released into the sea from shipping activities. The climate changes reflect on the Mediterranean climate and on the sea level. Sea level rise for the next century (2100) is expected to be between 30 and 100 cm. A rise of 30 cm would flood 200 km2 of the Nile Delta and many other low territories. The sea is entirely within the Mediterranean Subregion of Palearctic and has many bigger islands (Sicily, Sardinia, Corsica, Balearic Islands, Crete, Cyprus, many Greek and Dalmatian islands, and many smaller islands around Italy). The arachnofauna of these islands is relatively well studied and contains many interesting endemics. 350 7 Regional Arachnogeography Map 7.9 Secondary subdivision of the Primary Mediterranean Center (De Lattin 1967) 1. Atlantomediterranean Secondary Center; 2. Adriatomediterranean SC; 3. Pontomediterranean SC; 4. Tyrrhenian SC; 5. Canarian SC; 6. Mauretanian SC; 7. Cyrenaican SC; 8. Cretan SC; 9. Cyprian CS The climate changes reflect on the Mediterranean climate and on the sea level. Sea level rise for the next century (2100) is expected to be between 30 and 100 cm. A rise of 30 cm would flood 200 square kilometers of the Nile Delta and many other low territories. In the world there are several “Mediterranean regions” (Eurafrican Mediterranean, California, Chile, New South Wales in Australia, the Cape area in South Africa). They are similar in their climate and other patterns. Their biogeographic characteristics have been outlined by Vitali-di-Castri (1973). 7.7 7.7.1 Balearic Islands Geography, General Zoogeography, and Paleogeography An archipelago in the Western Mediterranean was formed by four largest islands (Mallorca, 3640 km2; Menorca, 696 km2, highest point 358 m; Ibiza, 572 km2, highest point 475 m; and Formentera, 83 km2) and several smaller islands (Cabrera, etc.) The last two larger islands form a group, called Pityusas. The highest point of the archipelago is on Mallorca (1445 m in the Serra de Tramuntana) (Colom 1957, Gautier et al. 1994, Pons and Palmer, 1996). 7.7 Balearic Islands 351 During the Last Glacial Maximum, the Emile Baudot Escarpment and the two submarine mounts, dels Oliva and Ausiàs Marc, formed a microarchipelago between the “Pitiusa” and the “Gran Balear” (Mateu et al. 2004). 7.7.2 Arachnogeography In their review of the endemic animals of Balearic Islands, Pons and Palmer (1996) enumerate 1 species of scorpions (a valid species Euscorpius balearicus Caporiacco, 1950, not subspecies of “carpathicus”), 1 sp. of Palpigradi, 13 spp. or ssp. of Pseudoscorpiones, 4 of Opiliones, 32 spp. of spiders, and 5 spp. of Acari. To them must be added Anelasmocephalus balearicus Martens et Chemini, 1988 (missed), and Trogulus balearicus Schönhofer et Martens, 2008 (described later). Palpigradi Peyerimhoff (1906) described from Mallorca one of the first known Palpigradi – Eukoenenia draco draco. Another subspecies (E. draco zariquieyi Condé, 1951) has been described from Catalonia. Scorpiones The Balearic scorpion was defined as valid autochthonous and endemic species Euscorpius balearicus Caporiacco, 1950 (Mallorca, Menorca, Cabrera, Dragonera) (Caporiacco 1950, Gantenbeim et al. 2001). Pseudoscorpiones On the islands are registered pseudoscorpions of 18 genera and the families Chthoniidae, Neobisiidae, Syarinidae, Garypidae, Geogarypidae, Atemnidae, Olpiidae, Cheliferidae, Chernetidae, and Withiidae (Beier 1959, 1961, Estany 1977, Lagar 1972, Mahnert 1977, 1978b, 1989, 1993, Tullgren 1900, Zaragoza 2000, Zaragoza and Vadell 2013, Pons and Palmer 1996). Endemics are: Chthonius campaneti Zaragoza et Vadell, 2013 (Mallorca, cave) Chthonius (Ephippiochthonius) cabreriensis Mahnert, 1993 (Cabrera, Ila del Connils) Ch. (E.) ponsi Mahnert, 1993 (Cabrera) Ch. (E.) balearicus Mahnert, 1977 (Mallorca, troglophile) Ch. (E.) bellesi Mahnert, 1989 (Menorca, troglobite) Acanthocreagris balearica Beier, 1961 (Menorca) Neobisium monasterii Mahnert, 1977 (Mallorca) N. ischyrum balearicum Beier, 1939 (Menorca, Eivissa, Cabrera) Roncus neotropicus Redikorzev, 1937 (?= R. balearicus Beier, 1961) (Mallorca, Eivissa) R. vidali Lagar, 1972 (Mallorca, troglobite) Allochernes pityusensis Beier, 1961 (Eivissa) Pselaphochernes balearicus Beier, 1961 (Mallorca) Dactylochelifer balearicus Beier, 1961 (Menorca, Eivissa) 352 7 Regional Arachnogeography D. besucheti Mahnert, 1976 (Mallorca) Opiliones On the Balearic Islands are known nine genera of Opiliones (two Laniatores and seven “Palpatores”) of eight genera (Ptychosoma, Scotolemon, Trogulus, Anelasmocephalus, Homalenotus, Odiellus, Phalangium, Paropilio, Leiobunum) and the families Phalangodidae, Trogulidae, Nemastomatidae, Phalangiidae, and Sclerosomatidae (Martens and Chemini 1988; Prieto 2003; Rambla 1972, 1977a, 1977b, 1979; Schönhofer and Martens 2008). Eupnoi (endemics) Fam. Phalangiidae Phalangium clavipus Roewer, 1911 – Mallorca Eudasylobus ferrugineus (Thorell, 1876) – Mallorca and Ibiza Metaphalangium abstrusum (L. Koch, 1882) – Baleares Dyspnoi (endemics) Fam. Trogulidae Anelasmocephalus balearicus Martens et Chemini, 1988 – Mallorca Trogulus balearicus Schönhofer et Martens, 2008 – Ibiza Laniatores Endemic (and only) Laniatores (Phalangodidae) are Scotolemon krausi Rambla, 1972, on Ibiza and Ptychosoma balearicum Rambla, 1977, on Mallorca (cave). Araneae According to the Iberian spider checklist of Cardoso and Morano (2010), there are 185 species of 122 genera and 37 families of spiders known from Balearic Islands (Cardoso and Morano 2010, Melic 2001, Morano 2004). The most numerous families are Linyphiidae (18 gen., 21 spp.), Salticidae (14 gen., 19 spp.), Araneidae (13 gen., 20 spp.), and Theridiidae (13 gen., 26 spp.). The endemic genus is Chatzakia Lissner et Bosmans, 2016 (Gnaphosidae) 7.8 7.8.1 Sicily, Sardinia, Corsica, and Elba Geography, General Zoogeography, and Paleogeography The three big islands Sicily, Sardinia, and Corsica and the smaller nearby islands of the Tuscan Archipelago (Elba, 223.5 km2, Montecristo, Carpalia, Pianos, etc.) as well as the other small archipelagoes (Aeolian and Pontine islands) are important as parts of the former microplate, bridge with North Africa (of Sicily) and large enough area of speciation (Shu 1974, Shu et al. 1973, Shu et al. 1977, La Greco 1957, 1961). Their mountains are high (Volcano Etna in Sicily, 3329 m; Monte Cinto in Corsica, 2706 m; and Punta La Marmora, 1834 m in Sardinia). 7.8 Sicily, Sardinia, Corsica, and Elba 353 Corsica was formed approximately 250 million years ago with the uplift of a granite backbone on the western side. It is the most mountainous island in the Mediterranean, a “mountain in the sea” (with 20 summits of more than 2000 m). Approximately 3500 km2 of the total surface area of 8680 km2 is dedicated to nature reserves (Parc Naturel Régional de Corse). The island is 90 km from Italy and 170 km from the Côte d’Azur in France. It is separated from Sardinia to the south by the Strait of Bonifacio, a minimum of 11 km wide. Sardinia is the second largest island in the Mediterranean Sea, with an area of 23,821 km2, with Gennargentu Ranges at the center of the island. Sicily is separated from the Italian region of Calabria by the Strait of Messina, about 3 km wide in the north and about 250 m deep. The total area of the island is 25,711 km2, while the autonomous region of Sicily (which includes smaller surrounding islands) has an area of 27,708 km2. Along the northern coast, mountain ranges of Madonie (2000 m), Nebrodi (1800 m), and Peloritani (1300 m) represent an extension of Apennines (Fig. 7.1). Sicily and its small surrounding islands have some highly active volcanoes. Etna covers an area of 1190 km2. The Aeolian Islands in the Tyrrhenian Sea, to the northeast of mainland Sicily, exhibit a volcanic complex including Stromboli. Currently active also are the three volcanoes of Vulcano, Vulcanello, and Lipari, usually dormant. The small islands in the Strait of Sicily (Lampedusa, Linosa, and Pantelleria) have been described in the paper of Massa (Ed.). The only island connected with Africa back to 18,000 years ago is Lampedusa (20.2 km2). Fig. 7.1 Mount Etna rising over suburbs of Catania (Wikipedia) 354 7.8.2 7 Regional Arachnogeography Arachnogeography On these islands are recorded 11 orders and 8 suborders of Arachnida. Lacking are the orders Ricinulei, Amblypygi, Schizomida, Thelyphonida, and Holothyrida and the suborders Palaeoamblypygi and Mesothelae. Only in Sicily are represented Solifugae (Biton Karsch, Daesiidae), and there are Opilioacarida and some scorpions. Palpigradi Rémy (1949) reported from Corsica Eukoenenia mirabilis and E. berlesei. Eukoenenia patrizii Condé, 1956 (endemic), was described from Sardinia. The first known species of the order E. mirabilis was described from Sicily by Grassi and Calandruccio (1885) and was recorded from Sardinia by Roewer (1953). From Sardinia Condé ьха Heurtault (1993) described a second troglobitic (endemic?) species – E. grafittii. As a whole, six species of Palpigradi have been recorded from seven islands of the Mediterranean: Sicily, Sardinia, Mallorca, Iraklia nr. Naxos, Kythira, Corfu, and Malta (Bertrand 1980, Condé 1956, 1987). Solifugae Only on Sicily are known two species of Daesiidae (Biton ehrenbergi Karsch and B. velox Simon) (Chemini 1995). Scorpiones From Sicily, Sardinia, Corsica, and the smaller archipelagoes around them are recorded (Caporiacco 1950, Fet 2010, Roewer 1953) the following scorpions: Fam. Euscorpiidae Euscorpius (Tetratrichobothrius) flavicaudis (DeGeer, 1778) – Corsica E. (E.) oglasae Caporiacco, 1950 – Montecristo Island (Tuscan Archipelago) (endemic) E. (E.) sicanus (C.L. Koch, 1837) – Sicily (t.t.), Sardinia E. (E.) “carpathicus” corsicanus Caporiacco, 1950 – Corsica (probably separate species) Pseudoscorpiones From Sardinia are known 72 sure spp. and ssp. of pseudoscorpions of 25 genera and 11 families (Beier 1955, 1956, 1956, 1959, 1973, Lazzeroni 1969, Callaini 1983a, 1983b, 1989, Gardini 1981, 1994, 2000, von Helversen 1968). From Sicily are known 45 sure spp. and ssp. of pseudoscorpions of 18 genera and 8 families (Beier 1961b, 1975; Callaini 1981, 1991; Gardini and Rizzerio 1987). From Corsica are known 37 sure spp. and ssp. of pseudoscorpions of 18 genera and 8 families (Beier 1948; Callaini 1981; Gardini 2000; Heurtault 1975; Mahnert 1978c; Schawaller 1981). Here is the list of the 15 endemic pseudoscorpions known from the three islands: Fam. Chthoniidae Chthonius (Ephippiochthonius) corsicus Callaini, 1981 – Corsica 7.8 Sicily, Sardinia, Corsica, and Elba 355 Ch. (E.) aegatensis Calaini, 1981 – Sicily Ch. (E.) giustii Callaini, 1981 – Corsica Ch. (E.) grafittii Gardini, 1981 – Sardinia Ch. (E.) poseidonis Gardini, 1981 – Sardinia Ch. (E.) remyi Heurtault, 1975 – Corsica Ch. (E.) siscoensis Heurtault, 1975 – Corsica Ch. (Ch.) multidentatus Beier, 1963 – Sicily Spelyngochthonius sardous Beier, 1955 – Sardinia S. grafittii Gardini, 1994 – Sardinia S. beieri Gardini, 1994 – Sardinia ? Troglochthonius doratodactylus Helversen, 1968 – published from Sardinia but, according to Martens and Helversen (1972) and Harvey (1990), mislabeled Fam. Neobisiidae Acanthocreagris aelleni Mahnert, 1978 – Corsica A. corsa Mahnert, 1978 – Corsica Neobisium mahnerti major Callaini, 1981 – Corsica N. (N.) pacei Callaini, 1991 – Sicily Roncus aetnensis Gardini et Rizzerio, 1987 – Sicily R. carusoi Gardini et Rizzerio, 1987 – Sicily R. siculus Beier, 1975 – Sicily R. zoiai Gardini et Rizzerio, 1987 – Sardinia Fam. Chernetidae Pselaphochernes scorpioides (Herman, 1804) – Sicily Lasiochernes siculus Beier, 1961 – Sicily Opiliones (Juberthie 1958, Marcellino 1970, 1974, 1975a, 1980, 1983, Roewer 1956, Simon 1872) Cyphophthalmi Two of the three spp. of the genus Parasiro Hansen et Sørensen, 1904 (Sironidae), live in Corsica (the third in Spain and continental France). In the other islands, Parasiro minor was recorded by Brignoli (1968) in Sardinia. Eupnoi Fam. Sclerosomatidae Mastobunus Simon, Metasclerosoma Roewer (Sardinia, Sicily), Nelima Roewer (Sicily), etc. Fam. Phalangiidae Dicranopalpus brevipes Marcellino – Sicily (endemic) D. wiehlei Kraus – Sardinia, Sicily Phalangium L., Metaphalangium Roewer, Opilio Herbst, 356 7 Regional Arachnogeography Dasylobus argentatus (Canestrini) – Corsica, Sardinia D. gestroi (Thorell) – Sardinia Dyspnoi Fam. Trogulidae Anelasmocephalus Simon – Corsica, Sardinia, Sicily (A. pusillus Simon) Calathocratus Simon – Sicily (C. africanus Lucas) Trogulus Latreille – Corsica (T. acuaticus Simon) Fam. Dicranolasmatidae Dicranolasma Sørensen – Sicily (D. wiehlei Kraus, D. soerensenii Thorell) Fam. Nemastomatidae Histricostoma Kratochvíl et Miller – Sicily, Corsica, Sardinia (H. argenteolunulatum Canestrini) Laniatores Fam. Travuniidae Buemarinoa Roewer, 1956 – endemic genus, Sardinia (cave) Fam. Phalangodidae The genus Scotolemon is represented in Corsica (Scotolemon terricola Simon – found also in Algeria and Sardinia) and Sicily and Sardinia (S. doriai Pavesi). Ptychosoma vitellinum Soerensen (Phalangodidae) – Sicily, Sardinia (Brignoli 1968, Marcellino 1970, 1975) Araneae According to the list of Marinu and Verneau (2002, online), in Corsica have been recorded 515 spp. of spiders, belonging to 198 genera and 37 families. The catalogue of spiders in Sardinia (Pantini et al. 2013) includes 43 families, 229 genera, and 495 species (Alicata and Cantarella 2000, Brignoli 1969, 1974, 1978, Gasparo 1999, Kraus 1955, Thermes 1972, Bosmans and Colombo 2015). In their catalogue of Sardinian spiders, Pantini et al. (2013) affirm that “…it is interesting to make a comparison with the fauna of nearby Corsica. In fact, the number of species in the Sardinian-Corsican area totals 700 species and the number present on each island is nearly the same, but actually the similarity between the two faunas is not very high (Sørensen’s index of similarity = 0.63).” Some 319 spp. are shared between the two islands. Among the Sardinian spiders, 10.9% are endemic. Endemic genera: Fam. Dysderidae Holissus Simon, 1882 – Corsica Sardostalita Gasparo, 1999 – Sardinia 7.9 Malta 357 Chorological spectrum of Sardinian araneofauna, after Pantini, Sassu, and Serra (2013) According to Brignoli (1981), “…la région sardo-corse a eu une importance extrême pour l’histoire du peuplement de toute l’Italie, non seulement parce qu’elle a joué le rôle de réfuge pour des espèces de climat tempéré (et peut-être aussi pour les thermophiles), mais aussi parce que, à travers elle (ainsi que à travers la région ligurienne-provençale) la faune ancienne de la Méditérranée occidentale a peuplé la peninsule.” Opilioacarida Opilioacarus italicus (With, 1904) was described from Sicily (Palermo) and is known also from Sardinia (Brignoli 1967). 7.9 7.9.1 Malta Geography, General Zoogeography, and Paleogeography The Maltese archipelago is situated in the center of the Mediterranean, 93 km south of Sicily and 288 km east of Tunisia. It consists of five islands: Malta (highest point Ta’ Dmejrek, 253 m), Gozo, and the small Comino, Cominoto, and Filfla. The whole area of the archipelago is 316 km2. The islands were formed from the high points of a land bridge between Sicily and North Africa that became isolated. The archipelago lies on the edge of the African tectonic plate where it meets the Eurasian plate (Schembri 1993, 2003). According to the WWF, the territory of Malta belongs to the ecoregion of “Mediterranean forests, woodlands, and scrub.” The climate is subtropical Mediterranean. 358 7.9.2 7 Regional Arachnogeography Arachnogeography The small islands with limited number of habitats shelter 1 endemic sp. of Palpigradi, 1 endemic scorpion, 22 pseudoscorpions, 3 suborders of opilions, 74 spp. of spiders, and some mites. Nothing is very sensational in the zoogeography of Arachnida (Schembri 2003). Palpigradi The only species known from Malta is Eukoenenia christiani Condé, 1988, from a cave. Scorpiones According to the list of Kovaøik (1999), in Malta live four species of scorpions, all living also on the European continent: Buthus occitanus, Euscorpius carpathicus, E. italicus, and E. flavicaudis. However, according to Fet (2010), E. carpathicus is confined to Romania, and the Maltese former subspecies E. carpathicus sicanus (C.L. Koch, 1837) is now a full species (Fet et al. 2003), known from Malta, Central and Southern Italy, Sardinia, Sicily, Greece, North Africa, and Madeira. Other sources indicate that in Malta live three species: Buthus occitanus, B. tunetanus (?), and Euscorpius sicanus. After Kritscher (1992) and Schembri (2003), on Maltese islands lives only one scorpion: Euscorpius sicanus. Pseudoscorpiones In Malta have been recorded 22 species of pseudoscorpions, belonging to 17 genera and 9 families (Beier 1973, Gardini and Rizzerio 1987, Mahnert 1975, 1982, Schembri 2003). Endemic species are Chthonius maltensis Mahnert, 1975; C. girgentiensis Mahnert, 1982; and Roncus melitensis Gardini et Rizzerio, 1987. Minniza algerica Beier (known also from Lampedusa) is the connection with North Africa and Chernes siciliensis Beier with Sicily. Opiliones On Maltese islands have been recorded five spp. of Opiliones, but there are known several others which are unpublished (Schembri 2003). Most species are known also from Sicily (Marcellino 1974, Thaler 1996). Eupnoi Fam. Phalangiidae Metaphalangium cirtanum (C.L. Koch) Opilio canestrinii (Thorell) Phalangium targionii (Canestrini) Dyspnoi Fam. Dicranolasmatidae Dicranolasma soerensenii Thorell Laniatores Fam. Phalangodidae Ptychosoma vitellinum Soerensen 7.10 Central and Northern Europe, Great Britain, Ireland, Island, and Faroe Islands 359 Araneae The literature on Maltese spiders was reviewed first by Baldacchino et al. (1993) (74 spp. of 21 families) and then by Bosmans and Dandria (1995) (83 spp. of 22 families). These authors describe several new species, seemingly endemic to Maltese islands. Most species are known also from Italy, and some are SiculoMaltese endemics (Brignoli 1969, Cantarella 1982, Kritscher 1996). Acari None is zoogeographically interesting. 7.10 7.10.1 Central and Northern Europe, Great Britain, Ireland, Island, and Faroe Islands Geography, General Zoogeography, and Paleogeography In this area are included the countries north of Spain, peninsular Italy and the Balkan Peninsula (north of Danube), and east to the borders with Russia, Belarus, and Ukraine. Included are also the British Isles, Island, Faroes, and other northern islands. This is one of the best studied parts of the Earth. Dominated by the Alps and Carpathians, Central Europe has been glaciated, with consequences on its arachnofauna (Hulten 1937). However, it is worth noticing the article of Malicky et al. (1983), advocating the existence of Central European faunal elements (distribution excluding the Mediterranean region and Asia largely or totally). Some of the examples are among the spiders and the opilions (by K. Thaler). The idea of the authors is that part of the fauna has “overwintered” during the Pleistocene glaciations. 7.10.2 Arachnogeography Besides the tropical orders, in Central and Northern Europe lack Solifugae and Opilioacarida, and there are very few scorpions of only one genus (Euscorpius Thorell). In this area the arachnids of the Alps are especially well studied, with many papers on high-altitude spiders, mites, opilions, and pseudoscorpions (Beron 2008b, with extensive bibliography on the high mountain Arachnida; 2016, Brinck 1966, Franz 1954, Freytag 1962, Meyer and Thaler 1995, Komposch 2011, Muster 2000, 2001, Schmölzer 1999, 2001, Strand 1906, Thaler, many papers). Palpigradi So far from mainland Europe (except for the three southern peninsulas) has been recorded eight species of Palpigradi, all belonging to Eukoenenia Börner. They are known from France, Austria, Hungary, and Romania (Christian 1998, Hansen 1926, Peyerimhoff 1902). Scorpiones The natural distribution of scorpions in Europe, north of the Danube, is limited to a line across Romania, Austria, and Czech Republic (?) (Braunwalder 2001, Caporiacco 1950, etc.). According to the concept of Fet and Soleglad (2002), the “widely distributed” before Euscorpius carpathicus (Linnaeus, 1767) is 360 7 Regional Arachnogeography confined to a small area in Southwest Romania. The small population in Slapy (Central Bohemia) is considered sometimes introduced or autochthonous (Kovarik 1999). The distribution of this species, still endemic for Romania, is distributed in two clusters in the Carpathians (Gherghel et al., 2016). According to Fet (2010), in Europe north of the three southern peninsulas are recorded four species of scorpions: Fam. Euscorpiidae Euscorpius (E.) carpathicus (Linnaeus, 1767) – Romania E. (Alpiscorpius) alpha Caporiacco, 1950 – Alps (Italy, Switzerland) E. (A.) germanus (C.L. Koch, 1837) – Austria, Slovenia, Switzerland, NE Italy E. (A.) gamma Caporiacco, 1950 – Austria, Slovenia, NE Italy Pseudoscorpiones In the Central and Northern Europe (the area described above) have been recorded pseudoscorpions of ten families: Chthoniidae, Neobisiidae, Syarinidae, Geogarypidae, Larcidae, Cheiridiidae, Cheliferidae, Atemnidae, Chernetidae, and Withiidae (Beier 1952, 1963, Beier and Franz 1954, Gulièka 1977, Kaisila 1949, Lehtinen 1964, Lohmander 1939, Meinertz 1964, 1962, Rafalski 1967, Uddström and Rinne 2014). From Czech Republic and Slovakia have been recorded 58 spp. of 22 genera and 8 families (Christophoryová et al. 2012). In Austria live 71 spp. of 24 genera and 10 families (Mahnert 2004, actualized), and from Finland are recorded (Uddström and Rinne 2014) 17 spp. of 5 families. Endemic genera of pseudoscorpions in Central and Northern Europe: Fam. Neobisiidae Occitanobisium Heurtault, 1977 – France (one sp.) Roncobisium Vachon, 1967 – France (two spp.) Simonobisium Heurtault, 1974 – France, Italy (one sp.) Anthrenochernes Lohmander, 1939 – Sweden (one sp.) Opiliones In West Europe north of the three southern peninsulas and the Pyrenees and west of Russia and Belarus are registered Opiliones of ten families: Sironidae, Cladonychiidae, Phalangodidae, Phalangiidae, Sclerosomatidae, Nemastomatidae, Trogulidae, Dicranolasmatidae, Ischyropsalididae, and Sabaconidae (Bezdecka 2008, Blick and Komposh, 2004, Cawley 2002, Heinäjoki 1944, Kauri 1980, Komposch 1999, 2011, Komposch and Gruber 2005, Lehtinen 1964, Pack-Beresford 1926, Rafalski 1956, Roewer 1934, Schuster 1975, Spoek 1975, Spungis 2008, Staręga 1976a, Stol 1993, Szalay 1968, Tumšs 1963, Vanhercke 1999, Wijnhoven 2005, Winiarska 2008). According to Stol (2007), a total of 24 Nordic species are currently known, of which 17 are from Norway, 19 from Denmark, 20 from Sweden, 12 from Finland, 5 from the Faroe Islands, and 4 from Iceland. Komposch (2004) lists for Hungary 33 spp. of Opiliones of 22 genera and 6 families. For Austria Komposch (2005) lists 61 (s) spp. of Opiliones of genera and 8 families. Among them there are two endemic (sub)species for Austria, Nemastoma bidentatum relictum Gruber et 7.10 Central and Northern Europe, Great Britain, Ireland, Island, and Faroe Islands 361 Martens and N. schuelleri Gruber et Martens, and nine subendemic species. Again Komposch (2004) lists from Hungary 33 species of 23 genera and 6 families. From the other countries, the figures are as follows: Belgium (Vanhercke L. 1999) – 27 (sub)spp. of Opiliones of four families Netherlands (Spoek 1975; Wijnhoven, 2005) – 27 (sub)spp. of Opiliones of five families Switzerland (Martens 1978) – 49 (sub)spp. of Opiliones of six families Germany (Martens 1978) – 49 (sub)spp. of Opiliones of six families Poland (Rafalski 1960, 1961; Rafalski and Staręga 1997; Staręga 2000; Winiarska 2008) – 36 (sub)spp. of Opiliones of eight families Czech Republic (Šilhavý 1956, Klimeš 2000) – 33 spp. of Opiliones of 15 genera and 5 families Slovakia (Šilhavý 1956, Klimeš 2000) – 33 spp. of Opiliones of 22 genera and 8 families Cyphophthalmi There is only one species (Siro carpaticus Rafalski, 1956), described from Poland, known also from Slovakia. The “Palpatores” form the bulk of the European opiliofauna. In Central Europe are known the families Phalangiidae, Sclerosomatidae, Nemastomatidae, Trogulidae, Dicranolasmatidae, Ischyropsalididae, and Sabaconidae. Eupnoi Fam. Phalangiidae – Phalangium L., Amilenus Martens, Dicranopalpus Doleschal, Lacinius Thorell, Mitopus Thorell, Odiellus Roewer, Oligolophus C.L. Koch, Paroligolophus Lohmander, Egaenus C.L. Koch, Opilio Herbst, Dasylobus Simon, Megabunus Meade, Lophopilio Hadži, Platybunus C.L. Koch Fam. Sclerosomatidae – Leiobunum C.L. Koch, Gyas Simon, Astrobunus Thorell, Nelima Roewer Dyspnoi Fam. Nemastomatidae – Mitostoma Roewer, Nemastoma C.L. Koch, Paranemastoma Redikorzev, Carinostoma Kratochvil, Hystricostoma Kratochvil Fam. Trogulidae – Trogulus Latreille, Anelasmocephalus Simon Fam. Dicranolasmatidae – Dicranolasma Sørensen Fam. Ischyropsalididae – Ischyropsalis C.L. Koch Fam. Sabaconidae – Sabacon Simon Laniatores Roewer (1935) made a review of the European Laniatores. The species of the three southern peninsulas and the Mediterranean islands are excluded; in Central Europe are known to live Laniatores from the genera Arbasus Roewer (Travuniidae), Scotolemon Lucas (Phalangodidae). From the family Cladonychiidae, one genus (Holoscotolemon Roewer, 1915) is represented with nine species in Austria, Romania, Hungary, France, and Italy. The other seven species of four genera live in the USA. Endemic opilionid genera for Central Europe are: Fam. Cladonychiidae Holoscotolemon Roewer, 1915 362 7 Regional Arachnogeography Araneae The spider fauna of Central and Northern Europe (within the described limits) is well known and represented by ca. 37 families, after Aakra and Hange (2000), Agnarsson (1996), Almquist (2005), Blick et al. (2004), Bosmans (2009), Bosmans and Vanytven (2001, Internet), Buchar (1992), Buchar et al. (1995), Buchar and Ružicka (2002), Eskov (1994), Gajdos et al. (1999), Hauge (1989), Komposch (2011), Koponen (1995, 1996, 2005), Kronestedt (Version 2001), Larsen and Scharff (2003), Marinu and Verneau (2002), Merrett et al. (1985), Merrett and Millidge (1992), Merrett and Murphy (2000), Mikhailov (1997, 1998, 1999, 2000), Milošević (2002), Roberts (19895), Thaler (1976, 1980, 1988), Thaler and Buchar (1994, 1996), Nentwig et al. (2013), Newlands (1978), Platnick (2014), Proszynski and Staręga (1971), Samu, Szinetar (1999), Vilkas (1992), Weiss and Urak (2000), and Wiehle (1953): Scytodidae, Sicariidae, Pholcidae, Dysderidae, Oonopidae, Segestriidae, Eresidae, Oecobiidae, Palpimanidae, Uloboridae, Anapidae, Araneidae, Linyphiidae, ?Synaphridae, Tetragnathidae, Theridiidae, Theridiosomatidae, Lycosidae, Oxyopidae, Pisauridae, Miturgidae (Zorinae), Agelenidae, Amaurobiidae, Anyphaenidae, Cybaeidae, Dictynidae, Hahniidae, Sparassidae, Zodariidae, Clubionidae, Titanoecidae, Gnaphosidae, Philodromidae, Thomisidae, Salticidae, Corinnidae, and Liocranidae. This number is not very big in this well-studied territory (for comparison, only on the Balkan Peninsula are registered 48 families of spiders). No family is endemic, and many of them are widespread in the world. In Great Britain are known 658 spp. of spiders (Duffey 2010), in France 1569 spp. (Le Peru 2007), in Switzerland 875 spp. (Maurer and Hänggi 1990), in Germany 925 spp. (Koponen 1991), in Poland 809 spp. (Rozwalka and Stanska 2008), in Sweden 704 spp. (Kronestedt 2001), and in Norway 535 spp. (Hauge 1989). Ireland shares all its spider species (377 names) with Great Britain but possesses only 59.6% of that fauna (van Helsdingen 1995). What concerns the Czech Republic, according to Buchar (1995), is that it “may be divided in two zoogeographically and geomorphologically distinct territories – Moravia and Silesia form division between the Bohemian Highlands and Carpathian depressions. The Basin of Vienna extends into Moravian territory from south and builds a way for the immigration of Mediterranean and sub Mediterranean termitophilous species. 761 species of spiders have been collected – 61 species of them are found in Moravia only 70% of those Moravian spiders occur the very xerotherm habitats.” A special case is Iceland, situated between Europe and North America. In this northern country are known the orders Opiliones, Pseudoscorpiones, Araneae, Ixodida, Mesostigmata, Trombidiformes, and Sarcoptiformes. Nine orders are missing. What concerns the spiders, Braendegård (1958) recorded 78 species of 9 families and confirmed the opinion of Lindroth (1931) that “the Icelandic insect [and spider] fauna was almost exclusively of palaearctic origine” and that “the boundary between the nearctic and the palaearctic geographical respect must be the Danmarkstraede.” From the 78 spp., 54 belong to the family Linyphiidae, containing more than half of all Arctic spiders, according to the same author. Acariformes Ref.: Beron (2008b, 2016, Luxton 1995) Prostigmata Fam. Teneriffiidae – the genus Mesoteneriffia Irk is known from the Alps. 7.11 Arachnida of the Atlantic Islands (Macaronesia, St. Helena, Ascension) 363 Oribatida Luxton (1995) published checklist of 135 genera and 303 spp. of Oribatida in the British Isles, with zoogeographical notes on their distribution. 7.11 Arachnida of the Atlantic Islands (Macaronesia, St. Helena, Ascension) 7.11.1 Canary Islands, Azores, and Madeira 7.11.1.1 Geography, General Zoogeography, and Paleogeography Four archipelagoes in the Atlantic Ocean (Canary Islands, Azores, Madeira, Cabo Verde) are called Macaronesia. The northernmost of them are the Azores. The nine volcanic islands of the archipelago have a total surface area of 2346 km2, and the highest peak is Mount Pico (2351 m). During the last several centuries, the forests (laurisilva) have been destroyed (about 95% of them), and it is believed that more than half of the insects and other invertebrates having existed originally (until the discovery of the uninhabited islands in fourteenth century) are gone or heavily endangered. The Azores are situated 1360 km west of Portugal, about 1510 km northwest of Morocco, and about 1925 km southeast of Newfoundland. Another archipelago, a Portugal territory, is Madeira, comprising the islands of Madeira (741 km2), Porto Santo, Desertas, and Selvagens, 520 km from the African coast. The archipelago of the Canary Islands consists of seven main islands, located between 100 and 500 km from the African coast. They are the Tenerife (2034 km2), Fuerteventura (1659 km2), Gran Canaria (1560 km2), El Hierro (278 km2), Palma (706 km2), Lanzarote (846 km2), and Gomera (370 km2). Pico de Teide on Tenerife is the highest mountain of all Atlantic Ocean islands (3718 m). All islands emerged in Miocene, Fuerteventura and Lanzarote being the oldest (Dietz and Sproll 1970b). For Macaronesia s. str. (Canary Islands, Azores, Madeira, Selvagens), we find on the EEA (2002) analysis of the geographic regions the following distribution of land: meadows 25%, forest 15%, cultivated land 14%, and land with few or no vegetation 34%. 7.11.1.2 Arachnogeography In 2001 was published a volume containing database on the terrestrial animals of the Canary Islands. The Arachnida are distributed as follows (Bacallado 1984, Schenkel 1938, Kunkel (Ed.) (1976)): Palpigradi – one sp. (omitted) Araneae – 453 spp. (292 endemic) Pseudoscorpiones – 44 (23 endemic) (according to Mahnert (2011), on the islands are known 56 spp. and subspp. of pseudoscorpions) Scorpiones – one sp. (introduced) 364 7 Regional Arachnogeography Opiliones – five spp. Palpigradi – one sp. Schizomida – one sp. (introduced) Solifugae – one sp. (endemic) Acari – 244 spp. Total – 799 spp. and 23 ssp., 393 spp. and 11 ssp. endemic (actually, many more than 800 spp.). Especially the Acari are certainly many more. Lacking from the islands are the orders Ricinulei, Uropygi, Opilioacarida, Holothyrida, and non-introduced Scorpiones and Schizomida. Azorean Islands – Araneae, Opiliones, Pseudoscorpiones, Acari, etc. Madeira – Palpigradi (one endemic sp.), Araneae, Opiliones, Pseudoscorpiones, Acari, etc. Palpigradi Eukoenenia mirabilis (Grassi) was the first Palpigradi, recorded from the Canary Islands (Tenerife) (Condé 1990). From Madeira is known Eukoenenia madeirae Strinati et Condé, 1996. Solifugae The only known species on the Canary Islands is Eusimonia wunderlichi Pieper, 1977 (Karschiidae). The other 14 spp. of Eusimonia are spread from North Africa to Central Asia. From Cabo Verde has been described Ammotrechella diaspora Roewer, 1934 (Ammotrechidae), a purely American genus and family. Scorpiones No scorpions on the Azores. On the Canary Islands have been found several species (incl. Centruroides gracilis), but probably introduced (Fet 2010). Also according to Fet (2010), “Euscorpius sicanus record from Madeira (Fet et al. 2003) could be either introduction from the Mediterranean, or a local relict.” Otherwise, Macaronesia did not seem to harbor native scorpions (Crucitti 2004, Fet et al. 2003, Fet 2010). Exception is Hottentotta caboverdensis (Buthidae) in Cabo Verde. Pseudoscorpiones The distribution of Pseudoscorpiones on the Atlantic Ocean islands is already well known and rich with zoogeographically remarkable taxa (Beier 1961, 1965, 1970, 1975, 1976, Estany 1979, von Helversen 1965, Mahnert 1980, 1989, 1990, 1993, 1997, 2002, 2011, Pieper 1981, 1991, Schenkel 1938, Tullgren 1900, Vachon 1961, Zaragoza, 1986, 2000, full bibliography, Zaragoza et al. 2004). They are distributed as follows: Canary Islands: 48 spp. of 13 genera and 8 families (Chthoniidae, Syarinidae, Geogarypidae, Olpiidae, Cheliferidae, Cheiridiidae, Chernetidae, Withiidae) Azores: two genera and two families (Neobisiidae, Syarinidae) Madeira and Selvagens: 23 spp., 5 genera, and 3 families (Chthoniidae, Syarinidae, Cheliferidae) 7.11 Arachnida of the Atlantic Islands (Macaronesia, St. Helena, Ascension) 365 The distribution of different families and genera is as follows: Fam. Chthoniidae Chthonius C.L. Koch – Canary Islands, Madeira Paraliochthonius Beier – Canary Islands, Madeira Tyrannochthonius Chamberlin – Canary Islands Lagynochthonius Beier – Canary Islands (six spp.) Fam. Geogarypidae Geogarypus Chamberlin – Canary Islands Fam. Olpiidae Calocheirus Chamberlin – Canary Islands (four of the six species in the genus; the remaining two live in Sudan, Israel, and Saudi Arabia) Fam. Syarinidae Microcreagrina Beier – Madeira, Canary Islands (three spp. of Microcreagrina live on the Canary Islands, one on Madeira, and one is widespread in the Mediterranean) Microcreagrella Beier – Madeira, Azores (different subspecies of M. caeca (Simon)) Pseudoblothrus Beier – Azores (two spp.) Fam. Cheiridiidae Cheiridium Menge – Canary Islands Fam. Chernetidae Pselaphochernes Beier – Canary Islands Fam. Cheliferidae Canarichelifer Beier – Canary Islands (endemic genus) Pseudorhacochelifer Beier – Madeira, Canary Islands (different spp.) Rhacochelifer Beier – Canary Islands Fam. Withiidae Withius Kew – Canary Islands Opiliones On the islands are known six genera of Opiliones (Laniatores, Eupnoi) of the families Pyramidopidae and Phalangiidae (Kauri 1963; Rambla 1956, 1975, 1993; Roewer 1936; Sharma et al. 2011; Spoek 1959; Staręga 1984). Four of them are endemic. Endemic on the islands are: 366 7 Regional Arachnogeography Laniatores Fam. Pyramidopidae Maiorerus randoi Rambla, 1993 – Fuerteventura, cave (endemic genus and species) Eupnoi Fam. Phalangiidae (Phalangiinae) Bunochelis Roewer, 1923 (endemic genus) Bunochelis canariana (Strand, 1911) (syn. B. altenai Spoek, 1959 – Tenerife) – Canary Islands B. spinifera (Simon, 1878) – Canary Islands Ramblinus Starega, 1984 (endemic genus) R. spinipalpis (Roewer, 1911) – Madeira Parascleropilio Rambla, 1975 (endemic genus) P. fernandezi Rambla, 1975 – Canary Islands 7.11.1.3 Araneae On the Canaries are known at least 453 spp. (292 endemics) of spiders (Berland 1935a, Denis 1941, Wunderlich 1987, 1991, 1992, 1993, 1995). On Madeira have been recorded 164 species of spiders (Denis 1962, Wunderlich 1987, Cardoso and Crespo 2008), including 40 single-island endemics. Analysis of the spider biodiversity patterns on the Azoreans has been done by Borges and Wunderlich (2008). Some endemic genera of spiders in Macaronesia: Fam. Pisauridae Cladycnis Simon, 1898 – Canaries Fam. Nesticidae Canarionesticus Wunderlich, 1992 – Canaries Fam. Linyphiidae Afribactrus Wunderlich, 1995 – Azores Acorigone Wunderlich, 2008 – Azores Canariellanum Wunderlich, 1987 – Canary Islands (four spp.) Frontiphantes Wunderlich, 1987 – Madeira (one sp.) Lomaita Bryant, 1948 – Hispaniola Fam. Gnaphosidae Canariognapha Wunderlich, 2011 – Canary Islands Macarophaeus Wunderlich, 2011 – Madeira, Canary Islands Fam. Pholcidae Ossinissa Dimitrov et Ribera, 2005 – Canary Islands (one sp.) 7.12 Cabo Verde 367 Fam. Theridiidae Eurypoena Wunderlich, 1992 – Canary Islands (one sp.) Grancanaridion Wunderlich, 2011 – Canary Islands, Gran Canaria (one sp.) Macaridion Wunderlich, 1992 – Madeira (one sp.) 7.12 7.12.1 Cabo Verde Geography, General Zoogeography, and Paleogeography Ten bigger and three smaller volcanic islands are located in the central Atlantic Ocean, 570 kilometers off the coast of western Africa (Senegal, Mauretania). The total area of is 4033 km2 and the highest point is Pico de Fogo (2829 m.), an active volcano. Islands date from eight million (in the west) to 20 million years (in the east). Climate is semidesert. The nature is strongly degraded by humane activities after the discovery of the islands in 1456. The archipelago is part of the Macaronesian ecoregion. 7.12.2 Arachnogeography On the islands are known members of Solifugae, Scorpiones, Pseudoscorpiones, and Araneae. The other orders are not recorded. It is to note that the Solifugae are represented by a member of the American family Ammotrechidae, but the scorpions and pseudoscorpions are of Afrotropical character. Solifugae Only one sp. has been recorded: Ammotrechella diaspora Roewer, 1934 (Ammotrechidae). The other members of genus Ammotrechella live in North and Central America and in West Indies. No Ammotrechidae in the Old World. Scorpiones Lourenço and Ythier (2006) described a new, and supposedly endemic, species for Cabo Verde islands (Hottentotta caboverdensis, Buthidae). Pseudoscorpiones On the archipelago are reported five spp. of pseudoscorpions of the families Atemnidae, Olpiidae, Garypidae, and Withiidae. The fauna is a mix of Afrotropical and Mediterranean elements (Ellingsen 1906, Vachon 1956). Araneae Berland (1936), having studied the spiders collected by A. Chevalier in 1934, brought the number of species of spiders to 48 species of 15 families. Now from Cabo Verde are reported 58 endemic species of spiders, including Luxuria lymphatica (jumping spider). Australoechemus Schmidt et Piepho, 1994 (Gnaphosidae); Koinothrix Jocqué, 1981 (Linyphiidae); and Wesolowskana Koçal et Kemal, 2008 (Salticidae) are endemic genera. 368 7.13 7.13.1 7 Regional Arachnogeography St. Helena and Ascension Geography and Paleogeography (General) St. Helena is an island of volcanic origin in the South Atlantic Ocean. It is part of the British overseas territory of St. Helena, Ascension, and Tristan da Cunha. St. Helena measures about 16 by 8 kilometers, with 122 km2 surface area with the highest point Diana’s Peak at 818 m. Ascension has a surface area of 91 km2 and Tristan da Cunha 98 km2. Important contribution to knowledge on the fauna of St. Helena were the four volumes, result of the Belgian mission in 1965. The Arachnida have been analyzed by Benoit (1977 – several papers). Ashmole P. and M.J. Ashmole (1997, 2000) contributed to the protection of the seabirds on Ascension and made some important discoveries of blind cave animals on the island. 7.13.2 Arachnogeography The fauna is well studied. It is not very rich as the islands are far from any mainland and also because of the human impact on the vulnerable environment. Lacking are Palpigradi, Ricinulei, Holothyrida, Opilioacarida, Amblypygi, Thelyphonida (Uropygi), and Schizomida. Better known are the pseudoscorpions and the spiders, with several endemic genera (Ashmole and Ashmole 1997, 2000). Scorpiones Only one cosmotropical species is known on the islands: Isometrus maculatus (de Geer) (Benoit 1977a). Pseudoscorpiones On both islands are represented the families Garypidae, Withiidae, Olpiidae, and Cheiridiidae (Beier 1951, 1961, 1977a, Mahnert 1993). Endemic genera for St. Helena are: Fam. Olpiidae Hemisolinus Beier, 1977 – one sp. Fam. Withiidae Scotowithius Beier, 1977 – one sp. Sphallowithius Beier, 1951 – one sp. From Ascension Island have been recorded five (including endemic) species: Withius ascensionis Beier, 1961; Stenowithius duffeyi Beier, 1961; Garypus titanius Beier, 1961 (the largest pseudoscorpion); and Apocheiridium cavicola Mahnert, 1993 Araneae After Benoit (1977a), from St. Helena are recorded 98 spp. of spiders (48 endemics) of 25 families (Merrett and Ashmole 1997, Millidge and Ashmole 1994, Platnick 1993). 7.13 St. Helena and Ascension 369 Endemic genera for St. Helena: Fam. Dictynidae Helenactyna Benoit, 1977 (two spp.) Fam. Miturgidae Helebiona Benoit, 1977 Fam. Eutichuridae Tecution Benoit, 1977 (three spp.) Fam. Theridiidae Zercidium Benoit, 1977 (one sp.) Fam. Linyphiidae Napometa Benoit, 1977 (two spp.) Fam. Lycosidae Dolocosa Roewer, 1960 Fam. Gnaphosidae Benoitodes Platnick, 1993 (two spp.) Fam. Thomisidae Bonapruncinia Benoit, 1977 (one sp.) Fam. Salticidae Paraheliophanus Clark et Benoit, 1977 (four spp.) Endemic genus and species for Ascension is the blind cave spider Catonetria caeca Millidge et Ashmole, 1994 (Linyphiidae). Acariformes Prostigmata – Ref.: Naudo (1977) Fam. Anystidae – endemic species Chaussieria benoiti Naudo Ch. brevis Naudo Ch. dissimilis Naudo Ch. sanctaehelenae Naudo Oribatida Ref.: Luxton (1995) Parasitiformes Benoit (1977) recorded from St. Helena two spp. of Ixodidae, one of Laelapidae and one of Macronyssidae (all parasitic). 370 7.14 7.14.1 7 Regional Arachnogeography Russia (North of Caucasus), Belarus, Ukraine, Siberia, Altai, China (North of Yangtze), and Mongolia Geography, General Zoogeography, and Paleogeography Ukraine, Belarus, and Russia, north of Caucasus, and of Central Asian mountains are vast plains and taiga forest. The valley of Yenisei River is considered to be the important frontier between eastern and western Siberia (Johannsen 1955). The Johannsen’s Line starts from Severnaya Zemlya Islands and Taymyr Peninsula, follows the east bank of Yenisei River, and reaches Altai. Another frontier is the Reinig’s Line along the east bank of Lena River and Aldan River and over the mountains of Stanovoy and Yablonovy into Tian Shan. The paleogeography of the glaciation of Northeast Asia was outlined by Glushkova (1992), the zoogeography of the Far East of Russia by Kolosov (1980), and of North and Central Asia by Serge (1993). Map 7.10 The Johannsen’s Line (oblique striated, determined by recent ecological factors) and Reinig’s Line (vertically striated, determined by historical factors) (from De Latin 1967) 7.14 Russia (North of Caucasus), Belarus, Ukraine, Siberia, Altai, China (North… 7.14.2 371 Arachnogeography From this vast area are missing the orders Amblypygi, Schizomida, Ricinulei, Opilioacarida, and Holothyrida. Thelyphonida (Uropygi) are found only on the very far east of China (the only record of Uropygi in the Maritime Province of Russia is considered a chance specimen brought by Man from China – Mikhailov, 2016a). The northern limits of several groups (Uropygi, Scorpiones, Solifugae) are running through this area (Ukraine, Siberia, Mongolia, China). There are several very useful reviews and checklists of the spiders of Russia and the former Soviet Union (FSU), done by Russian specialists, mainly by K.G. Mikhailov, who was kind enough to keep me informed about the progress of the arachnology of Northern Asia. Solifugae Only one species is known in European Russia and Ukraine: Galeodes araneoides (Pallas 1772) (Galeodidae) (Birula 1912, 1917, 1922). In Central Asia, Altai, North China, and Mongolia are known 26 spp. of Karschia Walter (Karschiidae) (Gromov 2004). They live in Turkmenistan, Uzbekistan, Tajikistan, Kyrgyzstan, Kazakhstan, China, Iran, Turkey, Armenia, and Azerbaijan. In Mongolia are recorded five species (Galeodes kozlovi Birula, 1911, and G. mongolicus Roewer, 1934 (from family Galeodidae), and Karschia gobiensis Gromov, 2004; K. mongolica Roewer, 1933; and Eusimonia turkestana Kraepelin, 1899 (from family Karschiidae)), marking part of the northern limit of the order in Asia. As a whole, from the countries of the former USSR are known 18 spp. of Solifugae (Mikhailov 2016), mostly from Transcaucasia and Central Asia. Thelyphonida (Uropygi) One species (Typopeltis amurensis Tarnani, 1889) was described from the Far East of Russia (Primorsky Krai, Olga Bay, acc. to Harvey, online, nomen dubium, acc. to Mikhailov 2016a, chance specimen brought with human transport from China). Scorpiones According to the catalogue of Fet (1988, with suppl.) and Mikhailov (2016a), in the present territory of Russia (including Crimea) are known five spp. of scorpions: three from fam. Euscorpiidae – Euscorpius tauricus (C.L. Koch, 1837), endemic for Crimea (see Fet, 2003); E. italicus (Herbst), east shore of Black Sea, north of Georgia; and E. mingrelicus (Kessler, 1874) – and two from fam. Buthidae, Mesobuthus caucasicus (Nordmann, 1840), west shore of Caspian Sea (Birula 1917, Nenilin and Fet 1992), and M. eupeus (C.L. Koch, 1839). From Mongolia are known three species: Mesobuthus martensii (Karsch, 1879), M. eupeus (C.L. Koch, 1839), and M. caucasicus (Nordmann, 1840) Pseudoscorpiones In the area are recorded pseudoscorpions of ca. 20 genera and the families Chthoniidae (Chthonius, Mundochthonius), Neobisiidae (Neobisium, Roncus, Bisetocreagris, Microcreagris, Stenohya), Syarinidae, Geogarypidae, Atemnidae (Atemnus, Diplotemnus), Olpiidae, Cheiridiidae (Cheiridium), Chernetidae (Megachernes, Wyochernes, Chernes, Allochernes, Pselaphochernes, 372 7 Regional Arachnogeography Dendrochernes), Cheliferidae (Centrochelifer, Chelifer, Dactylochelifer), and Withiidae (Withius) (Beier 1966, 1969, 1973, 1979, Ćurčić 1983, 1985, Krumpál and Kiefer 1982, Redikorzev 1922, 1934, 1949, Schawaller 1985, 1986, 1989). From Mongolia have been recorded 28 spp. of 14 genera and the families Neobisiidae, Atemnidae, Cheliferidae, Chernetidae, and Withiidae. The most widespread is Dactylochelifer Beier (seven spp.). From the huge territory of Russia (17045 million km2) are recorded only 48 spp. (Mikhailov 2016a). From South Korea are known 21 spp. of 11 genera and the families Chthoniidae, Pseudotyrannochthoniidae, Neobisiidae, Syarinidae, Garypidae, Cheiridiidae, and Chernetidae. There are no information about North Korea (material has been collected, but not yet identified). Some endemic genera: Gobichernes Krumpál et Kiefer – Mongolia Opiliones In this large territory are known harvestmen of more than 30 genera (Chevrisov 1979, Chemeris 2000, Chemeris and Logunov 2001, Chemeris et al. 1979, 1980, Farzalieva and Esyunin 2000, Gritsenko 1979b, 1980, Kratochvil 1959, Martens 1989, Šilhavý 1966, Staręga 1964, 1965, 1978, Suzuki 1941c, Tchemeris et al. 1998, Tsurusaki 1998, Tsurusaki and Crawford 2001, Tsurusaki et al. 2000) and the families Phalangiidae, Sclerosomatidae, Sabaconidae, Taracidae, Trogulidae, Nemastomatidae, and Ischyropsalididae. One sp. of Holoscotolemon (Cladonychiidae, Laniatores) is known from the Ukrainian Carpathians (Chevrisov 1979). Most others are genera of the former “Palpatores.” Cyphophthalmi are not known from the area. Eupnoi Fam. Phalangiidae – Oligolophus C.L. Koch, Mitopus Thorell, Acanthomegabunus Tsurusaki et al., Rilaena Šilhavy, Phalangium L., Zachaeus C.L. Koch, Homolophus Banks, Opilio Herbst, Odiellus Roewer, Metaplatybunus Roewer, Lophopilio Hadzi, Platybunus C.L. Koch, Egaenus C.L. Koch, Tchapinius Roewer (Kamchatka) Fam. Sclerosomatidae – Metagagrella Roewer, Psathyropus L. Koch (Far East), Leiobunum C.L. Koch, Gyas Simon, Nelima Roewer Dyspnoi Fam. Sabaconidae – Sabacon Simon (two spp. – S. crassipalpis (L. Koch), Siberia; S. sergeidedicatus Martens, Altai; S. okadai Suzuki, Mandjuria Fam. Taracidae – Taracus Simon (one sp. T. birsteini Ljovuschkin, 1971 from the Far East, cave; the other seven spp. live in the USA) Fam. Trogulidae – Trogulus Latreille (one sp.), Dicranolasma Sørensen, Platybessobius Roewer Fam. Nemastomatidae – Paranemastoma Redikorzev, Nemastoma C.L. Koch Fam. Ischyropsalididae – Ischyropsalis C.L. Koch Laniatores Fam. Cladonychiidae 7.14 Russia (North of Caucasus), Belarus, Ukraine, Siberia, Altai, China (North… 373 Holoscotolemon Roewer – Ukrainian Carpathians, H. jaqueti (Corti, 1905) Araneae After studying of 505 spp. of spiders in Northern Siberia, Eskov (1986a, thesis) concludes that the Arctic arachnofauna contains 98 spp. (more than 30% are limited to the northern parts of the tundra zone). The Yenisei Line, running on the end of Middle Siberian plateau, is a very clear biogeographical frontier within Holarctic (a superposition of eastern limits of areas of the European-Western Siberian species and the western limits of areals of the East Siberian-American species). After the distribution of spiders in Northern Holarctic are delimitated three base regions, European, Angaran, and Canadian, and two transitional regions, West SiberianLaplandian (from Enisey to Stegman’s Line, connecting Altai with Fennoscandia) and Beringian (from Kolima to Mackenzie). ) (Eskov 1986b, 1994, Izmailova 1989, Marusik et al. 1995, Marusik et al. 1992, Marusik et al. 1993, Marusik et al. 1993, Marusik and Kovblyuk 2011, Marusik et al. 2000). The araneid fauna of the former Soviet Union has been summarized in the catalogues of Mikhailov (1996, 1997, 1998, 1999, 2000, 2002, 2013). By December 31, 2015, 3374 and 2397 spider species have been reported from the FSU territories and Russia, respectively (Mikhailov 2016b). From Ukraine are known 1016 spp. (end 2015). From the relatively small territory of Tuva (170,000 km2), Marusik et al. (2000) recorded 614 spp. (36% of them in Linyphiidae). From Ural are known ca. 800 spp. The monograph of Song et al. (1999) enumerates for the entire vast territory of China (including Taiwan) 2361 spp. of spiders of 450 genera and 56 families (almost half of the total number of 114 spider families). Two of the families (Dipluridae and Dysderidae) have not been recorded for continental China (only from Taiwan). From the families of Chinese spiders, most genera (over 20) have Linyphiidae (100), Salticidae (71), Araneidae (37), Thomisidae (29), Theridiidae (27), and Gnaphosidae (26). Since 1999 have been published many new papers (mostly by Chinese specialists) and added many new genera and species to the list. Meng et al. (2008) outlined the biogeographical patterns of Chinese spiders, using the parsimony analysis of endemicity. They provided slightly different figures for the families (58, Zoridae included) and species (2858) of the Chinese spiders. Most genera have Linyphiidae (112), Salticidae (88), Araneidae (41), Thomisidae (30), Theridiidae (36), Thomisidae (30), and Gnaphosidae (30). These figures certainly have been changed since 2008. Meng et al. (2008) indicate that “Traditionally [based on vertebrates], China has been divided into two realms and seven biological provinces. The Palearctic realm includes the northeastern, northern, Inner-MongoliaXinjiang, and Qinghai-Tibetan biological provinces. The Oriental realm includes the Southwestern, Central, and Southern biological provinces. This division, originally established by Cheng & Zhang (1959), was later modified by Zhang & Zhao (1978) and Zhang (1998, 2004).” No endemic families. Some endemic genera of spiders in the described area: Fam. Oonopidae Spinestis Saaristo et Marusik, 2009 – Ukraine (one sp.) 374 7 Regional Arachnogeography Fam. Linyphiidae Typhochrestoides Eskov, 1990 – Northern Russia Opilioacarida – missing Acariformes Oribatida The checklist of Chen, Liu, and Wang (2014) indicates that in China (incl. Hong Kong and Taiwan) are known 599 species and subspecies of Oribatida of 275 genera and 97 families (Wang, Wen, and Chen 2002, 2003). Holothyrida – missing 7.15 7.15.1 Sakhalin and Kuril Islands Geography, General Zoogeography, and Paleogeography The island of Sakhalin (72,492 km2) is detached from the Asian continent by the Strait of Tartary (Tartar Strait, 4–20 m deep and 7.5 km wide at the narrowest part, called Nevelskoy Strait) and from Hokkaido by Soya (La Pérouse) Strait. The island is forested, with the highest point Mount Lopatin (1609 m). The Strait of Tartary is freezing often and gives way to migrations by different animals. The Kuril Islands are a chain of 56 islands stretched on 1300 km between Kamchatka and Hokkaido. The highest point is Alaid volcano, 2339 m on Atlasov Island, the archipelago has a surface area of 10,503 km2. Only the southern islands are forested. On the islands there are over 100 volcanoes (40 active), summits of stratovolcanoes. The climate is harsh, with long winters. Some researchers (Sergeev 1993) define Sakhalin (together with Hokkaido) as a province within the Manchurian Subregion of the Palearctic. 7.15.2 Arachnogeography The “southern” orders are not represented in the described chain of islands. The orders reported from this area are Araneae, Opiliones, Pseudoscorpiones, Mesostigmata, Ixodida, Trombidiformes, and Sarcoptiformes. Opiliones Suzuki (1941b, 1956) published papers on the opilions of southern Sakhalin, Kunashir, and Iturup. In another paper, he (Suzuki 1967b) concluded that all five species of opilions on Sakhalin belong to the Holarctic fauna. Between Sakhalin and Hokkaido is situated the Hatta Line. Tsurusaki and Crawford (2001) analyzed the opilions of Kuril Islands and concluded that on the islands are distributed 12 species and that Miyabe Line (= Iturup Strait lying between Iturup and 7.16 Caucasus and Transcaucasia 375 Urup) is of certain significance both for the distribution of the flora and the fauna of the islands. “Along the chain of islands, the number of species per island gradually decreased from Hokkaido toward the northeast: Hokkaido (20 species), Kunashir (10), Iturup (nine), Urup to Ketoi (four, except for Simushir where there are five), and Ushishir to Kharimkotan (two).” According to them, Nipponopsalis yezoensis and Mizozatus flavidus are the most widespread in the islands. The much larger Sakhalin has richer, but underexplored, fauna of Opiliones (Suzuki 1967b, Gricenko 1979), Phalangiidae (Oligolophus thienmuschanensis Wang, Phalangium opilio L.), and others. Araneae In May 2001 in Sapporo there was a Symposium on Kuril Island Biodiversity. Some lectures and posters on spiders by Tanaka, Kamura, Logunov, and Marusik and especially by Marusik and Crawford (2001) outlined the biogeography of this group on the islands. Marusik et al. (1993) published a checklist of spiders from Sakhalin and Kuril Islands. Endemic genera of spiders on the Kuril Islands: Fam. Hahniidae: Pacifantistea Marusik, 2011 7.16 7.16.1 Caucasus and Transcaucasia Geography, General Zoogeography, and Paleogeography The Caucasus Mountains is a mountain system between the Black Sea and the Caspian Sea. It includes the Greater Caucasus Mountain Range and the Lesser Caucasus Mountains. The Greater Caucasus Range extends from the Black Sea nearly to Baku on the Caspian Sea, while the Lesser Caucasus runs parallel to the greater range, at a distance averaging about 100 km south. The Meskheti Range is a part of the Lesser Caucasus system. The Greater and Lesser Caucasus ranges are connected by the Likhi Range, which separates the Kolkhida Lowland from the Kura-Aras Lowland. The Lesser Caucasus and the Armenian Highland constitute the Transcaucasian Highland. The highest peak in the Caucasus range is Mount Elbrus in the Greater Caucasus, which rises to a height of 5642 meters above sea level. Annual precipitation in the Western Caucasus ranges from 1000–4000 mm. The absolute maximum annual precipitation is 4100 mm on the Meskheti Range in Ajaria. Snow cover in several regions (Svaneti and Northern Abkhazia) may reach 5 meters. The Mt. Achishkho region, which is the snowiest place in the Caucasus, often records snow depths of 7 meters. The formation of the present structure of the Caucasus and adjacent areas is connected with the unidirectional compression of this area and the rotation of the compressional axis from a northeastern to a sub-meridional direction 376 7 Regional Arachnogeography in the course of the Cimmerian-Alpine cycle (Gamkrelidze 1986). “The mountainous Crimea and Caucasus Major appeared as islands in the Sarmat Sea 10-5.4 million years ago…; Caucasus Minor, Asia Minor and Balkan compounded the single massif” (Kovblyuk 2004). 7.16.2 Arachnogeography The environment north and south of the Caucasus is quite different, and some group disappear or are strongly reduced north of the mountain (Solifugae, Scorpiones). Some orders and suborders are missing in the area: Amblypygi, Uropygi, Schizomida, Opiliones, Cyphophthalmi, Laniatores, Opilioacarida, and Holothyrida. Palpigradi Christian (2014) described the species Eukoenenia vargovitshi from Abkhazia as an element bridging the apparent gap between Romania and India. Solifugae From Caucasus and Transcaucasia have been recorded five families of Solifugae (Aliev and Gadzhiev 1983, Birula 1936a, 1936b, Morin 1928). Fam. Daesiidae – Bitonissus schelkovnikovi (Birula, 1917) Fam. Karschiidae – Karschia (K.) caucasia (L. Koch, 1878) Fam. Galeodidae – Galeodes araneoides (Pallas, 1772) Fam. Gylippidae – Gylippus (Paragylippus) caucasicus caucasicus Birula, 1907 Fam. Rhagodidae – Rhagodes caucasicus Birula, 1905 Fam. Rhagodidae – Rhagodes caucasicus Birula, 1905 Scorpiones In Caucasus and Transcaucasia are known six spp. of scorpions (Birula 1899, 1917a, Fet 1988, Kinzelbach 1980, etc.) as follows: Fam. Buthidae Androctonus crassicauda (Olivier, 1807) Mesobuthus caucasicus (Nordmann, 1840) M. eupeus (C.L. Koch, 1839) Fam. Euscorpiidae Euscorpius (Polytrichobothrius) italicus (Herbst, 1800) E. (Alpiscorpius) mingrelicus (Kessler, 1874) Fam. Iuridae Calchas nordmanni Birula, 1899 Pseudoscorpiones According to Dashdamirov and Schawaller (1992), in Caucasus have been recorded 66 spp. of pseudoscorpions, belonging to 29 genera and 10 families: Chthoniidae, Neobisiidae, Atemnidae, Cheliferidae, Chernetidae, 7.16 Caucasus and Transcaucasia 377 Geogarypidae, Garypinidae, Olpiidae, Cheiridiidae, and Withiidae (Daday 1889, Redikorzev 1926, 1930, Schawaller 1983, 1994b, Schawaller and Dashdamirov 1988). In Armenia are known 18 spp. of 12 genera and the families Chthoniidae, Neobisiidae, Garypidae, Cheiridiidae, Olpiidae, Cheliferidae, and Chernetidae. In Georgia are known 48 spp. of 16 genera, and 6 families. Remarkable are the two troglobitic Neobisium (Blothrus), described by Lapschoff (1940). In Azerbaijan are known 50 spp. of 26 genera and 10 families (Dashdamirov 1990, actualized). In this paper is done a zoogeographical analysis with 11 zoogeographical complexes. Asia Minor – Caucasian elements are 23.0% of species; endemic Caucasian complex includes six species (17.9%); Palearctic type of areal comprises 15.4%; Mediterranean elements are 10.3%; European complex is 10.3%. Two species are cosmopolitic. Opiliones The opilionids of Caucasus were (and still are) explored by several specialists: Mcheidze (1964), Martens (2006), Snegovaya (1999, 2011), Snegovaya and Chemeris (2005, 2016), Staręga (1966), and others. Cyphophthalmi and Laniatores – not recorded Eupnoi Fam. Phalangiidae – Odiellus Roewer, Lacinius Thorell, Phalangium L., Metaplatybunus Roewer, Opilio Herbst, Rilaena Šilhavý, etc. Fam. Sclerosomatidae – Nelima Roewer Dyspnoi Fam. Trogulidae – Trogulus Latreille, Calathocratus Simon (= Platybessobius Roewer) Fam. Dicranolasmatidae – Dicranolasma Sørensen Fam. Nemastomatidae – Nemastoma C.L. Koch, Paranemastoma Redikorzev, Histricostoma Kratochvil, Mitostoma Roewer, Mediostoma Kratochvil, Nemaspela Šilhavý, Giljarovia Kratochvil, Caucnemastoma Martens, Vestiferum Martens Endemic genera in Caucasus area are Caucnemastoma Martens and Vestiferum Martens. Fam. Gylippidae – Gylippus (Paragylippus) caucasicus caucasicus Birula, 1907 Fam. Rhagodidae – Rhagodes caucasicus Birula, 1905 Scorpiones In Caucasus and Transcaucasia are known six spp. of scorpions (Birula 1899, 1917a, Fet 1988, Kinzelbach 1980, etc.) as follows: Fam. Buthidae Androctonus crassicauda (Olivier, 1807) Mesobuthus caucasicus (Nordmann, 1840) M. eupeus (C.L. Koch, 1839) 378 7 Regional Arachnogeography Fam. Euscorpiidae Euscorpius (Polytrichobothrius) italicus (Herbst, 1800) E. (Alpiscorpius) mingrelicus (Kessler, 1874) Fam. Iuridae Calchas nordmanni Birula, 1899 Pseudoscorpiones According to Dashdamirov and Schawaller (1992), in Caucasus have been recorded 66 sp. of Peudoscorpions, belonging to 29 genera and 10 families: Chthoniidae, Neobisiidae, Atemnidae, Cheliferidae, Chernetidae, Withiidae (Daday 1889, Redikorzev 1926, 1930, Schawaller 1983, 1994b, Schawaller and Dashdamirov 1988). In Armenia are known 18 sp. of 12 genera and the families Chthoniidae, Neobisiidae, Garypidae, Cheiridiidae, Olpiidae, Cheliferidae, Chernetidae. In Georgia are known 48 sp. of 16 gen, and six families. Remakkable are the two troglobitic Neobisium (Blothrus), described by Lapschoff (1940). In Azerbaidjan are known 50 sp. of 26 genera and ten families (Dashdamirov 1990, actualized). In this paper is done a zoogeographical analysis with 11 zoogeographical complexes. Asia-Minor – Caucasian elements are 23.0% of species; endemic Caucasian complex includes six species (17.9%); Palearctic type of areal comprises 15.4%; Mediterranean elements are 10.3%, European complex – 10.3%. Two species are cosmopolitic. Opiliones The opilionids of Caucasus were (and still are) explored by several specialists: Mcheidze (1964), Martens (2006), Snegovaya (1999, 2011), Snegovaya and Chemeris (2005, 2016), Staręga (1966), and others. Cyphophthalmi and Laniatores – not recorded Eupnoi Fam. Phalangiidae – Odiellus Roewer, Lacinius Thorell, Phalangium L., Metaplatybunus Roewer, Opilio Herbst, Rilaena Šilhavý, etc. Fam. Sclerosomatidae – Nelima Roewer Dyspnoi Fam. Trogulidae – Trogulus Latreille, Calathocratus Simon (= Platybessobius Roewer) Fam. Dicranolasmatidae – Dicranolasma Sørensen Fam. Nemastomatidae – Nemastoma C.L. Koch, Paranemastoma Redikorzev, Histricostoma Kratochvil, Mitostoma Roewer, Mediostoma Kratochvil, Nemaspela Šilhavý, Giljarovia Kratochvil, Caucnemastoma Martens, Vestiferum Martens, Endemic genera in Caucasus area are Caucnemastoma Martens and Vestiferum Martens. 7.17 Turkey, Cyprus, Sinai, the Dodecanese Islands, Iran, Afghanistan, the Near East,… 379 Araneae Over 1000 species belonging to 46 families are known from the Caucasus (Buchar and Thaler 1998, Ovcharenko 1978, 1979, Tanasevitch 1987, 1990). The species richest families are as follows: Linyphiidae (~180), Salticidae (122), Gnaphosidae (>100), Lycosidae (>100), Theridiidae (80), Dysderidae (70), and Thomisidae (70). In the different families, endemism values vary from 0 to 100%. The average level of endemism in the Caucasus is about 22%, and the highest level of endemism among species-rich families was found in the Dysderidae, being around 60% (Marusik et al. 2006). According to Mikhailov and Mikhailova (2006), quantitatively Gnaphosidae is the second most abundant family after the Lycosidae. According to Marusik and Guseinov (2003), in Azerbaijan have been found over 600 spp. of spiders of 43–44 families – almost the whole number (51) of the spider families of the former USSR. From Armenia are known 127 spp. and from Georgia 456 spp. (Mikhailov 2002). 7.17 7.17.1 Turkey, Cyprus, Sinai, the Dodecanese Islands, Iran, Afghanistan, the Near East, Middle Asia, and the Arabian Peninsula Geography, General Zoology, and Paleogeography The described area includes Sinai, Asiatic Turkey, Cyprus, the Greek islands near the shore of Asia Minor (Rhodes, Karpathos, Kassos, Kos, Kalymnos, Samos, Chios, Mitilini, and some smaller islands), Iran, Afghanistan, Syria, Lebanon, Israel, Palestine, Jordan, Saudi Arabia, Yemen, Oman, Qatar, UAE, Kuwait, Iraq, Bahrain, Uzbekistan, Tajikistan, Kyrgyzstan, Turkmenistan, and Kazakhstan. The area is mostly dry, with deserts (Iran) and high mountains like Hindu Kush, Tian Shan, and Pamir, some volcanoes, etc. (Turkey, Ararat, 5165 m; Iran, Damavand, 5610 m in Alborz Mt.; Afghanistan, Noshaq, 7453 m, second highest summit of Hindu Kush, after Tirich Mir (7690 m in Pakistan); Kyrgyzstan, Jengish Chokusu (Pik Pobedy), 7439 m, Tian Shan; Tajikistan, Ismoil Somoni Peak in Pamir, 7495 m.). The highest summit of the Arabian Peninsula is Jabal an Nabi Shu’ayb, at 3666 m (in Yemen), and of Lebanon is Qurnat as Sawda’ (3088 m). Some sources: Horowitz (1975), Kosswig (1955), Krumsiek (1976), Kuznetsov (1950, 1957), Kusnezow (1924), Lopatin (1969), Vigna Taglianti et al. (1999) Anatolia is the region where three of the world’s 35 biodiversity hotspots meet, and interact: the Caucasus, Irano-Anatolian, and Mediterranean basin hotspots. One of the most distinctive biogeographic features that helps in understanding the biodiversity of Anatolia is the Anatolian diagonal which has long been recognised as a biogeographic boundary between the central and eastern Anatolian floras and faunas. (Gür 2016) Analyzing the zoogeography of the Levant, Por (1975) reached to the conclusion that the Levant province is a peculiar and complex “subtraction-transition zone” 380 7 Regional Arachnogeography (following the expression of Darlington, 1957). The Levant is “a stretch of land about 150 km wide, wedged in between the [Mediterranean] sea and the SyrioArabian desert, stretching from the mouth of the River Orontes and the Amanus and Taurus mountain ranges in the north, to the Isthmus of Suez in the south” (Por 1975). This “subtraction-transition zone” between the Palearctic and Ethiopian regions, born by the desertification, is called Paleo-eremic region. Usually the biota of the Levant is considered to be Palearctic (Mediterranean), but, according to Por (1975), the inclusion of the Old World deserts in the Holarctic is due to a certain “Europa-centrism” of the scientists. “According to this scheme, the Ethiopis begins only south of the Sahara, in the Savannas, where the last of the typical Palaearctic animals fade out. However, if at all, the connections of the Palaeo-eremic faunal inventory are much closer to the Ethiopis than to Palaearctis….Our [Israel’s] typical desert animals, such as scorpions, agamas, gerbils, sand grouse and gazelles, have Ethiopian rather than Holarctic connections, even if some species are so-called “Mediterraneans” (Por 1975). According to the systems of Bodenheimer (1935) and Por (1975), in the Levant could be distinguished four faunal elements: Palearctic, Paleo-eremic, Ethiopian, and Oriental, with clear prevalence (especially in the north) of the Palearctics in most of the animal groups. The line along the foothills of Northern Galilee and the Golan Heights was called by Por (1975) “Nehring Line.” South of it follows a “transitional zone,” where Palearctic elements mingle with Paleo-eremic elements. More to the south, Por (1975) outlines a “Bodenheimer Line,” which is the end of the transitional zone. Follow Ramon Mountains and the mountains of Sinai. The “Ethiopian element does not prevail in any area. The Oriental species are even more scattered, without showing any geographical pattern. The endemic species among the animals are chiefly of Palaeo-eremic, Ethiopian and also of Oriental origins… this suggests a higher age of the tropical element, and perhaps a younger and more expansive character of the Holarctic species” (Por). In conclusion, “The Levant province is a meeting place and transitional area between the Palaearctic, Oriental and Ethiopian zoogeographic regions.” The broad “Paleo-eremic desert belt serves as a filtering barrier between the three faunal regions.” Turkey is located on cross section of the Boreal, Euro-Siberian, Irano-Turan, and Mediterranean elements from Central Asia, Mesopotamia, Caucasus, and Balkans (Akman 1993). 7.17.2 Arachnogeography In this large, mostly arid, area are known the orders Palpigradi (only three spp.) and Solifugae and many genera and species from the families Galeodidae, Daesiidae, Karschiidae, Gyllipidae, and Rhagodidae, Scorpiones (Pseudochactidae, Akravidae, Buthidae, Euscorpiidae, Hemiscorpiidae, Scorpionidae, Iuridae), Pseudoscorpiones (Atemnidae, Cheiridiidae, Cheliferidae, Chernetidae, Chthoniidae, Garypinidae, Garypidae, Geogarypidae, Menthidae, Lechytiidae, Neobisiidae, Olpiidae, 7.17 Turkey, Cyprus, Sinai, the Dodecanese Islands, Iran, Afghanistan, the Near East,… 381 Withiidae), Opiliones (Sironidae, Phalangiidae, Sclerosomatidae, Trogulidae, Dicranolasmatidae, Nemastomatidae, Phalangodidae), Amblypygi (six spp. – four Charinidae and two Phrynichidae), Araneae (50 families), and Opilioacarida (one sp. of Paracarus from Middle Asia, endemic genus Adenacarus from Yemen). In the described area, there is only one endemic family (Akravidae). Six orders of class Arachnida were recorded from Lebanon (El-Hennawy 2013) in addition to Acari. They are Araneae (38 families, 109 genera, 165 species), Scorpiones (2 families, 10 genera, 12 species), Pseudoscorpiones (7 families, 8 genera, 10 species), Opiliones (3 families, 8 genera, 10 species), Solifugae (1 family, 1 genus, 2 species), and Palpigradi (1 family, 1 genus, 2 species). The total is 52 families, 137 genera, and 201 species. Most of this fauna is clearly Palearctic. The south of the Arabian Peninsula (Yemen, Oman) is usually considered part of Afrotropical Region (Oman also with Oriental elements). Missing from it are the orders Ricinulei, Uropygi, Schizomida, and Holothyrida and the suborders Palaeoamblypygi and Mesothelae (Gromov and Kopdykbaev 1994, Whittick 1941). Palpigradi Almost nothing is known about these tiny and rarely collected animals from the vast and mostly dry area from Turkey and Arabia to Afghanistan (Condé 1965, 1974). The only exceptions are Eukoenenia juberthiei, Condé, 1974, from Lebanon and the strange Leptokoenenia gerlachi, Condé, 1965, from Farasan Island near Saudi Arabia (type of a new genus, found next year also in DR Congo). Solifugae In the vast area of Central Asia and the Near East (Israel, Jordan, Lebanon, Iraq, Kuwait, Syria, Turkey), the Solifugae are widespread. Recorded are the families Galeodidae, Daesiidae, Karschiidae, Gyllipidae, and Rhagodidae (Aliev and Gadzhiev 1983, Birula 1941, Caporiacco 1948, El-Hennawy 1998 2007, Gromov 2004, Lawrence 1953, Prendini et al. 2006, Roewer 1934, Simon 1872, Turk 1948). In Iran are recorded 67 spp. of Solifugae from five families (Kraus 1959; Roewer 1952 1960, with suppl., Harvey 2013b). In Turkmenistan are found 31 spp. (Gromov 1998 and others). In Afghanistan 41 spp. of Solifugae are known so far (Lawrence 1956 and suppl.). In Asia Minor are found 37 spp. of Solifugae: Daesiidae (Blossia Simon, Gnosippus Karsch, Gluviopsilla Roewer, Gluviopsis Roewer, Gluviopsida Roewer, Biton Karsch), Galeodidae (Galeodes Olivier), Gylippidae (Gylippus Simon), Karschiidae (Karschia Simon, Eusimonia Kraepelin, Barussus Roewer), Rhagodidae (Rhagodia Roewer), and Solpugidae (Solpugella Roewer). In some other countries in the area, the number of Solifugae spp. is as follows: Israel (65 spp.), Jordan (5 spp.), Saudi Arabia (14 spp.), Syria (17 spp.), and Lebanon (2 spp.) (Harvey, 2013b). It is clear that this fauna is understudied – the huge number of Solifugae in the tiny Israel is obviously due to the good specialists in this country. In Rhodes (Greece) live four spp. of Solifugae of two fam.: Gluviopsilla discolor Kraepelin, Gluviopsis rufescens Pocock, Gluviella rhodiensis Caporiacco [endemic genus] (Daesiidae), and Galeodes rhodicola Roewer [endemic] (Galeodidae). In Cyprus live five spp. of Solifugae of four families (Blick 2004). They are Biton ehrenbergi Karsch (Daesiidae), Galeodes graecus C.L. Koch (Galeodidae), 382 7 Regional Arachnogeography Gylippus cyprioticus Lawrence, G. syriacus Simon (Gyllipidae), and Eusimonia furcillata Simon (Karschiidae). Only G. cyprioticus is endemic. The number of Solifugae from the republics of Middle Asia is 62 (Mikhailov 2016a, inf. from Gromov in litt.). In total, from the described area, we have the following: Endemic genera: Gluviola Roewer – G. armata Birula, 1905 (Iran) Gluviopsida Roewer – G. taurica Roewer, 1933 (Turkey) Haarlovina Lawrence – H. nielsi Lawrence, 1956 (Afghanistan) Mumaella Harvey – M. robusta Lawrence, 1956 (Afghanistan) Ceratobiton Delle Cave et Simonetta – C. styloceros Kraepelin, 1899 (Israel and Jordan) Gluviella Caporiacco – G. rhodiensis Caporiacco, 1948 (Rhodes) Galeodumus Roewer – G. colognatoi Roewer, 1960 (Afghanistan) Roeweriscus Birula – R. paradoxus Birula, 1937 (Iran) Zombis Simon – Z. pusiola Simon, 1882 (Israel) Acanthogyllipus Birula – A. judaicus Kraepelin, 1899 (Israel) Rhagodelbus Roewer – Rh. bucharicus Birula, 1935 (Uzbekistan) Rhagodax Roewer – Rh. wadidaba Roewer, 194 (Jordan) Rhagodospus Roewer – Rh. babylonicus Birula, 1935 (Iraq) Rhagoduja Roewer – Rh. finnegani Roewer, 1933 (Iran) Rhagodula Roewer – Rh. nigra Roewer, 1941 (Israel) Scorpiones The list of Vachon (1966) covers most of the countries in this sector. In contains 78 spp. of the families Buthidae, Euscorpiidae (to use the modern classification), Scorpionidae (incl. Diplocentridae), and Iuridae. Since this time many amendments and new data occurred, the most striking being the description of a new relict species, genus, and family Pseudochactidae Gromov, 1998. The genus remains endemic, but new genera of the family have been described since from the caves of Laos and Vietnam (Birula 1917, Cruccitti and Cicuzza 2000, Farzanpay 1988, Fet 1985, Fet et al. 2011, Francke 1981, Gantenbein et al. 2000, Gromov 1998, 2001, Habibi 1971, Khazim Al-Asmari et al. 2013, Kaltsas et al. 2008, Kovařik 1993, Levy and Amitai 1980, Lourenço 1996, Lourenço and Vachon 1996, Parmakelis et al. 2006, Vachon 1947, 1951c, 1979a, Warburg et al. 1978). In the review of Vachon and Kinzelbach (1987) to the 78 spp. in 1966 have been added another 28, or in total up to 1987 from the Near East have been recorded 106 spp. of scorpions. Have been added the many species described by Levy et al. (1973) and also by Vachon (1974, 1980) and Francke (1980). Have been described also the new Kraepelinia Vachon, 1974, and Habibiella Vachon, 1974, from Iran. From Turkmenistan Fet (1994) are published six genera (Mesobuthus, Orthochirus, Liobuthus, Anomalobuthus, Pectinibuthus, Kraepelinia) and seven spp. of scorpions, all from fam. Buthidae. According to Fet (1994), all seven spp. belong to the Saharo-Gobian (desert Palearctic) genera. As a whole, from the republics of the former USSR are known 18 spp. of Scorpiones (Mikhailov 2016a, inf. from Fet, in litt.). 7.17 Turkey, Cyprus, Sinai, the Dodecanese Islands, Iran, Afghanistan, the Near East,… 383 Fet et al. (2009) described a new species, Calchas gruberi, from the Greek islands Megisti and Samos and from Anatolia. Kovařík et al. (2010) revised the genus Iurus Thorell (Iuridae), described two new species from Turkey, and concluded that in Turkey live four endemic species of this genus (I. kraepelini, I. kinzelbachi, I. asiaticus, and I. kadleci). This and several earlier papers on Turkish scorpions (Tolunay 1958, Vachon 1947, 1966, Fet and Braunwalder 2000) brought the number of scorpions in this country to 26 spp. of 12 genera from the families Euscorpiidae, Buthidae, and Iuridae. The biogeographic analysis of Crucitti (1999a) says that the Anatolian Peninsula shows its nature of crossroads of different faunistic stocks, as highlighted by its scorpion fauna too. Yağmur et al. (2011) described from Cyprus the new (and most probably endemic) species Buthus kunti (Buthidae). According to the monograph of Levy and Amitai (1980, with suppl.), in Palestine live scorpions of three (modern) families: Buthidae, Scorpionidae, and the recently described from dead specimens family Acravidae. The present-day fauna of Israel contains 19 recorded spp. of 12 genera: Akrav, Androctonus, Birulatus, Buthacus, Buthus, Compsobuthus, Hottentotta, Leiurus, Mesobuthus, Orthochirus, Nebo, and Scorpio. In Jordan Amr and El-Oran (1994) have registered 13 spp. of scorpions of 10 genera and 2 families: Buthidae and Scorpionidae (incl. Diplocentridae). Later this fauna was completed to 20 spp. of the same genera as in Israel, without Akrav. Nebo Simon is considered now (not by all specialists) to belong to Scorpionidae. Extremely interesting is the archaic family Pseudochactidae, from which one species (Pseudochactas ovchinnikovi Gromov, 1998) has been described from Tajikistan and Uzbekistan and another one (P. mischi Soleglad, Kovarik, et Fet, 2012) from Afghanistan. The genus and the two species are endemic from this area. In Oman (Vachon 1977, 1980) were found members of the genera Apistobuthus Finnegan, Butheolus Simon, Buthotus Vachon (Buthidae), and Nebo Simon (Diplocentridae). Now (2017) the fauna of Oman consists of 39 spp., including several endemics, of 16 genera and 3 families. On Rhodes Island live Mesobuthus gibbosus (Brullé) of Buthidae and Iurus dufoureius (Brullé) of Iuridae (Gruber 1963). El-Hennawy (2009) updated the list of scorpions recorded from Saudi Arabia to 23 spp. and 3 subspp., belonging to 3 families: Buthidae (18 spp. of 10 genera), Hemiscorpiidae (1 sp.), and Scorpionidae (4 spp., 3 subspp.). Another survey (Khazim Al-Asmari et al., 2013) provides the following figures: 22 spp., 3 subspp., and 3 families – Buthidae (18 spp. of 10 genera), Hemiscorpiidae (1 or 2 spp.), and Scorpionidae (1 sp., 2 subspp.) (Table 7.1). Endemic genera for this area are: Fam. Pseudochactidae Pseudochactas Gromov, 1998 (Tajikistan, Uzbekistan, Afghanistan) Fam. Buthidae Neohemibuthus Lourenço, 1996 (Iran) Paraorthochirus Lourenço et Vachon, 1995 (Iran) 384 7 Regional Arachnogeography Table 7.1 Scorpiones in Asia Minor, the Near East and Central Asia Country Number of sp. Taxa Fam. Akravidae Akrav Levy Fam. Buthidae Afghanobuthus Lour. Androctonus Ehr. Anomalobuthus Kr. Apistobuthus Finn. Birulatus Vachon Buthacus Birula Buthus Leach Compsobuthus Vach. Hottentotta Birula Iranobuthus Kovarik Isometrus Ehr. Kraepelinia Vachon Leiurus Ehrenberg Liobuthus Birula Mesobuthus Vachon Odontobuthus Vachon Orthochirus Karsch Polisius Fet et al. Psammobuthus Birula Razianus Farzanpay Sassanidiothus Far. Asia Minor Syria Lebanon Jordan Israel Iraq 26 20 10 19 19 20 Afgan. Uzb. Kaz. Kir. Tadz. 29 8 5 2 6 – – + – – – + – – – + – – – + – + 1 + – – – + – – – + 1 – – + – – – + – – – + – – – + – 1 3 3 4 3 1 2 – – – – – – – – – – – 1 1 – – – – – – – – – – – – – – 1 – 1 1 – 1 – – – – – – 2 3 – 5 1 – 1 1 2 5 2 1 4 2 – 2 1 – 3 1 – – – – – – – – – – – 2 1 1 1 1 3 5 – – – 1 – – – – – – – – – – – – – 1 – – – – – – – – – – – – – – – – – – – 1 2 1 2 1 – – – – – – – 4 – 2 – 1 – – – – – 3 – 2 1 2 1 2 – 2 – 2 – – – – – 2 – – – – – 1 – – 1 1 2 9 2 1 – 1 – – – – – – – – – – – – – – – – – – – – – 1 – – – – – – – – – – – – – – – – – 2 – – – – (continued) 7.17 Turkey, Cyprus, Sinai, the Dodecanese Islands, Iran, Afghanistan, the Near East,… 385 Table 7.1 (continued) Country Vachoniolus Levy et al. Fam. Euscorpiidae Euscorpius Thorell Scorpiops Peters Fam. Hemiscorpiidae Hemiscorpius Peters Fam. Scorpionidae Nebo Simon Scorpio L. Fam. Iuridae Calchas Birula Iurus Thorell Neocalchas Yagmur et al. Protoiurus Soled.et al. Country Number of species Taxa Fam. Buthidae Androctonus Ehr. Anomalobuthus Kraepelin Apistobuthus Finn. Babycurus Karsch Buthacus Birula Butheolus Simon Buthus Leach Compsobuthus Vachon Femtobuthus Lowe Hottentotta Birula Asia Minor Syria Lebanon Jordan Israel Iraq – – – – – – Afgan. Uzb. Kaz. Kir. Tadz. – – – – – + – – – – (+) – – – – – 6 – – – – (+) – – – – – – – – – – – – – – – – + – – – – – – – – – – – – – – – 1 – – – – – – + + + + + – – – – – 1 – + 4 1 1 1 1 – – – – 1 1 – – – – 1 1 – – – – – 1 – – – – 1 1 + 2 – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – 3 – – – – – – – – – – Turkm. 7 Saudi Arabia Qatar 29 6 UAE 14 Bahrain 4 Yemen 37 Oman 39 + – 1 + 4 – + 1 – + 1 – + 1 – + 2 – + 2 – – 1 1 1 – 1 1 – – – – – 1 1 – – – – 3 4 – 5 2 – – 1 3 1 – 4 2 – – – – 2 1 6 1 2 – 4 – – – – – – 1 – 2 – 2 – 4 4 (continued) 386 7 Regional Arachnogeography Table 7.1 (continued) Country Kraepelinia Vachon Leiurus Ehrenberg Liobuthus Birula Mesobuthus Vachon Microbuthus Kraep. Odontobuthus Vach. Orthochiroides Kov. Orthochirus Karsch Parabuthus Pocock Pectinibuthus Fet Picobuthus Lowe Vachoniolus Levy et al. Fam. Euscorpiidae Euscorpius Thorell Fam. Hemiscorpiidae Hemiscorpius Peters Fam. Scorpionidae Heteronebo Pocock Nebo Simon Pandinus Thorell Scorpio Linnaeus Turkm. 1 Saudi Arabia Qatar – – UAE – Bahrain – Yemen – Oman – – 4 – – – 3 2 1 1 – – – – – – – – – – – – – – – – – 1 3 – – – – – – 2 – – – – – 2 – 2 2 – – – 1 4 – 1 – – – 1 – 1 – – – – 1 – – – – – 1 – – – – – – – 2 3 – – – – – (+) – – – – – – (+) – – + – + – + + – 1 – 1 – 3 3 – + + – – + + – – – – – 2 – – – – – – 1 – – 1 – – – – – – 4 1 1 3 – – Fam. Akravidae Akrav Levy, 2007 – Israel Amblypygi From the described area are known (Delle Cave 1986, 1989, Delle Cave et al. 2009, Kovařík and Vlasta 1996, Kritscher 1959, Pocock 1895, Weygoldt et al. 2002, Whittick 1941, Miranda et al. 2016) only six species of Amblypygi: 7.17 Turkey, Cyprus, Sinai, the Dodecanese Islands, Iran, Afghanistan, the Near East,… 387 Fam. Charinidae Charinus ioanniticus Kritscher, 1959 – Turkey, Israel, the Dodecanese islands Kos and Rhodes (known also from Egypt) Charinus israelensis Miranda et al., 2016 – Israel Charinus omanensis Delle Cave, Gardner, et Weygoldt, 2009 – Oman (cave) Charinus dhofarensis Weygoldt, Pohl, et Polák, 2002 – Oman [Sarax mediterraneus Delle Cave, 1986 – Kos and Rhodes] – misidentification (Seiter et al., 2015) Fam. Phrynichidae Phrynichus dhofarensis Weygoldt, Pohl, et Polák, 2002 – Oman Ph. jayakari Pocock, 1894 – Hadramaut These are among the species outlining the northern limit of Amblypygi. Actually, Yemen and Oman and its province Dhofar are part of the south of Arabian Peninsula, considered usually as part of the Afrotropical Region. Schizomida From the area described Schizomida occurs only in the Arabian Peninsula (Enigmazomus eruptoclausus Harvey in Oman) (Harvey 2006b). Pseudoscorpiones From the whole of Turkey have been published 102 spp. of pseudoscorpions from 34 genera and the families Chthoniidae, Neobisiidae, Garypinidae, Lechytiidae, Geogarypidae, Cheiridiidae, Atemnidae, Olpiidae, Cheliferidae, Chernetidae, and Withiidae (Beier 1949, 1963, 1965a, 1967, 1969c, 1973d, Harvey 2011, Kunt et al. 2008). From Afghanistan (Beier 1959, 1960, 1961, 1967, 1971, Harvey 2011, 2013, actualized) have been recorded 32 spp. of 22 genera and the families Neobisiidae, Olpiidae, Garypinidae, Ideoroncidae, Atemnidae, Cheliferidae, and Chernetidae. From the Dodecanese and the other islands near Asia Minor have been published pseudoscorpions by Caporiacco (1948), Mahnert, Beier (1962a), and others. From Iraq and Kuwait have been published only three spp. of three genera and the families Cheliferidae and Olpiidae. Pseudoscorpions from Middle Asia have been published by Dashdamirov and Schawaller (1992b, 1993a, 1993b, 1995) and others. In Kazakhstan, Uzbekistan, Turkmenistan, Tajikistan, and Kyrgyzstan are registered scorpions of 30 genera and 10 families: Chthoniidae, Pseudotyrannochthoniidae, Atemnidae, Cheliferidae, Chernetidae, Garypinidae, Ideoroncidae, Neobisiidae, Cheiridiidae, and Olpiidae. According to Mikhailov (2016a), from the former USSR are known 144 spp. of pseudoscorpions. The pseudoscorpions of Iran have been identified by Beier (1971), Nassirkhani and Shoushtari (2015), and Nassirkhani (several papers). Now (2017) from this country are known to live 41 spp. of 23 genera and 10 families: Chthoniidae, Atemnidae, Cheliferidae, Chernetidae, Garypinidae, Ideoroncidae, Neobisiidae, Cheiridiidae, Menthidae, and Olpiidae. 388 7 Regional Arachnogeography From Arabian Peninsula (Saudi Arabia, the Emirates, Bahrain, Qatar, Oman, Yemen – without Socotra) are known pseudoscorpions of the families Chthoniidae, Ideoroncidae, Olpiidae, Garypidae, Garypinidae, Menthidae, Atemnidae, Chernetidae, Cheliferidae, and Withiidae (Mahnert 1980, 1991). According to Mahnert (1991), “the results obtained in pseudoscorpions [of Arabian Peninsula] seem to be well in concordance with those of other animal groups: clear affinities to the Afrotropical region are recognizable in the southwestern part of the peninsula; the species of the central and northern parts are highly influenced by palearctic elements. The southeastern corner yielded oriental elements.” In Israel, Palestine, Lebanon, and Syria are known 37 spp. of Pseudoscorpiones from the families Chthoniidae, Neobisiidae, Geogarypidae, Atemnidae, Chernetidae, and Cheliferidae (Beier 1955, 1963b, Harvey 2011, Mahnert 1974). Endemic genera and species are Paramenthus shulovi Beier, 1963, and Ayalonia dimentmani Ćurčić, 2008. According to Beier (1963b), Israel’s fauna is predominantly Palearctic, but with a few species (Lamprochernes savignyi, Nudochernes spalacis, Myrmecowithius wahrmani) reminding the Afrotropical fauna and with genus Apolpium reminding the Neotropical fauna. In Cyprus have been found 12 species of 11 genera and 8 families, and the endemic genus and species is Mesatemnus cyprianus (Beier et Turk, 1952) (Atemnidae). Opiliones From the Dodecanese are registered the families Phalangiidae (Eupnoi), Trogulidae, Dicranolasmatidae, and Nemastomatidae (Dyspnoi) (Roewer 1924; Caporiacco 1926, 1929, 1948; Gruber 1963, 1966, 1978; Martens 1965). From Turkey are known Opiliones of three of the four suborders (Gruber 1969, 1998; Bayram et al. 2010; Mitov 2012, Kurt et al. 2010; Kurt et al. 2013, and others). According to Bayram et al. (2010), 50 species plus 3 subspecies of Opiliones from 25 genera in 6 families inhabit Turkey (Sironidae, Phalangiidae, Sclerosomatidae, Trogulidae, Dicranolasmatidae, Nemastomatidae). A more recent checklist (Kurt 2014) enumerates 88 spp. and 7 ssp. of 35 genera and 7 families, adding the family Ischyropsalididae. From this number 14 spp. are known from European Turkey. According to Mikhailov (2016a), from the former USSR are known 178 spp. of Opiliones. Opiliones in the described area: Cyphophthalmi Fam. Sironidae Cyphophthalmus Joseph is known from Asia Minor (Gruber 1969). Eupnoi Fam. Phalangiidae – Phalangium L., Metaphalangium Roewer, Buresilia Šilhavý, Dasylobus Simon, Egaenus C.L. Koch, Homolophus Banks, Lacinius Thorell, Redikorcevia Snegovaya et Staręga, Mitopus Thorell, Oligolophus C.L. Koch, 7.17 Turkey, Cyprus, Sinai, the Dodecanese Islands, Iran, Afghanistan, the Near East,… 389 Opilio Herbst, Platybunoides Šilhavý, Platybunus C.L. Koch, Rafalskia Staręga, Rilaena Šilhavý, Zachaeus C.L. Koch Fam. Sclerosomatidae – Leiobunum C.L. Koch Endemic genera are: Redikorcevia Snegovaya et Staręga, 2008 (Phalangiidae) – Kazakhstan (one sp.) Dyspnoi Fam. Trogulidae – Trogulus Latreille (one sp., Turkey), Trogulocratus Roewer (one endemic sp., Rhodes – T. rhodiensis Gruber, 1963) Fam. Dicranolasmatidae – Dicranolasma C.L. Koch (four spp., Turkey, Iraq) Fam. Nemastomatidae – Giljarovia Kratochvil, Histricosoma Kratochvil, Mediostoma Kratochvil, Nemastoma C.L. Koch, Paranemastoma Redikorzev, Vestiferum Martens, Pyza Staręga (two spp., Turkey), Starengovia Snegovaya (one sp. from Kyrgyzstan) Endemic genera are: Fam. Nemastomatidae Starengovia Snegovaya, 2010 (one sp. from Kyrgyzstan) Laniatores From Laniatores is known the family Phalangodidae. Endemic genera are: Fam. Phalangodidae Haasus Roewer, 1949 – Israel (one sp.) Other sources: Caporiacco (1925), Gruber (1968, 1976, 1979), Logunov (2013), Mitov (2013), Roewer (1949), Šilhavý (1967), Snegovaya (2004, 2008), Snegovaya and Staręga (2008) Araneae In Turkmenistan 335 species of spiders belonging to 162 genera and 38 families are found (Michailov and Fet 1994, Wesolowska 1996). With mountain uplift and aridization in the Pliocene, Kopet Dagh became a sublatitudinal dispersal pathway for mesophilic species. About half of araneofauna is represented by widely distributed species; of the other half, Iranian, Iranian-Turkestanian, European, European-Caucasian, and European-Mediterranean comprise the majority of the mountain araneofauna. Turanian desert species are predominant in the lowland deserts (Michailov and Fet 1994). The extensive field work of E.M. Andreeva in Tajikistan resulted in a general survey of the spiders (Andreeva 1975b), preceded by a zoogeographical analysis (Andreeva 1975a). By this time from Tajikistan have been recorded 260 species and subspecies of spiders (223 species), from them 50% being Central Asiatic endemics, 13.2% of Mediterranean elements. Others are widely distributed species (8.9% Palearctic, 7.3% Holarctic, and 2.3% cosmopolitan elements). The figure of Mediterranean elements (13.2%) curiously coincides with the number of the 390 7 Regional Arachnogeography Mediterranean elements on Balkan Peninsula (13. 8%, acc. to Deltshev, 1999, 2004). Other supposition of Andreeva (1975a) is that “…it seems highly probable that the Central Asiatic spider fauna has originated locally from the widespread genera living there during the Palaeogene period. The high percentage of Central Asiatic endemics, about half the number of species occurring there, appears to be an important argument for this assumption.” Spasskiy (1952) presumed that numerous forms arrived into the Turanian Province from northern areas (connected with Palaeogene Turgaian mesophilous forests). The first substantial paper of the spiders of Afghanistan was due to Denis (1958) on the collection of 74 spp. (30 newly described) of the 3rd Danish Expedition to Central Asia. According to Zamani et al. (2017), the spider fauna of Iran is composed of 608 recorded spp./subspp. of 259 genera and 48 families. The actualized checklist of all Turkish spiders (Bayram et al. 2014) contains 1013 spp. of 330 genera and 53 families. Papers on the Turkish spiders were published also by Brignoli (several), Bayram (2002), Seyyar et al. (2008), Topçu et al. (2005), and others. Some recent papers on the spiders of some Greek islands near Asia Minor show how incomplete are the lists available for Turkey and other countries in the area: on Chios have been recorded 315 spp. (Russel-Smith et al. 2011) and on Lesbos 299 spp. (Bosmans et al. 2009). Even much less is known the spider fauna of Cyprus (some 70 spp., after Logunov 2013). From the described area are known spiders of 50 families (no endemic families). Some endemic genera in the area are: Fam. Zoropsidae Akamasia Bosselaers, 1997 – Cyprus (one sp.) Fam. Oonopidae Megabulbus Saaristo, 2007 – Israel Megaoonops Saaristo, 2007 – Israel Semibulbus Saaristo, 2007 – Israel (one sp.) Fam. Tetragnathidae Zygiometella Wunderlich, 1995 – Israel (one sp.) Fam. Caponiidae Iraponia Kranz-Baltensperger et al., 2009 – Iran (one sp.) Fam. Miturgidae Israzorides Levy, 2003 – Israel (one sp.) Fam. Palpimanidae Levymanus Zonstein et Marusik, 2013 – Israel (one sp.) 7.18 Socotra 391 Fam. Dysderidae Tedia Simon, 1982 – Israel, Syria (two spp.) Other sources: Andreeva (1975a, 1975b, 1976), El-Hennawy (2002), Mozaffarian and Marusik (2001), Mirshami et al. (2015), Simon (1890), Tanasevitch (1989), Saaristo (2007), and others Opilioacarida The only species of Opilioacarida in the former Soviet Union (Kazakhstan) was described by Redikorzev (1937) and assigned by Chamberlin and Mulaik (1942) to a new genus. Paracarus hexophthalmus (Redikorzev, 1937) is endemic for Central Asia, but a new (fossil) species (Paracarus pristinus Dunlop, Wunderlich, et Poinar, 2004) was described from Baltic amber (? Kaliningrad). This is also the northernmost Opilioacarida in the Eastern Hemisphere. Acariformes Oribatida In Iran have been recorded 380 spp. of Oribatida of 191 genera and 86 families (Akrami 2015). 7.18 7.18.1 Socotra Geography, General Zoogeography, and Paleogeography Socotra (Soqotra) is an archipelago of four islands (Socotra, Abd al Kuri, Samhah, and Darsah) in the Indian Ocean, at some 240 km from the Horn of Africa and 380 km from the Arabian Peninsula (politically part of Yemen). The surface area is 3796 km2 and the highest point is 1503 m. It is an isolated part of Gondwana, detached in Miocene, and has three geographical parts: the narrow coastal plain, a limestone plateau with karstic caves, and the Haghier Mountains. It has a tropical desert and semi-desert climate and a mean annual temperature over 18oC (De Geest 2005). Several endemic birds, no native mammals (except bats). High endemism in phytogeography, from 825 plant species 307 are endemic (37%). Several endemic birds, no native mammals (except bats). 7.18.2 Arachnogeography So far absent from the island are the orders Thelyphonida (Uropygi), Schizomida, Holothyrida, and Opilioacarida and the suborders Palaeoamblypygi, Cyphophthalmi, Eupnoi, Dyspnoi, and Mesothelae. 392 7 Regional Arachnogeography Solifugae There is one endemic species Gluviopsis balfouri (Pocock) known from the archipelago (Socotra, Samha, Abd al Kuri) (Pocock 1889). Scorpiones At least five species of scorpions are known from the Archipelago. Two belong to the Family Buthidae, the relatively large and most common yellowish Hottentotta socotrenis (Pocock, 1889) (recorded from Socotra and Samha), and the dark coloured and apparently more rare Orthochirus bicolor insularis (Pocock, 1889) (Socotra). Widespread on Socotra is the brownish Hemiscorpius socotranus Pocock, 1889. The specimens recorded on Samha and Darsa are more light in colour and need further studies to clarify their status. The remaining two forms belong to the Family Diplocentridae (Heteronebo forbesii Pocock, 1889; H. granti Pocock, 1889), and were supposed to be restricted to Abd al Kuri. However during the last expeditions we found Heteronebo on Socotra too. It is a small genus, but of great taxonomic and zoogeographical interest, because it has a peculiar distribution, with the two (or three ?) forms on the Archipelago, and the next relatives on Caribbean islands. (Pocock 1898) Pseudoscorpiones On the island are known 19 spp. of pseudoscorpions of 12 genera and the families Chthoniidae, Ideoroncidae, Garypidae, Geogarypidae, Olpiidae (7 spp.), Menthidae, Withiidae, and Cheliferidae (Mahnert 2007, he described 17 new species, for the time being considered endemic). According to him, the fauna has clear affinities with the East African pseudoscorpions and more discrete affinities with the Arabian fauna. Endemic genus: Pseudomenthus Mahnert, 2007 (three spp.), fam. Menthidae (Socotra is considered by Mahnert a hot spot for this rare family) Opiliones Laniatores From the island are known Opiliones of the family Biantidae (Hirst 1911, Loman 1902, Pocock 1903). The endemic harvestmen Hinzuanius flaviventris (Pocock, 1903) (Biantidae) is conspicuous in reddish color. Amblypygi From Giniba Cave in Socotra was described the blind endemic species Charinus stygochtobius Weygoldt et Van Damme, 2004. It was added to the two other Socotran species Charinus soqotranus Weygoldt, Pohl et Polak, 2002, and Phrynichus heurtaultae Weygoldt, Pohl, et Polák, 2002. Araneae From Socotra have been recorded 42 species of spiders (32) from 36 genera (4 endemics) and 22 families. Salticidae is the best represented, with 15 spp. (13 endemics) (Blackwall 1877, Hirst 1911, Pocock 1903, Lehtinen 1967, Saaristo and Van Harten 2002, Wesolowska and Van Harten 2002, Jocqué and van Harten 2015). 7.19 Karakorum, Hindu Kush, Pamir, Tian Shan, Himalaya, and Tibet 393 Endemic genera are: Fam. Barychelidae Atrophothele Pocock, 1903 (one sp.) Fam. Oonopidae Socotroonops Saaristo et van den Harten, 2002 (one sp.) Fam. Palpimanidae Scelidomachus Pocock, 1899 (one sp.) Fam. Linyphiidae Agelenella Lehtinen, 1967 (one sp.) Fam. Thomisidae Bassaniodes Pocock, 1903 (one sp.) Dimizonops Pocock, 1903 (one sp.) Parasitiformes Fam. Ixodidae (Hyalomma truncatum Koch, 1844; Boophilus annulatus Say, 1821) 7.19 7.19.1 Karakorum, Hindu Kush, Pamir, Tian Shan, Himalaya, and Tibet Geography, General Zoogeography, and Paleogeography This area includes some of the highest mountains and plateaus on Earth. The culminant point of Hindu Kush Tirich Mir (in Pakistan) reaches 7690 m. The glaciers in this mountain cover 6200 km2, and the snow line is at 4650 m on the northern slope and at 5400 m on the southern slope. On the southeast slopes of Hindu Kush, the upper forest limit reaches 3300–4000 m; above this altitude spread mountain pastures. Himalaya sensu stricto is called the system of Tertiary ranges south and west of Tsangpo (Brahmaputra) and east of the Indus River. North of them lie the ranges of “Transhimalaya” (Zanskar‚ Ladakh, and Karakorum) and Tibet‚ and in the south are situated the Siwalik and other lower mountains. Himalaya is the highest mountain system in the world‚ containing more than 500 summits higher than the Montblanc. This system includes also 10 of the 14 summits higher than 8000 m. Himalaya is a very young mountain. In the Pleistocene its altitude should have been only half of the present 8848 m. The collision between the Indian subcontinent and Eurasian continent started in Paleogene time and continues today. The Indian plate continues to move northward relative to Asia about 5 cm per year. 394 7 Regional Arachnogeography With an average elevation exceeding 4500 m, the Tibetan Plateau is the world’s highest and largest plateau, with an area of 2,500,000 square kilometers. The Tibetan Plateau is surrounded by massive mountain ranges. The plateau is bordered to the south by the Himalayan Range, to the north by the Kunlun Range which separates it from the Tarim Basin, and to the northeast by the Qilian Range which separates the plateau from the Hexi Corridor and Gobi Desert. To the east and southeast, the plateau gives way to the forested gorge and ridge geography of the mountainous headwaters of the Salween, Mekong, and Yangtze rivers in western Sichuan (the Hengduan Mountains) and southwest Qinghai (Bassoullet et al. 1977, Beron 2008a, Colchen 1981, Dobremez 1972, 1976, 1978, Le Fort 1996, Mani 1968, Reinig 1930, Schweinfurth 1957, Troll 1957, Vtorov 1966, Wissmann 1959). 7.19.2 Arachnogeography Caporiacco (1934–1935) initiated the studies on the arachnofauna of the highest parts of the planet, mainly in Karakorum. Beron (2008a, with suppl. Beron, 2016) summarized the information on the high-altitude Arachnida of these mountain systems (above 2200 m) (see also Martens 1983, 1984, 1987, 1993, Schmölzer 2001). The expeditions of J. Martens resulted in hundreds of new taxa and in several analyses (Martens 1981 and others). Solifugae From the area described have been recorded six families of Solifugae (Birula 1938, Hirst 1907). Fam. Rhagodidae – Rhagoderma Roewer, Rhagodia Roewer, Rhagodixa Roewer Fam. Karschiidae – Karschia tibetana Hirst, 1907 was described from altitude 4570 m in Tibet Fam. Daesiidae – Daesiola Roewer, Haarlovina Lawrence, Mumaella Harvey Fam. Galeodidae – Galeodes setulosus Birula, 1938 is known from Tajikistan (3000 m) In Pakistan are found 29 spp. of Solifugae of the families Galeodidae (Galeodes Olivier, 22 spp.) and Rhagodidae (Rhagodes Pocock, 4spp.; Rhagodia Roewer, 1 sp.; Rhagoduna Roewer, 1 sp.; Rhagoderma Roewer, 1 sp.) Amblypygi Fam. Charinidae – Charinus pakistanus Weygoldt, 2005 (Pakistan) Scorpiones In the described area are known scorpions of six families (Fet 1988, Lourenço 1997, 2003, Lourenço et al. 2005, Lourenço and Qi 2006, Mani 1959, Shi and Zhang 2005, Tikader and Bastawade 1983, Qi et al. 2005, Zhu et al. 2004, Di et al. 2013). Fam. Scorpiopidae – Scorpiops Peters (endemic high-mountain species in Tibet, Kashmir, and Himalaya), Euscorpiops Vachon (E. montanus Karsch, Himalaya) 7.19 Karakorum, Hindu Kush, Pamir, Tian Shan, Himalaya, and Tibet 395 Fam. Buthidae – Himalayotityobuthus Lourenço (endemic genus, H. martensi Lourenço, Kashmir), Mesobuthus Vachon Fam. Scorpionidae – Heterometrus Ehrenberg Fam. Bothriuridae – Cercophonius Peters (Indian Himalaya) Fam. Chaerilidae – Chaerilus Pocock (Himalaya) Fam. Hormuridae – Tibetiomachus Lourenço et Qi (Tibet) Pseudoscorpiones In Nepal and Bhutan are known 41 spp. of pseudoscorpions of 31 genera and 14 families: Chthoniidae, Lechytiidae, Tridenchthoniidae, Neobisiidae, Syarinidae, Geogarypidae, Olpiidae, Cheiridiidae, Pseudochiridiidae, Sternophoridae, Atemnidae, Cheliferidae, Chernetidae, and Withiidae (Beier 1959, 1959, 1974, 1976, Curčić 1980, Mahnert 1977, Mani 1959, Martens 1975, Schawaller 1983, 1987, 1988, 1991). Dashdamirov (2004) recorded from northern Pakistan 30 spp. of 20 genera and 9 families: Chthoniidae, Tridenchthoniidae, Geogarypidae, Olpiidae, Neobisiidae, Cheiridiidae, Atemnidae, Cheliferidae, and Chernetidae. To them is to be added Withiidae (Nannowithius pakistanicus Beier, 1978, described from Kashmir). The preliminary conclusion from this study is that “the false-scorpion fauna of the northern mountainous provinces of Pakistan can be concluded to be Palaearctic, with only few species derived from the Oriental region.” As a whole, from the 25 species of pseudoscorpions living in the Old World above 3500 m, 15 are known from Himalaya and Karakorum (Beron 2008a). Endemic genera from the described region are: Fam. Neobisiidae Nepalobisium Beier, 1974 – Nepal (one sp.) Fam. Tridenchthoniidae Rheodithella Dashdamirov et Judson, 2004 – N. Pakistan Fam. Chernetidae Bipeltochernes Dashdamirov, 2004 – N. Pakistan Opiliones As a result of the research mainly of Martens (1972, 1973, 1977, 1978, 1979, 1980, 1984, 1987, 1993), Roewer (1957), and Suzuki (1966a, 1966b, 1970), the extremely rich fauna of Opiliones of the Himalaya was revealed. Cyphophthalmi – not recorded Eupnoi Fam. Phalangiidae – endemic genera: Himalphalangium Martens (subendemic: five end. spp. in the described area, one sp. in Japan, one in Korea; other endem- 396 7 Regional Arachnogeography ics: Homolophus Banks (= Euphalangium Roewer) thienshanense (Šilhavý, 1967) and several other endemic spp., including H. nordenskioeldi (L. Koch), the highest living Opilion at 5600 m in Karakorum; Egaenus C.L. Koch (several endemic species) Fam. Sclerosomatidae – endemic genera: Pseudastrobunus Martens and Granulosoma Martens (Nepal); Gyoides Martens (6 spp., Nepal), Diangathia Roewer (1 sp., Nepal), Globulosoma Martens (2 spp., Nepal), Harmanda Roewer (15 spp. from Nepal and India); Himaldroma Martens (2 end. spp. from Nepal); Himalzaleptus Martens (1 sp., Nepal), Metaverpulus Martens (7 spp., Nepal, Bhutan), Nepalkanchia Martens (2 spp., Nepal); Octozaleptus Suzuki (1 sp., Nepal); Pokhara Suzuki (7 spp., Nepal); Rongsharia Roewer (3 spp., Nepal); Zaleptiolus Roewer (4 spp.); other endemics: Gagrella Stolička (several endemic species), Opilio Herbst (several endemic species in Nepal and Karakorum) (Gricenko 1975, Janetschek 1990, Staręga and Snegovaya 2008). Dyspnoi Fam. Sabaconidae – Sabacon Simon (six endemic species from Nepal, up to above 5000 m) Laniatores In the described mountain area are represented Laniatores from the families Sandokanidae (Oncopodidae), Podoctidae, Phalangodidae, Epedanidae, Assamiidae, and Biantidae. Fam. Sandokanidae (Oncopodidae) – Gnomulus Thorell Fam. Podoctidae – in India five species Fam. Phalangodidae (? Epedanidae) – endemic genus: Dhaulagirius Martens (Nepal) Fam. Assamiidae – endemic genera: Assaphala Martens (Nepal), Micrassamula Martens (Nepal), Nepalsia Martens (Nepal), Nepalsioides Martens (Nepal) Fam. Biantidae – Biantes Simon (many endemic species from Nepal and India) Araneae To the important paper of Caporiacco (1934) on the Arachnida of Karakorum have been added further studies of this mountain system (Beron 2008a, 2016). Andreeva (1975, 1976) summarized the information on the spiders of Tajikistan. According to Tikader (1970), the spider fauna of Sikkim consists of 65 species (45 of them described as new and presumably endemic) of 11 families. The spiders of the Himalaya above 2200 m belong to the families Anapidae, Oonopidae, Zodariidae, most of all Linyphiidae, Tetrablemmidae, Amaurobiidae, Lycosidae, Araneidae, Sparassidae, Clubionidae, Drassodidae, Dictynidae, Scytodidae, Hahniidae, Gnaphosidae, Sicariidae, Thomisidae, and Salticidae. The families found above 3500 m are underlined. Spiders (indet.) have been observed up to 6700 m, Acantholycosa baltoroi (di Caporiacco) – up to 6100 m. Ixodida Ref.: Clifford et al. (1975), Beron (2008a) Argas himalayensis Hoogstraal et Kaiser parasitizes Lerwa lerwa up to 4575 m. Ixodes berlesei Birula in Nepal is found up to 5488 m. Almost all genera of Himalayan Argasidae and Ixodidae are 7.20 Indian Peninsula 397 widespread. Special is Anomalohimalaya Hoogstraal, Kaiser, et Mitchell, high mountain inhabitant of Pamir, Tajikistan and Himalaya, up to 3800 m. 7.20 7.20.1 Indian Peninsula Geography, General Zoogeography, and Paleogeography Geographically, the Indian subcontinent is a peninsular region in South-Central Asia, delineated by the Himalaya in the north, the Hindu Kush in the west, and the Arakanese in the east and extending southward into the Indian Ocean with the Arabian Sea to the southwest and the Bay of Bengal to the southeast. This region rests on the Indian Plate and is isolated from the rest of Asia by mountain barriers (Alegre et al. 1984, Dobremez 1972, 1976, 1978, Hocutt 1987, Katz and Premoli 1979, Mani (Ed.) 1974, Sahni 1984, Sahni and Kumar 1974). Upland plain (Deccan Plateau) and plains near Ganges River are typical for the peninsula, and to the north India shares with Nepal the third highest mountain in the world – Kanchenjunga (8,586 m). Andaman and Nicobar Islands also belong to India. The onset of the collision of India with Asia occurred near the Paleocene-Eocene boundary (Gaetani and Garzanti 1991), some 55 million years ago. To quote Briggs (1989): “If India broke its contact with the other continents sometime around 148 m.y.a. and, if it existed as an isolated, oceanic continent until the early Miocene, its fossil terrestrial and shallow marine biotas should demonstrate the evolutionary effects of more than 100 m.y. of isolation. This means that India should have developed a peculiar biota with a high percentage of distinct genera and families. But, with the possible exceptions ...the expected preponderance of peculiar organisms has simply not been found.” 7.20.2 Arachnogeography The northern part of India is the chain of Himalaya where there is a transition between the Palearctic and the Indomalayan regions and even between two Kingdoms – the Holarctic and the Paleotropic. It is difficult to differentiate the genera and species of Arachnida in this area (Blasco 1981, Briggs 1989). In Peninsular India are known the orders Palpigradi, Solifugae (21 spp.), Scorpiones (117 spp.), Pseudoscorpiones (165 spp.), Opiliones (1 endemic genus in Cyphophthalmi, Dyspnoi are lacking), Amblypygi (4 endemic species), Uropygi (endemic genus), Schizomida (several species, 1 endemic genus), Araneae (1442 spp. of 59 families), and Opilioacarida (1 endemic genus) and many different mites. Lacking are also the orders Holothyrida and Ricinulei and the suborders 398 7 Regional Arachnogeography Palaeoamblypygi and Dyspnoi. In India are not recorded endemic families of Arachnida. In the western edge of the peninsula (Thar Desert) is passing the band separating Holarctic and Paleotropic Kingdoms (Map 8.2). Palpigradi From India have been recorded three taxa: Eukoenenia angusta hindua Condé, 1989; E. a. tamula Remy, 1960; and E. singhi Condé, 1989. Solifugae From India are known 21 spp. of Solifugae, belonging to the genera Galeodes Olivier, 14 spp. (Galeodidae), Gluviopsis Roewer (Daesiidae) and Rhagodеrma Roewer, Rhagodomma Roewer, Rhagodima Roewer, and Rhagodopa Roewer (Rhagodidae) (Pocock 1895). Endemic genera for India are: Rhagodomma Roewer, 1933 – one sp. Rhagodima Roewer, 1933 – one sp. Scorpiones In India are known 117 spp. of Scorpiones of the families Bothriuridae, Buthidae, Chaerilidae, and Scorpiopidae (see the list) (Ambalaparambil et al. 2010, Bastawade 2006, Lourenço 1996, 1997, 2003, Tikader and Bastawade 1983, Vachon 1982). Pseudoscorpiones From India are known about 160 spp. (and several subspecies) of pseudoscorpions of 61 genera. Altogether from India, Nepal, and Bangladesh are known 71 genera and the families Chthoniidae, Hyidae, Geogarypidae, Feaelidae, Atemnidae, Cheliferidae, Lechytiidae, Chernetidae, Garypidae, Geogarypidae, Ideoroncidae, Olpiidae, Cheiridiidae, Pseudotyrannochthoniidae, Sternophoridae, Syarinidae, and Withiidae (Beier 1973, 1974, Murthy and Ananthakrishnan 1977, Vachon 1982). Endemic genera for Peninsular India are: Ectoceras Stecker, 1875 – India (two spp.) Hygrochelifer Murthy et Ananthakrishnan, 1977 – India (two spp.) Opiliones In Peninsular India have been registered the families Stylocellidae (Cyphophthalmi), Phalangiidae, Sclerosomatidae (Eupnoi), and Epedanidae, Podoctidae, Sandocanidae, and Assamiidae (Laniatores), with several endemic genera (Roewer 1929a, 1929b, Giribet et al. 2007). Cyphophthalmi From India (Arunachal Pradesh) has been described an endemic genus and species Meghalaya annandalei Giribet, Sharma, et Bastawade, 2007 (Stylocellidae) Eupnoi Fam. Phalangiidae – Euphalangium Roewer Fam. Sclerosomatidae – Metagagrella Roewer 7.20 Indian Peninsula 399 Dyspnoi – not represented Laniatores Fam. Epedanidae (incl. Dibuninae, Acrobuninae, Sarasiniciinae, Sarasinellinae) – Aboriscus Roewer, Buparus Thorell Fam. Podoctidae (incl. Erecananinae and Ibaloniinae) – Eupodoctis Roewer, Pumbaraius Roewer, Tandikudius Roewer, Vandaravua Roewer Fam. Sandokanidae [= Oncopodidae, praeoccup.] – Gnomulus Thorell Fam. Assamiidae – Assamia Sørensen Amblypygi On the peninsula are known four species (endemics) (Bastawade et al. 2005, Gravely 1911, 1915, Pocock 1894). Fam. Charinidae Charinus Simon – Ch. bengalensis (Gravely, 1911) Sarax Simon – S. cochinensis (Gravely, 1915) Fam. Phrynichidae Phrynichus Karsch – Ph. andhraensis Bastawade, Rao, Javed, et Krishna, 2005; Ph. phipsoni (Pocock, 1894) Thelyphonida (Uropygi) From India are known two spp. of Labochirus Pocock, one of Hypoctonus Thorell, one of Thelyphonus Latreille, and Uroproctus assamensis (Stoliczka). According to Bastawade (2006), the Indian fauna of Uropygi consists of four genera and six species (Butler 1872, Pocock 1900, Stoliczka 1869, Gravely 1912 Harvey 2013 ). Schizomida From India (Maharashtra) has been described the endemic (so far) genus Neozomus with the species N. tikaderi (Cokendolpher, Sissom, et Bastawade, 1988). Also from India are known other species of “Schizomus” kharagpurensis Gravely, Ovozomus lunatus Gravely, and others (see Reddell and Cokendolpher 1995), requiring further study. Together with the species described by Bastawade (2002, 2004, 2006), in India are known seven spp. of Schizomida. A new genus (Gravelyzomus Kulkarni) has been described for “Schizomus” chaibassicus Bastawade, 2002, and a new sp. Schizomus arunachalicus Bastawade was found in Arunachal Pradesh (Bastawade 1985, 2002, 2004, 2006 Cokendolpher et al. 1988, Gravely 1911, 1912, 1915, 1925, Kulkarni 2012, Pocock 1900, Reddell and Cokendolpher 1995). Araneae According to the list of Indian spiders (Siliwal, Molur and Biswas 2005) in the whole of India are recorded 1442 species of 361 genera and 59 families (see also Brignoli 1972, Pocock 1900, Tikader 1970, 1987). The genera with more than 50 spp. are Araneidae (147), Gnaphosidae (134), Lycosidae (126), Oxyopidae (65), Salticidae (181), Sparassidae (80), Tetragnathidae (51), Theraphosidae (51), Theridiidae (52), and Thomisidae (154). Of the 1442 spp., 1002 are endemic to 400 7 Regional Arachnogeography mainland India. In the updated list of Siliwal and Molur (2007) are 1447 spp., 365 genera, and 60 fam. (20 genera are endemic to India). No endemic families. Opilioacarida The only representative of Opilioacarida in Indian subcontinent is Indiacarus pratyushi Das et Bastawade, 2007 – India (endemic genus and species). Acariformes Prostigmata Fam. Erythraeidae Microsmarialla coniferana Khot, 1963 (endemic genus and species) 7.21 7.21.1 Ceylon (Sri Lanka) Geography, General Zoogeography, and Paleogeography The Island of Ceylon, on which is situated the state Sri Lanka, has been covered with luxuriant tropical forests, but most of them have been replaced by tea plantations. The island is close to India and even is almost connected with the continent by several islands called Adam’s Bridge. The surface of the island is 65,610 km2. Geologists differ concerning how and when arose the 30 km-long Adam’s Bridge (Pathirana 1980). For us there is no doubt that Ceylon had a land connection with the continent. 7.21.2 Arachnogeography Despite the quasi-total loss of forests, replaced by tea plantations, this mountainous island still has quite interesting arachnofauna. Represented are all main orders and suborders (except Ricinulei, Palaeoamblypygi, Dyspnoi, and Mesothelae). Especially interesting is the genus Pettalus (from fam. Pettalidae, known from faraway southern areas like Chile, South Africa, and New Zealand). In India live Cyphophthalmi of different family (Stylocellidae). Remarkable is the holothyrid endemic genus Indothyrus Lehtinen (Holothyrida are not known from the nearby India). The Palpigradi indicate connections with Madagascar and Southeast Asia. Palpigradi The three species found in Sri Lanka (Remy 1961b) show connections with South Asia (Eukoenenia angusta in Thailand and India), but also with Madagascar and the Mascarene (Eukoenenia chartoni, Koeneniodes madecassus). Remy (1961b) was a supporter of the ideas of Jeannel for the existence of continental interconnections between Madagascar, the Seychelles, Sri Lanka, Mauritius, and India. 7.21 Ceylon (Sri Lanka) 401 Thelyphonida (Uropygi) Labochirus proboscideus Butler has been described from Sri Lanka and is an endemic species to this island (Butler 1872, Pocock 1900). Schizomida From Sri Lanka are known the first schizomid ever described: Schizomus crassicaudatus Pickard-Cambridge, 1972, and also several other “Schizomus” needing further study (buxtoni Gravely, formicoides Fernando, greeni Gravely, peradeniyensis Gravely, perplexus Gravely, suboculatus Pocock, vittatus Gravely) (Cokendolpher et al. 1988, Fernando 1957, Gravely 1911, 1912, 1915, Pickard-Cambridge 1972, Pocock 1900, Reddell and Cokendolpher 1995, Harvey 2013g). Scorpiones On the island are recorded 15 spp. of the genera Buthoscorpio Werner, Charmus Karsch, Hottentotta Birula, Isometrus Ehr., Lychas C.L. Koch (Buthidae), and Chaerilus Simon (Chaerilidae) (Lourenço 1997, Tikader and Bastawade 1983, Vachon 1982). Endemic species of scorpions in Sri Lanka is Lychas srilankensis Lourenço, 1997 – low level of endemicity. Pseudoscorpiones On Sri Lanka have been recorded 43 spp. of Pseudoscorpiones, belonging to 31 genera and 14 families: Chthoniidae (2 spp.), Pseudotyrannochthoniidae (2), Atemnidae (11), Feaellidae (1), Garypidae (1), Geogarypidae (3), Hyidae (1), Ideoroncidae (1), Sternophoridae (1), Olpiidae (5), Syarinidae (1), Cheliferidae (5), Chernetidae (7), and Withiidae (2) (Beier 1973a, Batuwita and Benjamin 2014). Some endemic species for the island are: Fam. Chthoniidae – Lagynochthonius brincki Beier, Afrochthonius ceylonicus Beier, A. reductus Beier Fam. Syarinidae – Ideobisium ceylonicum Beier Fam. Olpiidae – Olpium ceylonicum Beier There are no genera, recorded only from Sri Lanka. Opiliones On Sri Lanka have been recorded 26 spp. of Opiliones from Pettalidae, Sclerosomatidae, Podoctidae, and Assamiidae (Boyer and Giribet 2007, Giribet 2008, Karsch 1891, Roewer 1910, 1912, 1929, Šilhavý 1974a, 1974b, Sharma et al. 2009, Sharma and Giribet 2006, Thorell 1876) (after Kury, online). Cyphophthalmi On Sri Lanka were recorded four species of the remarkable Gondwanan endemic genus Pettalus Thorell, a base of the family Pettalidae, spread over Sri Lanka, New Zealand, Australia, South Africa, and Chile. Eupnoi From Sri Lanka is known the family Sclerosomatidae (Gagrella, Gagrellula) Dyspnoi – not represented Laniatores From Sri Lanka are known Laniatores from the families: Podoctidae – Eusitalces Roewer, Eurytromma Roewer, Neopodoctis Roewer 402 7 Regional Arachnogeography Assamiidae – Assamia Soerensen, etc. Trionyxellinae – India and Sri Lanka The endemic genera are in bold: Solifugae From Sri Lanka is known one sp. of Solifugae (Pocock 1895). Fam. Rhagodidae Rhagodes phipsoni (Pocock) (endemic sp.) Araneae In Sri Lanka have been recorded 383 spp. of spiders of 213 genera and 46 families (Benjamin et al. 2012, Brignoli 1972, Huber and Benjamin 2005, Pocock 1900, Siliwal et al. 2005, Tikader 1970, 1987). These figures include 271 spp. and 22 genera endemic for Sri Lanka. Benjamin, Channa and Bambaradeniya (2006) affirm that the known spiders on the island are 501 spp. of 46 families, including 106 spp. of Salticidae. Spider families in Sri Lanka: Agelenidae, Araneidae, Barychelidae, Clubionidae, Corinnidae, Ctenidae, Dictynidae, Dipluridae, Eresidae, Eutichuridae, Hahniidae, Hersiliidae, Idiopidae, Linyphiidae, Liocranidae, Lycosidae, Mimetidae, Mysmenidae, Nesticidae, Ochyroceratidae, Oonopidae, Oxyopidae, Palpimanidae, Philodromidae, Pholcidae, Phrurolithidae, Pisauridae, Psechridae, Salticidae, Scytodidae, Segestriidae, Sparassidae, Stenochilidae, Tetrablemmidae, Tetragnathidae, Theraphosidae, Theridiidae, Theridiosomatidae, Thomisidae, Titanoecidae, Trachelidae, Ubudidae, Uloboridae, Zodariidae, and Zoropsidae Some endemic genera are: Fam. Barychelidae Plagiobothrus Karsch, 1891 (one sp.) Fam. Ctenidae Diallomus Simon, 1897 (two spp.) Fam. Dictynidae Atelolathys Simon, 1892 (one sp.) Rhion O. P.-Cambridge, 1870 (one sp.) Fam. Tetrablemmidae Gunasekara Lehtinen, 1981 (one sp.) Fam. Zoropsidae Devendra Lehtinen, 1967 (three spp.) Fam. Linyphiidae Labullinyphia van Helsdingen, 1985 7.22 Andaman and Nicobar Islands 403 Fam. Hersiliidae Promurricia Baehr et Baehr, 1993 (one sp.) Fam. Tetragnathidae Atelidea Simon, 1895 (one sp.) Schenkeliella Strand, 1934 (one sp.) Fam. Mysmenidae Phricotelus Simon, 1895 (one sp.) Fam. Udubidae Campostichomma Karsch, 1892 (one sp.) Fam. Pholcidae Wanniyala Huber et Benjamin, 2005 Fam. Thomisidae Peritraeus Simon, 1895 (one sp.) Holothyrida Only Indothyrus greeni Lehtinen, 1995, is known from Sri Lanka (endemic genus and species). Ixodida According to Dilrukshi (2006), in Sri Lanka are registered 27 spp. of Ixodidae of 9 genera, mostly widespread (Nosomma monstrosum Nuttall et Warburton, 1908 is endemic to Ceylon). 7.22 7.22.1 Andaman and Nicobar Islands Geography, General Zoogeography, and Paleogeography The territory comprises two island groups in the Indian Ocean, the Andaman Islands and the Nicobar Islands, separated by the 10° N parallel, with the Andaman to the north of this latitude and the Nicobar to the south. The total land area of the Andaman Islands is approximately 6408 km2 and of Nicobar Islands – 1841 km2. The islands are situated in the Bay of Bengal, and geographically are part of Southeast Asia, 150 km north of Aceh in Indonesia and separated from Thailand and Burma by the Andaman Sea. The Andaman Islands were linked to the mainland during the low sea levels in glaciation time. 404 7.22.2 7 Regional Arachnogeography Arachnogeography So far Pseudoscorpiones, Scorpiones, Amblypygi and Araneae are recorded from these isolated and rarely visited islands. Some species are known from Madagascar and Aldabra. Low-level endemism. Pseudoscorpiones From the Andaman Islands, Beier (1981) recorded three species: Xenolpium madagascariense Beier, known from Madagascar and Aldabra; Anagarypus oceanusindicus Chamberlin, known from Aldabra and Chagos Archipelago; and Pseudochiridium clavigerum (Thorell), known from India and Indonesia. From the Nicobar Islands are known Catatemnus nicobarensis With, 1906, Garypus nicobarensis Beier, 1930. Scorpiones From the Andaman Islands (Little Andaman) is known the endemic Chaerilus andamanensis Lourenço, Duhem, et Leguin, 2011. Before that only the widespread Isometrus maculatus (De Geer) and Liocheles australasiae (Fabricius) have been recorded (Tikader and Bastawade 1983). Amblypygi From the Andaman Islands is known the family Charinidae – Charinus Simon, Sarax Simon. Araneae In the paper of Tikader (1977) and other papers are listed 20 families of spiders of 41 genera, found in the Andaman and Nicobar Islands. There are also 26 newly described (? endemic) species, among the 58 in the list. The families are Scytodidae, Filistatidae, Uloboridae, Amaurobiidae, Dictynidae, Theridiidae, Pholcidae, Linyphiidae, Tetragnathidae, Araneidae, Hersiliidae, Oxyopidae, Pisauridae, Lycosidae, Gnaphosidae, Heteropodidae, Thomisidae, Clubionidae, Salticidae, and Psechridae. 7.23 7.23.1 Southeast Asia (Burma, Thailand, Indochina, Peninsular Malaysia, China South of Yangtze, and Hainan) Geography, General Zoogeography, and Paleogeography The continental part of Southeast Asia is bordering India to the west and China to the north. On it are situated the countries Burma (Myanmar), Thailand, Laos, Cambodia, Vietnam, western Malaysia, and Singapore (connected with bridge to the 7.23 Southeast Asia (Burma, Thailand, Indochina, Peninsular Malaysia, China South… 405 continent). The development of the peninsula has been analyzed many times by Hall (1997, 1998, 2001, 2002), Hall and Holloway (Eds) (1998), and other authors. Indochina is (was) covered by lush rain forest and is very fast dwindling. The peninsula is crossed by big rivers, coming from the northern mountains. The northern part of Burma includes some parts of Himalaya (Peak Hkakabo Razi, 5881 m). In the other parts of the peninsula, the highest summit is Fansipan in Vietnam (3143 m). The loss of habitats, deforestation, and replacement of the rain forest by oil palm plantations are very serious treats to the rich and varied wildlife of the peninsula. The paleogeography of South China and the adjacent territories is presented in the Atlas of Wang Hongzhen, Ed. (1985). The Hainan Island had been connected with the mainland until Miocene, when faulting caused subsidence and brought about the formation of the Qiongzhou Strait (Wang Hongzhen, Ed., 1985). The biogeographic analysis of Hua Zhu (2016) indicates the low endemism of the flora (7 seven endemic genera out of 1283), and the conclusion is that “Hainan Island could have been adjacent to northern Vietnam and the Guangxi at least in Eocene.” “Today, the waters of SE Asia contain the highest marine faunal diversity in the world, and the islands of the region contain some of the most diverse collections of plant and animal species found on Earth. The period 30-0 Ma is of most interest to biogeographers since before then the separation between Asia and Australia was greater and for almost all land plants and animals it was probably not possible to cross this barrier… There were never continuous land links between Sundaland and Australia”. (Hall, 2001) Other sources: Blair Hedges (2006), Dao Van Tien (1978), Golonka et al. (2006), Hoffmann (2001), Lydekker (1896), Metcalfe (2002), Turner et al. (2001), Wallace (1860), Weber (1899) 7.23.2 Arachnogeography The Malayan part of the Indomalayan Region has a rich and varied arachnofauna. They are represented by Palpigradi (both families); Solifugae (one genus and species, the easternmost member of the order); scorpions of the families Buthidae, Scorpionidae, Chaerilidae (the center of speciation), Euscorpiidae, Hormuridae, and Pseudochactidae (the most remarkable recently described relict genera); Pseudoscorpiones; Opiliones; Araneae; Opilioacarida (two endemic genera); and other Acari. It is to mark the importance of the cave habitat for certain groups (Opilioacarida, Pseudochactidae). The area is also center of such groups as Stylocellidae (Opiliones) and Mesothelae (Araneae) (Zhang and Zhao 1978). Palpigradi Eight species have been recorded from Thailand: Prokoenenia asiatica Condé, 1994, (Prokoeneniidae) and six spp. of Eukoeneniidae – Eukoenenia angusta Hansen, 1901; E. deleta Condé, 1992; E. lyrifer Condé, 1992; E. siamensis Hansen, 1901; E. thais Condé, 1988; Koeneniodes leclerci Condé, 1992; and K. spiniger Condé, 1984. All of them are known only from this country (endemics). Condé 406 7 Regional Arachnogeography (1992) recorded Koeneniodes madecassus from Hong Kong (the first Palpigradi known from China). Solifugae The only member of Solifugae in Southeast Asia is Dinorhax rostrumpsittaci (Simon, 1877) (Vietnam, Melanoblossiidae). Scorpiones In the Indomalayan Region are recorded scorpions of 34 genera and six families Pseudochactidae. Lourenço (2007) added to the only known species of Pseudochactas Gromov from Central Asia another endemic genus and species Troglokhammouanus steineri Lourenço, 2007 from a cave in Laos. A third endemic genus and the species Vietbocap canhi Lourenço et Ðình Sắc Pham, 2010, V. thienduongensis Lourenço et Ðình Sắc Pham, 2012, and V. lao Lourenço, 2012 (Vietbocapinae, end. subfam.) have been found in caves in Vietnam and Laos. They are all relicts (Table 7.2). According to Di et al. (2011), Yunnan Province has the biggest scorpion biodiversity in the whole of China (nine spp.). From the Hainan Island have been recorded three widespread species (Di et al. 2011) and two endemic taxa. Pseudoscorpiones In Southeast Asia are recorded pseudoscorpions of 18 families: Chthoniidae, Lechytiidae, Tridenchthoniidae, Sternophoridae, Atemnidae, Ideoroncidae, Hyidae, Neobisiidae, Syarinidae, Pseudocheiridiidae, Geogarypidae, Garypinidae, Olpiidae, Parahyidae, Cheiridiidae, Cheliferidae, Chernetidae, and Withiidae (Beier 1951, 1967, Ćurčić 1983, Dashdamirov 1997, 2007, Ellingsen 1911, Gao and Zhang 2013a, 2013b, Mahnert 2003, 2009, Redikorzev 1938, Schawaller 1994b, 1995, Thorell 1889, With 1906). Nguyen Thi Dinh (20) published a list of the pseudoscorpions of Vietnam, containing 62 spp., of 32 genera and 13 families. Parahyidae is an endemic family (Singapore, Caroline Islands). Number of pseudoscorpion species in some of the countries in the area: Burma (Myanmar), 13; Thailand, 43; Vietnam, 62; Laos, 9; Cambodia, 15 (Harvey, 2013f) Endemic pseudoscorpion genera in Southeast Asia are: Fam. Garypinidae Caecogarypinus Dashdamirov, 2007 – Vietnam (one sp.) Fam. Ideoroncidae Shravana Chamberlin, 1930 – Thailand (one sp.) Fam. Cheliferidae Tetrachelifer Beier, 1967 – Vietnam (two spp.) 7.23 Southeast Asia (Burma, Thailand, Indochina, Peninsular Malaysia, China South… 407 Table 7.2 Scorpions in the Indomalayan Region Country Number of species Taxa Fam. Bothriuridae Cercophonius Peters Fam. Buthidae Androctonus Ehrenb. Buthacus Birula Buthoscorpio Werner Charmus Karsch Compsobuthus Vachon Hemibuthus Pocock Himalayotityobuthus Lourenço Hottentotta Birula Isometrus Ehr. Lychas C.L. Koch Mesobuthus Vachon Odontobuthus Vachon Orthochirus Karsch Thaicharmus Kovařik Vachonus Tikader et Bastawade Fam. Chaerilidae Chaerilus Simon Fam. Euscorpiidae Alloscorpiops Vachon Dasyscorpiops Vachon Euscorpiops Vachon Neoscorpiops Vachon Parascorpiops Banks Scorpiops Peters Fam. Hormuridae Chiromachetes Pocock Hormurus Thorell India Nepal Bhutan Bangladesh Burma Sri Lanka Mal. 117 11 4 5 11 15 27 Indon. 29 + 1 + 2 1 4 – – + – – – – – – – – – – – + – – – – – + – – – – – + – – 1 – – + – – – – – + – – – 3 3 – – – – – – – – 2 – – – – – 1 2 – 1 – – – – – – – – – – – – 6 13 15 – 1 – 1 4 – – – – – – – – 1 – – – – – 4 – – 1 6 1 – – – 3 5 1 – – 7 6 – – 6 2 – – – – – – – – – – – – – – 1 – – – – – – – + 6 + – + 2 + – – – + – + 1 + – – – + 1 + 1 – – + 11 + – + 8 – – – – – – – – 1 – 4 4 – – 2 – 1 – 4 – – – – – – – – – – – – – 1 – 10 + 2 2 – – 2 – – 1 + – – + – – – – – – + – – + – – – – – – – 2 (continued) 408 7 Regional Arachnogeography Table 7.2 (continued) Country Hormiops Fage Iomachus Pocock Liocheles Sundevall Fam. Scorpionidae Heterometrus Ehr. Rugodentus Bastawade et al. India – 4 2 + 23 1 Nepal – – – + 1 – Bhutan – – – – – – Bangladesh – – 1 – – – Burma – – 1 + 1 – Sri Lanka – – – + 3 – Mal. 1 – 1 + 3 – Indon. – – 3 + 3 – Country Number of species Taxa Fam. Buthidae Hottentotta Birula Isometrus Ehrenberg Lychas C.L. Koch Mesobuthus Vachon Orthochirus Karsch Thaicharmus Kovařik Razianus Farzanpay Fam. Chaerilidae Chaerilus Simon Fam. Euscorpiidae Alloscorpiops Vachon Euscorpiops Vachon Scorpiops Peters Fam. Hormuridae Hormiops Fage Hormurus Thorell Liocheles Sundevall Tibetiomachus Lourenço et al. Fam. Scorpionidae Heterometrus Ehr. Fam. Pseudochactidae Troglokhammouanus Lourenço Vietbocap Lour. et Pham Thailand Laos Vietnam Cambodia Philippines 20 15 25 5 14 China [50] Hainan 5 + – 3 3 – – 1 + – 1 4 – – – + – 3 1 – – – + – 1 1 – – – + – 2 3 – – – + 2 3 2 6 1 – + – 2 1 1 – – – + 2 + 2 – + 1 + 2 – + 6 + – – + 1 – – – + 3 – – 1 + 8 + – – – – – – 3 1 + – – 1 – 2 1 – – – – – 5 2 + 1 – 1 – – – – – – – – – – + – 2 1 – 11 11 + – – 1 1 – – + – – 1 – + 4 – + 2 + + 4 + + 2 – + 3 – + 3 – – – – – 1 – – – – – – 1 2 – – – – 7.23 Southeast Asia (Burma, Thailand, Indochina, Peninsular Malaysia, China South… 409 Opiliones Cyphophthalmi In Eastern Palearctic the suborder Cyphophthalmi is known only from Japan (Suzukielus sauteri Roewer) – endemic genus Suzukielus Juberthie. In the Southeast Asia are represented five of the six genera of fam. Stylocellidae, two of them endemic (Thorell 1882, 1890, Rambla 1991, 1994, Giribet 2000, Clouse 2012, Clouse et al. 2011, Schwendinger and Giribet 2005, Sharma and Giribet 2009, 2011, Clouse and Giribet 2010, Shear 1993, Clouse et al. 2010, Kury et al. 2009): Subfam. Fangensinae (endemic) Fangensis Rambla, 1994 – Thailand (three spp.) Giribetia Clouse, 2012 – Thailand (one sp.) Subfam. Stylocellinae Leptopsalis Thorell, 1882 – Malaya (three spp.) Stylocellus Westwood, 1874 – Malaya (one sp.) Miopsalis Thorell, 1890 – Malaya (two spp.) The species are also endemic, mostly known from the type localities. Map 7.11 Distribution of Stylocellidae (After Clouse 2012) 410 7 Regional Arachnogeography Three genera of fam. Stylocellidae are recorded in Malay Archipelago (19 spp.): Leptopsalis Thorell, 1882 – Sumatra (three spp.), Java (two spp.), Sulawesi (four spp.) Stylocellus Westwood, 1874 – Sumatra (one sp.) Miopsalis Thorell, 1890 – Borneo (nine spp.) Two species of Stylocellus Westwood have been found in the western part of New Guinea, crossing, thus, the Lydekker’s Line (Clouse and Giribet 2007). In Sri Lanka live three species of the endemic genus Pettalus Thorell, 1876 (fam. Pettalidae, known also from Madagascar, Chile, South Africa, Australia, and New Zealand) (Sharma and Giribet 2006, Sharma et al. 2009). From India (Arunachal Pradesh) has been described an endemic genus and species Meghalaya annandalei Giribet, Sharma, et Bastawade, 2007 (Stylocellidae). The family Pettalidae is not known so far from the Asiatic continent (Giribet 2000 and suppl.). Dyspnoi Eastern Palearctic is inhabited by members of the families Sabaconidae, Nemastomatidae, and Nipponopsalididae; present are also a few Ischyropsalididae and Trogulidae. Two more families (Dicranolasmatidae and Taracidae) are dwellers of Western Palearctic and the USA. (Schönhofer 2013). Fam. Sabaconidae – the only genus Sabacon Simon is widespread (USA, Europe, Japan, China, Siberia, Altai; several species have been described from Nepal – the high Himalaya up to above 5000 m, which form part of the boundary between Palearctic and Indomalayan regions). Tsurusaki and Daxiang Song (1993b) published two new species of Sabacon from Sichuan Province (China). Fam. Ischyropsalididae – found easternmost to Tajikistan. Fam. Nemastomatidae – in the Eastern Palearctic, the genera Mediostoma Kratochvil (Iran, Tajikistan) and Starengovia Snegovaya (Kyrgyzstan) reach the mountains of Central Asia. Martens (2016) published as new genus and sp. the first nemastomatine from South China (Yunnan, 3800 m) – Sinostoma yunnanicum. This was quite surprising finding of a Palearctic family far from the easternmost localities of the family (Tajikistan, Kyrgyzstan). Also, this is the maximal altitude reached by a member of Nemastomatinae (relict). From Southeast Asia are recorded only two other spp. of Dyspnoi from the family Nemastomatidae. The finding of Cladolasma [Dendrolasma] angka (Schwendinger et Gruber, 1992) (Ortholasmatinae) in Thailand is the second species of the genus Cladolasma Suzuki; the other species is C. parvulum Suzuki from Japan. Another Cladolasma – C. damingshan Zhang et Zhang (Zhang and Zhang 2013) – was described from Guangxi, China (the first representative of Nemastomatidae in China). Fam. Nipponopsalididae – three species of genus Nipponopsalis Martens et Suzuki from Korea and Japan (including Ryukyus and the Kuril Islands) (Schönhofer 2013) 7.23 Southeast Asia (Burma, Thailand, Indochina, Peninsular Malaysia, China South… 411 Fam. Trogulidae – easternmost to the northern Iran (Schönhofer 2013) Practically the Dyspnoi are missing in the Indomalayan Region. Tsurusaki and Daxiang Song (1993a) recorded the occurrence of Crosbycus dasycnemus in China (already known from Japan and North America). According to Schönhofer (2013), Crosbycus should belong to the family Taracidae, and its amphipacific populations “require investigation for specific distinction.” Suzuki (1972) analyzed some other cases of discontinuous distribution of opilions. Eupnoi In Eastern Palearctic have been found members of the families Caddidae, Phalangiidae, and Sclerosomatidae (Gricenko 1979a, Gritsenko 1979b, 1980; Starêga 1978; Tsurusaki et al. 2000). Fam. Caddidae – only one species of genus Caddo Banks (otherwise North American) is known from Japan. Even the species Caddo agilis Banks is shared between Japan and North America (Suzuki and Tsurusaki 1983). Fam. Phalangiidae – genera, represented in Eastern Palearctic: Lacinius Thorell (China), Mitopus Thorell (Japan, Mongolia), Oligolophus C.L. Koch (China), Egaenus C.L. Koch (Karakorum, Iran, Mongolia, Siberia, Tajikistan, Uzbekistan), Homolophus Banks (Altai, Korea, Siberia, Mongolia, China, Kazakhstan), Opilio Herbst (China, Mongolia, Iran, Japan, Kuril Islands, Karakorum, Kazakhstan), Scleropilio Roewer (= Scutopilio Roewer) (Central Asia), Acanthomegabunus Tsurusaki et al. (Siberia), Liropilio Gritsenko (Kazakhstan, Russia), Phalangium L. (? China), Rilaena Šilhavý (Iran), Thapinius Roewer (Kamchatka), Pamirphalangium Starêga (Pamir, nomen nudum). Many of these genera are known also from Europe. Himalphalangium was described by J. Martens from Nepal Himalaya, where the Eastern Palearctic meets the Indomalayan Region. Other important papers on the Opilions of Himalaya are due to Martens (1977 and others). Fam. Sclerosomatidae – genera, represented in Eastern Palearctic: Gagrellula Roewer (China, Japan), Harmanda Roewer (Nepal), Harmandina (China), Psathyropus L. Koch (Japan, Far East of Russia), Pseudogagrella Redikorzev (China), Systenocentrus Simon (Japan), Leiobunum C.L. Koch (Japan), Nelima Roewer (Japan), Pseudohomalenotus Caporiacco (Karakorum), Pygobunus Roewer (Japan). Many of these genera are known also from Europe. Some genera (Himaldroma, Nepalgrella, Nepalkanchia, Gyoides) were described by J. Martens from Nepal Himalaya (Martens 1983, 1984 and others). After Roewer and Suzuki, Tsurusaki (1991) reported opilions from Taiwan. From Southeast Asia are recorded many species of Eupnoi from the families Phalangiidae and Sclerosomatidae, particularly Gagrellinae (many papers of Thorell, Roewer, Suzuki, and other authors). Some endemic genera are Mitopiella Banks; Adungrella Roewer, Akalpia Roewer, Altobunus Roewer, Antigrella Roewer; Aurivilliola Roewer; Bakerinulus Roewer; Bastia Roewer; Baturitia Roewer; Biceropsis Roewer; Bonthainia Roewer; Bullobunus Roewer; Carinobius Roewer; Carmichaelus Roewer Ceratobunellus Roewer; Ceratobunoides Roewer; Cervibunus Roewer; Chasenella Roewer; Chebabius Roewer; Coonoora Roewer; Dentobunus Roewer, Diangathia 412 7 Regional Arachnogeography Roewer, Echinobunus Roewer; Euceratobunus Roewer, 1923; Eugagrella Roewer; Eusclera Roewer, Euzaleptus Roewer; Gagrella Stoliczka; Gagrellenna Roewer; Gagrellina Roewer; Gagrellissa Roewer; Gagrellopsis Sato et Suzuki; Gagrellula Roewer; Globulosoma Martens; Hamitergum Crawford; Harmanda Roewer; Harmandina Schenkel; Hehoa Roewer; Heterogagrella Roewer; Hexazaleptus Suzuki; Himaldroma Martens; Himalzaleptus Martens; Hologagrella Roewer; Hypogrella Roewer; Koyamaia Suzuki; Marthana Thorell; Melanopa Thorell; Melanopella Roewer; Melanopula Roewer; Metadentobunus Roewer; Metahehoa Suzuki; Metasyleus Roewer; Metaverpulus Roewer; Metazaleptus Roewer, Microzaleptus Roewer, Neogagrella Roewer, Nepalgrella Martens, Nepalkanchia Martens, Obigrella Roewer, Octozaleptus Suzuki, Oobunus Kishida, Orissula Roewer, Padangrella Roewer, Palniella Roewer, Paradentobunus Roewer, Paragagrella Roewer, Paragagrellina Schenkel, Paraumbogrella Suzuki, Pergagrella Roewer, Pokhara Suzuki, Prodentobunus Roewer, Psathyropus L. Koch, Pseudarthromerus Karsch, Pseudogagrella Redikorzev, Pseudomelanopa Suzuki, Pseudosystenocentrus Suzuki, Sarasinia Roewer, Sataria Roewer, Scotomenia Thorell, Sericicorpus Martens, Sinadroma Roewer, Syleus Thorell, Syngagrella Roewer, Systenocentrus Simon, Tetraceratobunus Roewer, Toragrella Roewer, Umbogrella Roewer, Umbopilio Roewer, Verpulus Simon, Verrucobunus Roewer, Xerogrella Martens, Zaleptiolus Roewer, Zaleptulus Roewer, Zaleptus Thorell Laniatores From the Southeast Asia and Malayan Archipelago are recorded Laniatores from more than 50 genera and the families Assamiidae, Biantidae, Podoctidae, Epedanidae, Sandokanidae (= Oncopodidae), Phalangodidae, Zalmoxidae, Tithaeidae, and Petrobunidae. The families in bold are endemic for the Indomalayan Region. Most of the others are not found in the Palearctic Region, and other families (Phalangodidae) are considered by some researchers to be relicts in the Palearctic. Other specialists (Martens 1972) disagree with the relict character of European Phalangodidae. Among the authors having worked on the Southeast Asian Laniatores, we should not miss Tsurusaki (1995), Schwendinger (1992, 2006), Sharma et al. (2012), Sharma and Giribet (2011), Suzuki (1969, 1977a, 1977b, 1982, 1985), and the numerous papers by Roewer (1912, 1927, 1931, 1935, 1938, 1940, 1949), Thorell, and other authors. Endemic genera are: Fam. Assamiidae Assamiinae: Assamiella Roewer, 1912, Burma (one sp.); Neassamia Roewer, 1935, Thailand (one sp.); Pechota Roewer, 1935, Malacca (one sp.); Popassamia Roewer, 1940, Burma (one sp.); Tavoybia Roewer, 1935, Malacca (one sp.). Dampetrinae: Cadomea Roewer, 1940, Malaysia (one sp.); Dongmolla Roewer, 1927, Vietnam (one sp.); Mermerus Thorell, 1876, Java, Borneo (two spp.); Nothippulus Roewer, 1923, Vietnam (one sp.); Nothippus Thorell, 1890, 7.23 Southeast Asia (Burma, Thailand, Indochina, Peninsular Malaysia, China South… 413 Sumatra, Malacca (three spp.); Pahangius Roewer, 1935, Malacca (one sp.); Paradampetrus Giltay, 1930, Sumatra (one sp.); Sudaria Roewer, 1923, Sumatra, Simalur, Sulawesi (four spp.) Fam. Podoctidae Dongmoa Roewer, 1927, Vietnam; Heteroibalonius Goodnight et Goodnight, 1947 (one sp.); Mesoceratula Roewer, 1949 (one sp.); Podoctellus Roewer, 1949, Malaysia (Johore) (one sp.); Podoctis Thorell, 1890, Pinang (one sp.); Sibolgia Roewer, 1923, Malaya (one sp.); Stobitus Roewer, 1949, Malaya (one sp.) Fam. Petrobunidae Petrobunus Sharma et Giribet, 2011 – Philippines (three spp.) Fam. Tithaeidae – 38 spp Istithaeus Roewer, 1949, Borneo; Kondosus Roewer, 1949, Borneo; Metatithaeus Suzuki, 1969, Borneo; Sterrhosoma Thorell, 1891, Sumatra; Tithaeomma Roewer, 1949, Burma; Tithaeus Thorell, 1890, Burma, Thailand, Malaya, Singapore, Sumatra, Krakatau, Java, Borneo, Sarawak, Timor Fam. Epedanidae Epedaninae: Alloepedanus Suzuki, 1985, Thailand (one sp.); Caletorellus Roewer, 1938, Thailand (one sp.); Epedanidus Roewer, 1945, Malaysia (Perak) (one sp.); Euepedanus Roewer, 1915, Thailand, Malacca (seven spp.); Heteroepedanus Roewer, 1912 (two spp.); Paratakaoia Suzuki, 1985, Thailand (two spp.); Plistobunus Pocock, 1903, Hong Kong, Hainan Island (two spp.); Pseudoepedanus Suzuki, 1969 (one sp.); Pseudomarthana P. D. Hillyard, 1985, Malaysia (one sp.); Thyreotus Thorell, 1889, Burma (two spp.); Toccolus Roewer, 1927, Vietnam (“Tonking”), etc. (three spp.); Zepedanulus Roewer, 1927, Malacca, Thailand, etc. (four spp.) Acrobuninae: Heterobiantes Roewer, 1912, Hong Kong (one sp.); Paracrobunus Suzuki, 1977 (two spp.) Sarasinicinae: Gintingius Roewer, 1938, Pahang (Malaya) (one sp.); Panticola Roewer, 1938 (placement is uncertain), Malacca (one sp.); Pasohnus Suzuki, 1976, was in Phalangodidae (one sp.); Sembilanus Roewer, 1938, Malacca (one sp.); Siponnus Roewer, 1927, Pulau Pinang (one sp.); Sungsotia Tsurusaki, 1995, Vietnam (one sp.); Tonkinatus Roewer, 1938, Vietnam (Tonking) (one sp.) Incertae sedis Buparellus Roewer, 1949 – Burma, Thailand (four spp.) Fam. Zalmoxidae Zalmoxis Soerensen is not endemic, but this is the only genus of Zalmoxidae in the Old World, with ca. 15 spp. in South Asia, including the Philippines (Sharma et al. 2012). 414 7 Regional Arachnogeography Fam. Sandokanidae (Oncopodidae) – 71 spp. Sandokan Thorell, 1876, Southeast Asia (10 spp.); Gnomulus Thorell, 1890, Southeast Asia, India, South China (53 spp.); Caenoncopus Martens et Schwendinger, 1998, Sumatra (three spp.); Palaeoncopus Martens et Schwendinger, 1998, Sumatra (three spp.); Biantoncopus Martens et Schwendinger, 1998, Leyte, Philippines (one sp.); Martensiellus Schwendinger, 2006, Borneo (one sp.) The distribution of Sandokanidae appears to be governed principally by limitations of this group’s dispersal ability. Four other laniatorid families —Assamiidae, Epedanidae, Podoctidae, and Zalmoxidae—are distributed throughout Sundaland, but all of these have greater range than Sandokanidae, and frequently demonstrate clear dispersal events (Giribet and Kury 2007). The restriction of Sandokanidae to Sundaland and the Philippines is suggestive of diversification in accordance with breakup of Sundaland’s components.” (Sharma and Giribet 2009) The opilionids of Kuril Islands have been analyzed by Tsurusaki and Crawford (2001), the ones from the Ryukyus – by Suzuki (1971, 1973). Amblypygi The Amblypygi are represented in Southeast Asia by at least 22 spp. of 5 genera (Harvey). Fam. Charinidae Catageus Thorell, 1889 (= Stygophrynus Kraepelin, 1895) – Burma, Thailand, Laos, Cambodia, Vietnam, Indonesia, Malaysia (C. pusillus Thorell, 1889= Stygophrynus cavernicola (Thorell, 1889), C. berkeleyi (Gravely, 1915), C. brevispina (Weygoldt, 2002), C. cerberus (Simon, 1901), C. dammermani (Roewer, 1928), C. longispina (Gravely, 1915), C. moultoni (Gravely, 1915), C. orientalis (Seiter et Wolf, 2017), C. sunda (Rahmadi et Harvey, 2008) Charinus Simon, 1892 – Ch. bengalensis (Gravely, 1911); Ch. pakistanus Weygoldt, 2005 Sarax Simon, 1892 – S. brachydactylus Simon, 1892 (Malaysia, Cambodia); S. buxtoni (Gravely, 1915) (Malaya, Singapore); S. davidovi Fage, 1946 (Vietnam, Laos, Cambodia); S. rimosus (Simon, 1901) (Malaya); S. singaporae Gravely, 1911 (Singapore); S. cavernicola Rahmadi et al., 2010; S. mardua Rahmadi, 2010; S. sangkulirangensis Rahmadi et al., 2010; S. yayukae Rahmadi et al., 2010 (the last four from Kalimantan) Weygoldtia Miranda, Giupponi, Prendini et Scharff, 2018 – one sp. W. davidovi (Fage, 1946) – Laos, Cambodia, Vietnam Fam. Charontidae Charon Karsch, 1879 – Ch. grayi (Gervais, 1842) (s. lato) (Singapore, Malaysia) 7.23 Southeast Asia (Burma, Thailand, Indochina, Peninsular Malaysia, China South… 415 Map 7.12 Distribution of Amblypygi in Asia and East Africa ■ – Phrynichus ● – Damon □ – Sarax X – Charinus – Catageus ! – Phrynus ♦ – Stygophrynus ◊ – Charon Ο – Euphrynichus Thelyphonida (Uropygi) The early papers of Oates (1889, 1890), Gravely (1912), and Hirst and Pocock (1902) described from Burma 13 out of the 17 Asiatic species in Hypoctonus Thorell. Teltus vanoorti Speijer, 1936, was described as endemic genus and species for Hainan, but Haupt (1996) synonymized the genus Teltus with Typopeltis Pocock. From South China have been recorded three species, Hypoctonus carmichaeli Gravely, H. granosus Pocock, and Typopeltis cantonensis Speijer, and from Taiwan and the Ryukyu T. crucifer Pocock. Another Typopeltis, living on the Ryukyus, is T. stimpsoni (Wood) (Yoshikura 1973; Haupt and Daxiang Song 1996a). Other Southeast Asian Uropygi are known from Bangladesh (3 spp.), Burma (12 spp.), Thailand (8 spp.), Laos (2 spp.), the Philippines (8 spp.), Indonesia (25 spp.), 416 7 Regional Arachnogeography Malaysia (7 spp.), Cambodia (3 spp.), Singapore (3 spp.), and Vietnam (5 spp.) (Haupt 2004, Kraepelin 1897, Rowland 1973, Tarnani 1894, 1900, Thorell 1888, Harvey 2013d). Haupt (2009a) synonymized Abaliella Strand, Minbosius Speijer, and Tetrabalius Thorell with Thelyphonus Latreille. The newest calculation (Harvey 2013d) lists for Southeast Asia (east of India and south of Yangtze, but including Bangladesh, the Philippines, Taiwan, and the Ryukyus) the genera Uroproctus Pocock (1 sp.), Hypoctonus Thorell (15 spp.), Typopeltis Pocock (12 spp.), Ginosigma Speijer (2 spp.), Glyptogluteus Rowland (1 sp.), Thelyphonoides Krehenwinkel et al. (1 sp.), and Thelyphonus Latreille (30 spp.) or in total 62 of the 110 spp. of Thelyphonida. Map 7.13 Distribution of order Thelyphonida (Uropygi) in Asia ● – Glyptogluteus Rowland ■ – Typopeltis Pocock ∆ – Labochirus Pocock ▼– Uroproctus Pocock ▲ – Hypoctonus Thorell ◊ – Thelyphonus Latreille O – Thelyphonoides Krehenwinkel et al. – Minbosius Speijer – Ginosigma Speijer 7.23 Southeast Asia (Burma, Thailand, Indochina, Peninsular Malaysia, China South… 417 Schizomida The list of Schizomida of South Asia is relatively long (34 spp. of 11 genera), but several of them require additional study for more accurate generic placement (Bastawade 2004, Brignoli 1974, Gravely 1911, 1912, 1924, Hansen and Soerensen 1905, Harvey 2013 , Kraepelin 1912, Reddell and Cokendolpher 1995, Cokendolpher et al. 2010, Rémy 1946, Shimojana 1981, Thorell 1889). It includes: Apozomus brignolii Reddell et Cokendolpher, 2000) – Marshall Islands (Ailuk) A. buxtoni Gravely, 1915 – Sri Lanka A. daitoensis (Shimojana, 1981) – Ryukyus (Minami-Daitô-jima) (endemic sp.) A. sauteri (Kraepelin, 1912) – Taiwan, Vietnam, Ryukyu Islands A. termitarium Cokendolpher, Sissom, et Reddell, 2010 (Malaysia, Taman Negara) A. yamasakii (Cokendolpher, 1988) – Taiwan A. zhensis Chen and Song, 1996 – China (Zhejiang) Bamazomus siamensis (Hansen, 1905) – Thailand, Hong Kong, Ryukyu Islands, Hawaii (Oahu) B. pileti (Brignoli, 1974) – W. Malaysia (endemic sp.) Burmezomus cavernicola (Gravely, 1912) – Burma Clavizomus claviger (Hansen, 1905) – Singapore, W. Malaysia Javazomus oculatus Reddell et Cokendolpher, 1995 – Java Neozomus tikaderi Reddell et Cokendolpher, 1995 – India Oculozomus biocellatus (Sissom, 1980) – Sumatra Orientzomus luzonensis (Hansen, 1905) – Philippines (Luzon) O. ralik Cokendolpher et Reddell, 2000 – Marshall Islands (Jibu) O. sawadai (Kishida, 1930) – Bonin Islands Ovozomus lunatus (Gravely, 1911) – India (Kolkata), Réunion, Seychelles O. peradenyiensis (Gravely, 1911) – Sri Lanka “Schizomus” modestus (Hansen, 1905) – W. Malaysia [also in Papua New Guinea, incl. New Britain] “S.” cambridgei (Thorell, 1889) – Burma “S.” procerus (Hansen, 1905) – Singapore “S.” peteloti (Rémy, 1946) – S. Vietnam (endemic sp.) “S.” vittatus Gravely, 1911 – Sri Lanka “S.” greeni Gravely, 1912 – Sri Lanka “S.” chaibassicus Bastawade, 2002 – India “S.” chalakudicus Bastawade, 2002 – India “S.” formicoides Fernando, 1957 – Sri Lanka “S.” crassicaudatus (O.P.-Cambridge, 1872) – Sri Lanka “S.” kharagpurensis Gravely, 1912 – India (West Bengal) “S.” perplexus Gravely, 1915 – Sri Lanka Trithyreus grassii (Thorell, 1889) – Burma (endemic sp.) T. sijuensis (Gravely, 1924) – India (Meghalaya) Zomus bagnalii (Jackson, 1908) – Singapore, W. Malaysia, Sarawak [also Kew Gardens in England] 418 7 Regional Arachnogeography Map 7.14 Distribution of order Schizomida in Asia Apozomus – ■ Bamazomus – ▲ Burmezomus – ▼ Clavizomus – ● Javazomus – Neozomus – ♣ Oculozomus – ☺ Orientzomus – ♠ Ovozomus – ♦ Schizomus – ◊ Trithyreus – □ Zomus – Araneae From continental Southeast Asia are known spiders from 58 families (none of them endemic) (Deeleman-Reinhold 1995, 2000, Jäger 2005, Meng et al. 2008, Tang and Li 2010a, b, Wang et al. 2010). From the Peninsular Malaysia, Norma-Rashid and Li (2009) list 425 species in 42 families and 238 genera of spiders. From the tiny Singapore, Song et al. (2002) announced the existence of 310 species of spiders of 186 genera and 39 families. From the list of Chinese spiders of Song et al. (1999) and the papers published afterward, considerable part consist of South Chinese species. Some endemic genera are: 7.23 Southeast Asia (Burma, Thailand, Indochina, Peninsular Malaysia, China South… Fam. Liphistiidae Liphistius Schiødte, 1849 – Southeast Asia (47 species) Nanthela Haupt, 2003 – Hong Kong, Vietnam (two spp.) Fam. Theraphosidae Ornithoctonus Pocock, 1892 – Myanmar, Thailand (three spp.) Fam. Dipluridae Leptothele Raven et Schwendinger, 1995 – Thailand (one sp.) Fam. Theridiosomatidae Chthonopes Wunderlich, 2011 – Laos (two spp.) Luangnam Wunderlich, 2011 – Laos (one sp.) Fam. Nemesiidae Atmetochilus Simon, 1887 – Myanmar (two spp.) Fam. Cyrtaucheniidae Angka Raven et Schwendinger, 1995 – Thailand Fam. Pisauridae Ilipula Simon, 1903 – Vietnam (one sp.) Fam. Liocranidae Jacaena Thorell, 1897 – Thailand, Laos, Myanmar, China (11 spp.) Fam. Tetrablemmidae Bacillemma Deeleman-Reinhold, 1993 – Thailand (one sp.) Fam. Dictynidae Penangolyna Wunderlich, 1995 – Malaysia (one sp.) Fam. Araneidae Perilla Thorell, 1895 – Myanmar, Vietnam, Malaysia (one sp.) Fam. Agelenidae Acutipetala Dankittipakul et Zhang, 2008 – Thailand (two spp.) Fam. Linyphiidae Thainetes Millidge, 1995 – Thailand (one sp.) Thaiphantes Millidge, 1995 – Thailand (two spp.) Fam. Oonopidae Bannana Tong et Li, 2015 – Yunnan (two spp.) 419 420 7 Regional Arachnogeography Fam. Selenopidae Siamsporops Dankittipakul et Corronca, 2009 – Thailand, Malaysia Fam. Zodariidae Euryeidon Dank. et Jocqué, 2004 – Thailand (six spp.) Heliconilla Dank et al., 2012 – Thailand, Vietnam, Malaysia, Singapore, Myanmar, Laos, China (nine spp.) Heradion Danki et Jocqué, 2004 – Thailand, Vietnam, Malaysia, Laos, China (ten spp.) Malayozodarion Ono et Hashim, 2008 – Malaysia (one sp.) Fam. Gnaphosidae Allomicythus Ono, 2009 – Vietnam Aracus Thorell, 1887 – Myanmar Fam. Lycosidae Zantheres Thorell, 1887 – Myanmar Fam. Thomisidae Boliscodes Simon, 1909 – Vietnam (one sp.) Haplotmarus Simon, 1909 – Vietnam (one sp.) Ocyllus Thorell, 1887 – Myanmar (two spp.) Pothaeus Thorell, 1895 – Myanmar (one sp.) Tagulinus Simon, 1903 – Vietnam (one sp.) Fam. Salticidae Angustaea Szombathy, 1915 – Singapore Ceclusa Thorell, 1895 – Myanmar The subfamily Onomastinae is endemic for the Indomalayan Region. Acariformes Prostigmata. Ref.: Mąkol and Gabryś (2005) Fam. Eutrombidiidae Caecothrombium deharvengi Mąkol et Gabryś, 2005 – Vietnam Oribatida According to the monograph of Vu Quang Manh (2015), in Vietnam have been recorded 320 spp. of Oribatida, zoogeographically zonated as follows: Oriental (60.3%), Palearctic (12.2%), Cosmopolitan (10.6%), Afrotropical (6.9%), Australian (5.0%), Neotropical (3.8%), Nearctic (0.9%), and Pacific (0.3%). Important contributions are made by Aoki (1965, 1967). 7.24 Taiwan 421 Parasitiformes Ixodida Petney et al. (2007) announced the presence of 104 spp. of ticks (Ixodida) of 12 genera in Southeast Asia. 7.24 7.24.1 Taiwan Geography, General Zoogeography, and Paleogeography Separated from the Asian continent by the 160 km-wide Taiwan Strait, the main island of the group is 394 km long and 144 kilometers wide. The southern end of the Ryukyu Islands of Japan is directly to the east; the Batanes Islands of the Philippines lie to its south across the Bashi Channel. The mountainous island spans the Tropic of Cancer and is covered by tropical and subtropical vegetation. Other minor islands and islets of the group include the Penghu Islands (Pescadores), Green Island, and Orchid Island. The area of the main islands is 35,801 km2 (land 32,260 km2, together with the smaller islands – 35,980 km2). Taiwan’s highest point is Yushan at 3952 meters, and there are five other peaks over 3500 meters. 7.24.2 Arachnogeography This island, recently detached from the continent (Wang Hongzhen, Ed., 1985), counts among its inhabitants many animals like mammals, snakes and others, clearly inherited from the mainland (Zhang and Zhao 1978). The high mountains (almost 4000 m) and the preserved vegetation in the national parks harbor many interesting arachnids – Thelyphonida (Uropygi), Schizomida, Pseudoscorpiones, Opiliones, and Araneae – and mites. Lacking (or not yet found) are Palpigradi, Ricinulei, Amblypygi, Holothyrida, and Opilioacarida. One amblypygid (Charon grayi) is known from a cave on the small Orchid Island or Botel Tobago (Kano, 1937). Thelyphonida (Uropygi) Whip scorpions from Taiwan are known as early as from the paper of Schwangart (1906). Fam. Thelyphonidae – Typopeltis crucifer Pocock, 1894 Schizomida Only two species (one of them endemic) are known so far from Taiwan (Cokendolpher 1988, Kraepelin 1911, Yamasaki and Shimojana 1974). 422 7 Regional Arachnogeography Fam. Hubba Apozomus sauteri rdiidae Apozomus sauteri (Kraepelin, 1912) – Taiwan, Vietnam, Ryukyu Islands yamazakii (Cokendolpher, 1988) – Taiwan (endemic sp.) Pseudoscorpiones In Taiwan are recorded ten spp. of pseudoscorpions of eight genera and the families Chthoniidae, Neobisiidae, Geogarypidae, Olpiidae, Cheiridiidae, Cheliferidae, Chernetidae, and Withiidae. Opiliones The opilionid fauna of Taiwan consists of 13 genera and the families Sclerosomatidae, Epedanidae, Phalangodidae, and Podoctidae (Roewer 1911, 1915, 1938, Suzuki 1944, 1974, 1977b – list of species, Tsurusaki 1991). Cyphophthalmi – not found Eupnoi Fam. Phalangiidae – Oligolophus C.L. Koch Fam. Sclerosomatidae – Gagrella Stoliczka, Pseudomelanopa Suzuki, Leiobunum C.L. Koch, Pseudogagrella Redikorzev, Psathyropus L. Koch (incl. Metagagrella), Metadentobunus Roewer, Nelima Roewer Dyspnoi – not known Laniatores Fam. Phalangodidae – Tokunosia Suzuki Fam. Epedanidae – Funkikoa Roewer, Takaoia Roewer, Kilungius Roewer, Mosfora Roewer (endemic genera in bold) Fam. Podoctidae – Metapodoctis Roewer, Lomanius Roewer Araneae From Taiwan are registered (Lee 1964, 1966; Chu and Okuma 1974, 1975, Chen 1996, Song D. X. and M. S. Zhu 1997, Song D. X. et al. 1999, Ono 2007) spiders of 37 families. All of them are known also in continental China, except for Dipluridae and Dysderidae. From the Taiwan spiders, there are five families of Mygalomorphae (Atypidae, Ctenizidae, Hexathelidae, Dipluridae, and Theraphosidae). The remaining 32 families are members of Araneomorphae. Endemic genera: Fam. Agelenidae: Femoracoelotes Wang, 2002 (two spp.) Fam. Anapidae: Enielkenie Ono, 2007 (one sp.) Fam. Selenopidae: Pakawops Crews et Harvey, 2011 (one sp.) Acariformes Oribatida – Tseng (1982, 1984) Parasitiformes Ixodida – The checklist of Robbins (2005) announced the existence in Taiwan of 33 spp. of Ixodida (4 Argasidae and 29 Ixodidae). Ixodes kuntzi Hoogstraal et Kohls, 1965, seems endemic to Taiwan. 7.25 The Malay Archipelago 7.25 7.25.1 423 The Malay Archipelago Geography, General Zoogeography, and Paleogeography Situated between the Indian and Pacific Oceans, the group of over 25,000 islands is the largest archipelago by area in the world. It includes Indonesia, the Philippines, Singapore, Brunei, East Malaysia, and East Timor. The islands of New Guinea are not included in definitions of the Malay Archipelago. In the present study, the volume of the notion “Malay Archipelago” includes only Indonesia (without Papua), East Timor, and northern Borneo (Sarawak, Sabah, and Brunei). The biggest islands are as follows: Borneo – area 743,330 km2, highest point Kinabalu (4095 m) Sumatra – area 473,481 km2, highest point Kerinci (3805 m) Sulawesi – area 174,600 km2, highest point Rantemario (3478 m) Java – area 132,187 km2, highest point Semeru (3676 m) Bali – area 5,633 km2, highest point Agung (3142 m) Lombok – area 4,725 km2, highest point Rinjani (3726 m) Flores – area 13,540 km2, highest point Poco Mandasawu (2370 m) Timor – area 30,777 km2, highest point Tatamailau (2963 m) We have to consider the analysis of Moss and Wilson (1998) concerning the biogeographical implications of the events on Sulawesi and Borneo in the Tertiary. According to them Wallacea is a biogeographical area situated between Asiatic and Australian flora and fauna with organisms of high level of endemicity. The land connection between Borneo and continental Southeast Asia might have existed during important section of Tertiary and could have allowed migrations of species. Western Sulawesi has been connected with East Borneo by Late Cretaceous and Early Eocene (more than 50 Ma) with option of dispersion of fauna. The ophiolites of east Sulawesi have been accreted to Sulawesi in the Late Oligocene, which resulted in a more extensive land of the large (174,600 km2) island. Microcontinental fragments accreted onto eastern Sulawesi in the Miocene to Pleistocene may have been emergent as they drifted towards Sulawesi and allowed island hopping or rafting for biota of Australian affinity. Island hopping routes for the dispersal of organisms between Borneo-Sulawesi and the Philippines may have existed along volcanic arcs, such as the long-lived North Sulawesi arc. (Moss and Wilson 1998) Here is the timing of events in the distribution of the elements of land in the area (according to Audley-Charles, 1984): 1. Australia/New Guinea splits from Antarctica (ca.53 Ìa). 2. Postulated formation of the Philippines by collision of an Asian continental fragment with an island arc (Îligocene). 3. Possible land connection(s) across Makassar Strait (mid-Miocene). 4. Collision between New Guinea and a Tertiary island arc (ca. 15 Ìa). 424 7 Regional Arachnogeography 5. Submarine collision between Gondwana (Sula Peninsula) and Laurasia at or near east Sulawesi (ca. 15 Ìa). 6. Island chain established between east Sulawesi and Australia (Late Miocene to Late Pliocene). 7. Collision between parts of Gondwanic Outer Banda Arc and Laurasian (volcanic) Inner Banda Arc (Late Miocene to Early Pliocene). 8. Gulf of Bone opens (about the same time). 9. Probable land connection(s) across south Makassar Strait (from Late Pliocene). Some sources: Golonka et al. (2006), Groves (1975), Keast (1983), Katili (1971, 1975, 1978), Mayr (1944), Wallace (1869), Weber (1899), Whitmore (Ed.) (1987), Wilson and Moss (1999) 7.25.2 Arachnogeography Zoogeographically one of the most remarkable areas is the so-called Wallacea, the transitional islands with elements of the Indomalayan and Papuan-Australian fauna. Its “mainland” is Sulawesi (Celebes), explored by the Swiss Fritz and his cousin Paul Sarasin in 1893–1902 (and later by many others). Roewer described the opilions collected by the remarkable naturalists. The Malay Archipelago is the richest area of fauna in the world. What concerns the arachnids, in the archipelago are recorded all the orders of Arachnida, except for Ricinulei, Holothyrida, and Opilioacarida. Remarkable is the absence of Holothyrida, well represented in the nearby New Guinea. Palpigradi Koeneniodes berndti Condé, 1988, was described from Malaysia (Borneo). From Indonesia are known seven species from Java, Sulawesi, and Sumatra (Condé 1989, 1990, 1992, 1994). Some of them are endemic (Eukoenenia maros, E. lienhardi (also in Brunei and Singapore), E. paulinae, Prokoenenia celebica, P. javanica), and others are known from Madagascar (Koeneniodes madecassus, K. frondiger). They belong to the families Eukoeneniidae (Eukoenenia Börner) and Prokoeneniidae (Prokoenenia Börner). Scorpiones In Indonesia are registered 29 spp. of 6 genera of 4 families of scorpions (see Table 7.2) – Buthidae, Chaerilidae, Hormuridae, and Scorpionidae. An eyeless forest litter scorpion was described from Halmahera (Chaerilus telnovi Lourenço, 2009), followed by another species of the same genus Chaerilus spinatus Lourenço et Duhem, 2010, from the cave Batu Lubang on the same island. Both are endemic to Halmahera. 7.25 The Malay Archipelago 425 Solifugae Fam. Melanoblossiidae, subfam. Dinorhacinae The genus Dinorhax Simon, 1879, is the only solpugid forming the eastern limit of the order. The species Dinorhax rostrumpsittaci (Simon, 1877) is known from Vietnam and Indonesia (Maluku). Pseudoscorpiones In Indonesia (incl. W. New Guinea) have been registered 88 spp. of pseudoscorpions and in Malaysia (incl. Peninsular), 38 spp. Altogether, in Malay Archipelago are recorded 45 genera of 14 families: Chthoniidae, Chernetidae, Cheliferidae, Atemnidae, Garypidae, Garypinidae, Geogarypidae, Hyidae, Olpiidae, Parahyidae, Pseudochiridiidae, Syarinidae, Tridenchthoniidae, and Withiidae (Beier 1932, 1935, 1952, 1953, 1954, Muchmore 1972, Simon 1899, Thorell 1883, Tullgren 1907, With 1906). Endemic genera for Malay Archipelago are: Fam. Cheliferidae: Aporochelifer Beier, 1953 – Flores (one sp.) Fam. Chernetidae: Chiridiochernes Muchmore, 1972 – Sulawesi (one sp.) Opiliones Clouse (2012), Loman (1892), Martens and Schwendinger (1998), Roewer (1913, 1914a, 1927, 1931, 1938, 1949), Schwendinger (2006), Shear (1993a), Thorell (1882, 1890, 1891) Cyphophthalmi Three genera of fam. Stylocellidae are recorded in Malay Archipelago (19 spp.) Leptopsalis Thorell, 1882 – Sumatra (three spp.), Java (two spp.), Sulawesi (four spp.) Stylocellus Westwood, 1874 – Sumatra (one sp.) Miopsalis Thorell, 1890 – Borneo (nine spp.) Eupnoi (Roewer, 1913, 1931) Fam. Phalangiidae – Mitopiella Banks, 1930 Fam. Sclerosomatidae – Altobunus Roewer, 1910; Dentobunus Roewer, 1910; Echinobunus Roewer, 1912; Eugagrella Roewer, 1910; Euzaleptus Roewer, 1911; Gagrella Stoliczka, 1869; Gagrellina Roewer, 1913; Gagrellissa Roewer, 1931; Gagrellula Roewer, 1910; Marthana Thorell, 1891; Melanopella Roewer, 1931; Metazaleptus Roewer, 1912; Neogagrella Roewer, 1913; Paragagrella Roewer, 1912; Zaleptus Thorell, 1876 Endemic genera are: Mitopiella Banks, 1930, Borneo (one sp.); Gagrellina Roewer, 1913, Sulawesi (one sp.) Gagrellissa Roewer, 1931 – Sunda Islands (one sp.) Dyspnoi – not known 426 7 Regional Arachnogeography Laniatores They are represented in Malay Archipelago (s.str.) by the families Biantidae, Podoctidae, Epedanidae, and Sandokanidae (= Oncopodidae preocc.). Endemic genera are: Fam. Epedanidae Epedaninae Epedanulus Roewer, 1913 – Sulawesi (one sp.) Epedanus Thorell, 1876 – Borneo, Java, Sarawak, Pinang, Sumbawa (eight spp.) Heteroepedanus Roewer, 1912 – (two spp.) Lobonychium Roewer, 1938 – Borneo (one sp.) Metepedanulus Roewer, 1913 – Sulawesi, Borneo (two spp.) Metepedanus Roewer, 1912 – Borneo (two spp.) Nanepedanus Roewer, 1938 – Borneo (one sp.) Parepedanulus Roewer, 1913 – Sulawesi (two spp.) Pseudoepedanus Suzuki, 1969 – (one sp.) Acrobuninae Acrobunus Thorell, 1891 – Sumatra, Borneo (three spp.) Anacrobunus Roewer, 1927 – Riau Islands (one sp.) Harpagonellus Roewer, 1927 – Sumatra (one sp.) Paracrobunus Suzuki, 1977 – Palawan, Philippines (two spp.) Sarasinicinae Acanthepedanus Roewer, 1912 – Sumatra (one sp.) Albertops Roewer, 1938 – Borneo (one sp.) Delicola Roewer, 1938 – Sumatra (one sp.) Koyanus Roewer, 1938 – Borneo (one sp.) Kuchingius Roewer, 1927 – Borneo (one sp.) Padangcola Roewer, 1963 – should possibly be in Epedaninae [Sumatra] (one sp.) Punanus Roewer, 1938 – Borneo (one sp.) Sarasinica Strand, 1914 – Borneo, Sulawesi (four spp.) Sinistus Roewer, 1938 – Borneo (two spp.) Siponnus Roewer, 1927 – Pulau Pinang (one sp.) Incertae sedis Beloniscellus Roewer, 1912 – Indonesia (six spp.) Beloniscops Roewer, 1949 – Sumatra (two spp.) Belonisculus Roewer, 1923 – Simalur Island (Sumatra) (one sp.) Beloniscus Thorell, 1891 – Singapore, Sumatra, Simalur Island, Java, “Sunda Islands” (14 spp.) Parabupares Suzuki, 1982 – Indonesia (one sp.) Sotekia Suzuki, 1982 – Indonesia (one sp.) 7.25 The Malay Archipelago 427 Fam. Sandokanidae [= Oncopodidae, praeoccup.] Caenoncopus Martens et Schwendinger, 1998 – Sumatra (three spp.) Martensiellus Schwendinger, 2006 – Borneo (one sp.) Palaeoncopus Martens et Schwendinger, 1998 – Sumatra (three spp.) Fam. Podoctidae (incl. Erecananinae and Ibaloniinae) Baramella Roewer, 1949 – Borneo (one sp.) Baramia Hirst, 1912 – Sarawak, Indonesia (four spp.) Baso Roewer, 1923 – Sumatra (one sp.) Basoides Roewer, 1949 – Sumatra (one sp.) Dino Loman, in Weber 1892 – Sumatra (one sp.) Gargenna Roewer, 1949 – Indonesia (one sp.) Heteroibalonius Goodnight et Goodnight, 1947 – Papua (one sp.) Mesoceratula Roewer, 1949 – (one sp.) Orobunus Goodnight et Goodnight, 1947 – Papua (one sp.) Pentacros Roewer, 1949 – Indonesia (one sp.) Gaditusa Roewer, 1949 – Borneo (one sp.) Idjena Roewer, 1927 – Java (one sp.) Japetus Roewer, 1949 – Borneo (one sp.) Lejokus Roewer, 1949 – Borneo (one sp.) Lundulla Roewer, 1927 – Borneo (one sp.) Podoctis Thorell, 1890 – Pinang (one sp.) Podoctomma Roewer, 1949 – Java (one sp.) Podoctops Roewer, 1949 – Sumatra (one sp.) Trencona Roewer, 1949 – Borneo (one sp.) Trigonobunus Loman, 1894 – Borneo (one sp.) Amblypygi From Malay Archipelago have been recorded until 2016 11 (may be more; see Harvey and West 1998) species of Amblypygi from 4 genera and 3 families (Gravely 1915, Harvey 2002, Kraepelin 1895, Rahmadi and Harvey 2008, Roewer 1928, Thorell 1888). Many of them were found in caves. No endemic genera, some endemic species. Quite puzzling was, however, to find in the cave Batu Cermin on the Flores Island the new species Phrynus exsul Harvey, 2002 – the first representative of the American family Phrynidae in the Old World. Rahmadi et al. (2010) added four new species of Sarax from Borneo. Fam. Charinidae Sarax Simon – S. cavernicola Rahmadi, Harvey, et Kojima, 2010; S. sangkulirangensis Rahmadi, Harvey, et Kojima, 2010; S. mardua Rahmadi, Harvey, et Kojima, 2010; S. yayukae Rahmadi, Harvey, et Kojima, 2010; S. sarawakensis (Thorell, 1888) (all Borneo or Kalimantan); S. javensis (Gravely, 1915) (Java) 428 7 Regional Arachnogeography Fam. Charontidae Charon Karsch – Ch. grayi (Gervais, 1842) (syn.? C. beccarii Thorell, 1888, and C. subterraneus Thorell, 1888), Indonesia (Ambon, etc.) Stygophrynus Kraepelin – S. (S.) dammermani Roewer, 1928 (Java); S. (S.) sunda Rahmadi et Harvey, 2008 (Java); S. (Neocharon) moultoni Gravely, 1915 (Sumatra, Kalimantan) Fam. Phrynidae: Phrynus Lamarck – Ph. exsul Harvey, 2002 (Flores) Thelyphonida (Uropygi) From Indonesia and Malaysian Borneo have been described or recorded 15 spp. of Thelyphonus Latreille, Hypoctonus gastrostictus Kraepelin, Tetrabalius nasutus Thorell, T. seticauda (Doleschall), Minbosius manilanus (Koch), Ginosigma lombokensis Speijer, and Chajnus renschi Speijer (endemic genus for the Sunda Islands). In conclusion, from the species from the Philippines and the three Labochirus and one Uroproctus from India and Sri Lanka, ca. 60 spp. of Uropygi are represented in the “Indomalayan Region” (without Melanesia – New Guinea, the Solomon Islands, and Vanuatu). Haupt (2009a) synonymized the genera Minbosius, Tetrabalius, and Abaliella with Thelyphonus, so now we know from Malayan Archipelago 21 spp. of Thelyphonida from the genera Thelyphonus Latreille (18 spp.), Hypoctonus Thorell (1), Ginosigma Speijer (1), and Chajnus Speijer (1). All of them (except Thelyphonus manilanus) are endemic for the various islands of the archipelago (Kraepelin 1897, Speijer 1931, 1936, Tarnani 1901, Thorell 1888). Schizomida Schizomida from the Malay Archipelago (s.s.): Clavizomus sp. – Java Javazomus oculatus (Cokendolpher et Sites, 1988) – Java (end. genus) Oculozomus biocellatus (Sissom, 1980) – Sumatra (end. genus) Zomus bagnalii (Jackson, 1908) – Sarawak [also Singapore, W. Malaysia, Kew Gardens in England] According to Reddell and Cokendolpher (1995), unidentified Schizomida are known also from Ambon, Kalimantan, Sulawesi, and Krakatau Islands. Araneae From Malay Archipelago is registered rich fauna of spiders (DeelemanReinhold 1995, 2001, Prószyn’ski and Deeleman-Reinhold 2010, and others). According to the checklist of Stenchly (2011), from Indonesia and New Guinea have been registered 58 families of spiders with 505 genera and 1954 spp., 499 spp. being found only in New Guinea. Six of the 58 families (Dipluridae, Lamponidae, Micropholcommatidae, Nicodamidae, Stiphidiidae, Titanoecidae) are found only in New Guinea. 7.25 The Malay Archipelago Some endemic genera of spiders from the Malay Archipelago (s.str.) Fam. Theraphosidae Phormingochilus Pocock, 1895 – Borneo, Sumatra (five spp.) Fam. Liocranidae Sudharmia Deeleman-Reinhold, 2001 – Sumatra (three spp.) Fam. Miturgidae Tamin Deeleman-Reinhold, 2001 – Borneo, Sulawesi (two spp.) Fam. Oonopidae Myrmopopaea Reimoser, 1933 – Sumatra (one sp.) Plectoptilus Simon, 1905 – Java (one sp.) Xyphinus Simon, 1893 – Borneo, Singapore, Malaysia Fam. Oxyopidae Tapponia Simon, 1885 – Malaysia, Sumatra, Borneo (one sp.) Fam. Phrurolithidae Plynnon Deeleman-Reinhold, 2001 – Sumatra, Borneo (three spp.) Fam. Phyxelididae Vytfutia Deeleman-Reinhold, 1986 – Sumatra, Borneo (two spp.) Fam. Tetrablemmidae Borneomma Deeleman-Reinhold, 1980 – Borneo (two spp.) Lamania Lehtinen, 1981 – Borneo, Bali, Malaysia, Sulawesi (seven spp.) Maijana Lehtinen, 1981 – Java (one sp.) Sabahya Deeleman-Reinhold, 1980 – Borneo (two spp.) Singalangia Lehtinen, 1981 – Sumatra (one) Fam. Tetragnathidae Mitoscelis Thorell, 1890 – Java (one sp.) Neoprolochus Reimoser, 1927 – Sumatra (one sp.) Fam. Anapidae Borneanapis Snazell, 2009 – Borneo (one sp.) Fam. Cycloctenidae Galliena Simon, 1898 – Java (one sp.) 429 430 7 Regional Arachnogeography Fam. Clubionidae Scopalio Deeleman-Reinhold, 2001 – Borneo (one) Fam. Eutichuridae Calamoneta Deeleman-Reinhold, 2001 – Java, Sumatra (two spp.) Summacanthium Deeleman-Reinhold, 2001 – Sulawesi (two spp.) Fam. Symphytognathidae Anapogonia Simon, 1905 – Java (one sp.) Fam. Linyphiidae Caenonetria Millidge et Russell-Smith, 1992 – Borneo Dumoga Millidge et Russell-Smith, 1992 – Sulawesi (two spp.) Eordea Simon, 1899 – Sumatra (one sp.) Erigophantes Wunderlich, 1995 – Borneo (one sp.) Phyllarachne Millidge et Russell-Smith, 1992 – Borneo Piesocalus Simon, 1894 – Java (one sp.) Fam. Araneidae Actinacantha Simon, 1864 – Sumatra, Java Friula O.P.-Cambridge, 1896 – Borneo (one sp.) Plicatiductus Millidge et Russell-Smith, 1992 – Sulawesi Racata Millidge, 1995 – Krakatau (one sp.) Mitrager van Helsdingen, 1985 – Java Fam. Theridiidae Borneoridion Deeleman et Wunderlich, 2011 – Borneo (one sp.) Deelemanella Yoshida, 2003 – Borneo (one sp.) Helvidia Thorell, 1890 – Sumatra (one sp.) Fam. Thomisidae Musaeus Thorell, 1890 – Sumatra (one sp.) Narcaeus Thorell, 1890 – Java (one sp.) Nyctimus Thorell, 1877 – Sumatra, Sulawesi (one sp.) Pseudamyciaea Simon, 1905 – Java (one sp.) Reinickella Dahl, 1907 – Java Scopticus Simon, 1895 – Java (one sp.) Fam. Salticidae Stichius Thorell, 1890 – Sumatra (one sp.) Thianella Strand, 1907 – Java (one sp.) Tisaniba Zhang et Maddison, 2014 (six spp.) Taivala Peckham et Peckham, 1907 – Borneo (one sp.) 7.26 The Philippines 431 Map 7.15 Outline of the Malayan Archipelago 7.26 7.26.1 The Philippines Geography, General Zoogeography, and Paleogeography The Philippines is an archipelago of 7107 islands with a total land area, including inland bodies of water, of approximately 300,000 km2. It is located between 116° 40′ and 126° 34′ E and 4° 40′ and 21° 10′ N and is bordered by the Philippine Sea to the east, the South China Sea to the west, and the Celebes Sea to the south. The island of Borneo is located a few hundred kilometers southwest, and Taiwan is located directly to the north. Most of the mountainous islands are covered in tropical rain forest and volcanic in origin. The highest mountain is Mount Apo (in Mindanao, 2954 m). Forest cover declined from 70% of the country’s total land area in 1900 to about 18.3% in 1999. Important paper on the zoogeography of the Philippines was written by Dickerson (1928), who introduced the word Wallacea. 7.26.2 Arachnogeography The archipelago is rich in Arachnida (the orders Palpigradi, Scorpiones, Pseudoscorpiones, Opiliones, Amblypygi, Thelyphonida, Schizomida, and Araneae and many mites). Missing are the orders Ricinulei, Solifugae, Opilioacarida, and Holothyrida and the suborders Palaeoamblypygi and Dyspnoi. Remarkable is the rich fauna of Uropygi. Palpigradi There is one species recorded: Koeneniodes deharvengi Condé, 1981. 432 7 Regional Arachnogeography Scorpiones The scorpion fauna of the Philippines consists of 14 species of the genera Isometrus Ehr. (Buthidae, 2 spp.), Lychas C.L. Koch (Buthidae, 3 spp.), Chaerilus Simon (Chaerilidae, 3 spp.), Liocheles Sundevall (Hormuridae, 1 sp.), Hormurus Thorell (Hormuridae, 2 spp.), and Heterometrus Ehr. (Scorpionidae, 3 spp.). There are no endemic genera (Kovaøik 2012). Pseudoscorpiones From the Philippines are recorded 41 spp. of pseudoscorpions of 29 genera and 13 families (Beier 1931, 1937, 1966, 1967), Chamberlin (1930), Harvey (2013f). Fam. Chthoniidae – Lagynochthonius Beier (two spp.), Tyrannochthonius Chamberlin (one sp.) Fam. Tridenchthoniidae – Compsaditha Chamberlin (one sp.), Ditha Chamberlin (one sp.), Dithella J.C. Chamberlin et R.V. Chamberlin (one sp.) Fam. Neobisiidae – Bisetocreagris Curcic (one sp.), Microcreagris Balzan (two spp.) Fam. Syarinidae – Alocobisium Beier (one sp.) Fam. Geogarypidae – Geogarypus Chamberlin (one sp.) Fam. Hyidae – Hya Chamberlin (one sp.) Fam. Pseudochiridiidae – Pseudochiridium With (one sp.) Fam. Olpiidae – Olpium L. Koch (one sp.) Fam. Atemnidae – Anatemnus Beier (two spp.), Atemnus Canestrini (one sp.), Oratemnus Beier (four spp.), Metatemnus Beier (one sp.), Paratemnoides Harvey (three spp.), Stenatemnus Beier (one sp.) Fam. Cheiridiidae – Apocheiridium Chamberlin (one sp.), Cryptocheiridium Chamberlin (one sp.) Fam. Cheliferidae – Lissochelifer Chamberlin (one sp.), Lophochernes Simon (two spp.), Metachelifer Redikorzev (one sp.) Fam. Chernetidae – Adelphochernes Beier (two spp.), Megachernes Beier (three spp.), Ceriochernes Beier (one sp.) Fam. Withiidae – Metawithius Chamberlin (one sp.), Withius Kew (one sp.) Endemic genera: Fam. Chernetidae: Adelphochernes Beier, 1937 – Mindanao, Mindoro (two spp.) Opiliones From the Opiliones of the Philippines are recorded 98 spp. of 3 suborders (Martens and Schwendinger 1998, Roewer 1926, 1954, 1955, Shear 1993a, Suzuki 1977a, 1982). Cyphophthalmi Fam. Stylocellidae – Miopsalis Thorell is recorded from Palawan, an island with special position within the Philippine Archipelago, also from Mindanao. Eupnoi Following Suzuki (1977a), we can see that on the archipelago is known only one family of “Palpatores,” with 20 genera and 63 species. 7.26 The Philippines 433 Fam. Sclerosomatidae Gagrellinae – Altobunus Roewer (1), Bakerinulus Roewer (1), Bastia Roewer (1), Bonthainia Roewer (1), Bullobunus Roewer (8), Ceratobunellus Roewer (1), Dentobunus Roewer (2), Eugagrella Roewer (2), Gagrella Stoliczka (21), Gagrellula Roewer (3), Harmanda Roewer (1), Hologagrella Roewer (1), Marthana Thorell (5), Metagagrella Roewer (2), Melanopula Roewer (1), Paragagrella Roewer (1), Pentazaleptus (2), Prodentobunus Roewer (1), Zaleptanus Roewer (2), Zaleptus Thorell (6) Dyspnoi – missing Laniatores From the (very conservative for the many monotypical genera) estimate of Suzuki (1977a and suppl.), the following 34 spp. of Laniatores have been recorded on the Philippine archipelago: Fam. Sandokanidae (Oncopodidae) Biantoncopus Martens et Schwendinger, 1998 – two spp. (Leyte, endemic genus) Gnomulus Thorell, 1876 (Pelitnus Thorell, 1891) – six spp. (Luzon, Mindanao, Leyte) Fam. Epedanidae Paracrobunus Suzuki, 1977 – two spp. (Palawan) Balabanus Suzuki, 1977 – one sp. (Balabac) Dumaguetes Roewer, 1927 – one sp. Fam. Podoctidae Oppodoctis Roewer, 1927– one sp. (Luzon, endemic genus) Hoplodino Roewer, 1915– one sp. (Dak) Ibalonius Karsch, 1880 – five spp. (Luzon, Bohol, Mindanao) Bonea Roewer, 1914 – two spp. (Luzon, Palawan) Fam. Biantidae Dibunus Loman, 1906 – ten spp. (Luzon, Bohol, Negros, Mindanao) Fam. Assamiidae Simalurius Roewer, 1923 – one sp. (Palawan) Amblypygi As a result of his visit to the Philippines in 1890, Simon (1892) described the first Amblypygi from the islands, including the new genus Sarax (Charinidae). As a whole, on the archipelago are represented three spp. of two genera and two families (Giupponi and Miranda 2012, Harvey 2003, 2013). Fam. Charinidae: Sarax Simon, 1892 – S. brachydactylus Simon, 1892; S. curioi Giupponi et Miranda, 2012 (endemic) 434 7 Regional Arachnogeography Fam. Charontidae: Charon Karsch, 1879 – Ch. grayi (Gervais, 1842) Thelyphonida (Uropygi) From the Philippines we used to know five genera of Uropygi: Glyptogluteus Rowland (Panay, one sp.), Minbosius Speijer (one sp.), Mimoscorpius Pocock (one sp.), Abaliella Strand (one sp.), and Thelyphonus Latreille (three spp. on Luzon and Mindanao). As Abaliella and Minbosius were synonymized under Thelyphonus by Haupt (2009a) and a new genus and species were described from Panay (Thelyphonoides panayensis Krehenwinkel Curio, Takud et Haupt, 2009), the situation changed, and now in the Philippines are known four genera with eight spp., including seven species endemic to the islands. “Minbosius” manilanus (C.L. Koch, 1843) is found also in New Guinea and the Moluccas. The fauna is mostly of Southeast Asiatic (Indomalayan) extraction, but with high endemism. The former genera Abaliella and Minbosius have mostly Papuan affinities (Kraepelin 1897, Krehenwinkel et al. 2009, Rowland 1973b, Speijer 1936). Schizomida The record for the Philippines consists only of Orientzomus luzonicus (Hansen, 1905), endemic sp. for Luzon, and “Trithyreus sp.” Undetermined material is known from Mindanao and Negros (Hansen and Sørensen 1905, Reddell and Cokendolpher 1995). The genus Orientzomus Cokendolpher et Tsurusaki is found also on the islands Mariana and Bonin. Araneae In the Philippines are represented ca. 40 families of spiders (Simon 1892, Huber 2011, Barrion and Litsinger 1995, Elias and Nuneza 2016). Endemic genera of spiders: Fam. Linyphiidae Apobrata Miller, 2004 Fam. Selenopidae Amamanganops Crews et Harvey, 2011 (one sp.) Fam. Uloboridae Astavakra Lehtinen, 1967 Fam. Theridiidae Achaearyopa Barrion et Litsinger, 1995 Landoppo Barrion et Litsinger, 1995 Fam. Thomisidae Pycnaxis Simon, 1895 (one sp.), Taypaliito Barrion et Litsinger, 1995 (one sp.) Acariformes The mite fauna of the Philippines is well known, thanks to L. Corpuz-Raros (2005 and many other papers). There are many endemic genera of various families. 7.27 Japan (the Ryukyus Excluded) 7.27 7.27.1 435 Japan (the Ryukyus Excluded) Geography, General Zoogeography, and Paleogeography Japan is a stratovolcanic archipelago extending along the Pacific coast of Asia. Measured from the geographic coordinate system, Japan is 36° north and 138° east. The major islands are (from north to south) Hokkaido, Honshu (the “mainland”), Shikoku, and Kyushu. There are 2456 islands, including the Bonin Islands, Daitô Islands, Minami-Tori-shima, Okinotorishima, Ryukyu Islands, and Volcano Islands. The climate varies from tropical in south to cool temperate in Hokkaido. The Japanese islands are the summits of mountain ridges uplifted near the outer edge of the continental shelf. About 73 percent of Japan’s area is mountainous. The highest point is Mount Fuji, 3776 m. Three mountain chains – the Hida, Kiso, and Akaishi Mountains – form the Japanese Alps (Nihon Arupusu), several of those peaks are higher than 3000 meters. The highest point in the Japanese Alps is Mount Kita at 3193 meters. The Bonin Islands, known in Japan as the Ogasawara Guntô, are an archipelago of over 30 subtropical and tropical islands, some 1,000 kilometers south of Tokyo. The total area of the islands is 73 km2. In Japan, as in most other places, the zoogeography was based mostly on mammals. The conclusions of Dobson (1994) are that: Distribution patterns among the terrestrial mammal species of Sakhalin and the main islands of Japan are shown to fall into 12 clear groups. The most fundamental distributional break (Blakiston’s Line) is that separating Hokkaido and Sakhalin to the north, with their boreal fauna typical of northern Eurasia, from ‘Hondo’ (Honshu, Shikoku, Kyushu) to the south, which demonstrates a high degree of endemism and supports a small number of Indo-Malayan elements. Distribution patterns may be explained by considering the Quaternary geohistory of the area, particularly the formation of land bridges and the changes in climatic conditions during this period. Hondo underwent two main periods of land bridge connection to the Asiatic mainland. The first, prior to the Pleistocene, allowed immigration of forms which have since developed into distinctive elements of the endemic fauna (‘Old Hondo Endemics’). The second, during the Middle Pleistocene, brought in widespread Palaearctic species as well as components from South-East Asia (‘Early Colonists’), some of which have since undergone vicariant speciation (‘New Hondo Endemics’); it also allowed several of the Old Hondo Endemics to extend their range to the mainland and Hokkaido (‘Expanding Hondo Endemics’). Sakhalin and Hokkaido have been more intimately connected to the mainland (most recently until less than 10,000 years ago), such that endemism is very restricted. Species groups here are the ‘Late Colonists’, cold-adapted tundra species which expanded with the glacial advances, but which are now restricted in distribution, and ‘Recent Colonists’, postglacial forest species which recolonized before the severance of land bridges. Moving the other way were ‘Expanding Northern Endemics’, which arose in Hokkaido or Sakhalin during the last glacial and colonized the adjacent mainland before severance of land links. The northern Ryukyu Islands are separated from Kyushu by the so-called Myake Line. 436 7.27.2 7 Regional Arachnogeography Arachnogeography From Japan (Ryukyu excluded) are missing the orders Palpigradi, Solifugae, Ricinulei, Opilioacarida, and Holothyrida. As the country is a chain of islands from the Indomalayan Ryukyus to the harsh climate of Hokkaido, the fauna varies from one area to another (Haupt 2003). The study of Haupt (1993) was “based on four groups of ground-living arachnids: whip scorpions and spiders (Hexathelidae, Ctenizidae, Mesothelae). A close zoogeographical connection was found between the Yaeyama Islands and nearby Taiwan. The central and northern parts of the Ryukyu Islands are faunistically closest to Kyushu. No distinct border between an Oriental and a Palearctic fauna can be detected.” Scorpiones In Japan are recorded only two widespread species of scorpions: Isometrus maculatus (Buthidae) and Liocheles australasiae (Hormuridae). Pseudoscorpiones In Japan (including Ryukyus) are recorded 68 pseudoscorpions, belonging to 13 families (none of them are endemic) (Beier 1952, J.C. Chamberlin 1929, 1938, Čurčić 1979, Ellingsen 1907, Kishida 1966, Morikawa 1955, 1957, 1960; 17 papers of Japanese pseudoscorpions from 1952 to 1972), Sakayori, 1999, 2000, 2002, Sato (1978, 1979, 1982, 30 papers of Japanese pseudoscorpions from 1976 to 1988). Endemic genera for Japan: Fam. Neobisiidae: Pararoncus J.C. Chamberlin, 1938 – seven spp. Fam. Cheliferidae: Kashimachelifer Morikawa, 1957 – one sp. (Honshu) Fam. Olpiidae: Nipponogarypus Morikawa, 1955 – one sp. (Enoshima and the Ryukyus) Opiliones Several authors (Martens and Suzuki 1966, Miyosi 1957, Sato and Suzuki 1939, Shear 2010a, Suzuki, papers from 1939 and 1991, Suzuki and Tsurusaki 1983) have contributed to our understanding that in Japan (Ryukyus and Bonin excluded) are recorded Opiliones of the families Sironidae, Caddidae, Phalangiidae, Nemastomatidae, Sabaconidae, Nipponopsalididae, Travuniidae, Triaenonychidae, Phalangodidae, and Podoctidae. Cyphophthalmi From Japan is known only one (endemic) genus and species. Fam. Sironidae: Suzukielus Juberthie, 1970 Eupnoi Fam. Phalangiidae – Opilio Herbst, 1798 Dyspnoi Fam. Caddidae (Caddo Banks, known also from eastern North America) 7.27 Japan (the Ryukyus Excluded) 437 Fam. Nemastomatidae (Cladolasma Suzuki, known also from Thailand and China) Fam. Sabaconidae – Sabacon Simon (nine spp.) Fam. Nipponopsalididae (one genus Nipponopsalis Martens et Suzuki, known from Japan, the Kuril Islands, and Korea) Laniatores In mainland Japan Laniatores are represented by eight genera and three families (Travuniidae, Phalangodidae, Podoctidae). Endemics for the mainland Japan are the genera: Fam. Travuniidae Yuria Suzuki, 1964 – one sp. (inc. sedis) Nippononychinae (endemic subfamily) Nippononychus Suzuki, 1975 – one sp. Metanippononychus Suzuki, 1975 – four spp. Izunonychus Suzuki, 1975 – one sp. Paranonychinae (USA, Canada, Japan) Kainonychus Suzuki, 1975 – one sp. Fam. Phalangodidae Proscotolemon Roewer, 1916 – one sp. Fam. Podoctidae Iyonus Suzuki, 1964 – one sp. Idzubius Roewer, 1949 – one sp. Thelyphonida (Uropygi) Typopeltis stimpsonii (Wood, 1862) from the islands Tokunoshima to Amakusa and the south of Kyushu, the only Thelyphonida from the mainland Japan (Yoshikura 1973) Araneae The araneofauna of Japan (total), according to Ono (2009), counts more than 1500 spp. In Japan (Ryukyus excluded) are known at least 50 families of spiders (Yaginuma 1961, Chikuni 1989, Kamura and Hayashi 2009, Kamura and Irie 2009, Komatsu 1961, Nishikawa 2009, Ono (ed.) 2009, Saito and Ono 2001, Tanikawa 2009, Tanikawa and Ono 2009). Some endemic genera: Fam. Leptonetidae: Masirana Kishida, 1942, Falcileptoneta Komatsu, 1970 Acariformes Oribatida. According to the checklist of Fujikawa et al. (1993, with suppl.), in Japan (entire) are known oribatid mites of 286 genera and 735 spp. 438 7.28 7.28.1 7 Regional Arachnogeography Ryukyu Islands Geography, General Zoogeography, and Paleogeography Ryukyu Islands (Ryûkyû-shotô), known in Japanese as the Nansei-shotô, lit. “Southwest Islands,” and also known as the Ryukyu Arc are a chain of more than 100 volcanic Japanese islands that stretch 1100 km southwest from Kyushu to Taiwan: the Ôsumi, Tokara, Amami, Okinawa, and Sakishima Islands (further divided into the Miyako and Yaeyama Islands), with Yonaguni the southernmost. The largest of the islands is Okinawa. The surface of the archipelago is 4642 km2, and the highest point is at 1936 m (Mt. Miyanoura-dake). The two largest islands are Okinawa (1,204 square km) and Amami Great Island (712 square km). The islands have a subtropical climate with mild winters and hot summers. Precipitation is very high and is affected by the rainy season and typhoons. Except the outlying Daitô Islands, the island chain has two major geologic boundaries, the Tokara Strait between the Tokara and Amami Islands and the Kerama Gap between the Okinawa and Miyako Islands. Between 1.6 and 1.3 Ma, the East China Sea area, including most of the Okinawa Trough, may have been subaerial. At that time, the Ryukyu Arc region may have been a part of the Eurasian continent. Extensive subsidence may have occurred at the second stage, at about 1.3 Ma, in the Early Pleistocene. The present Ryukyu Arc (Ryukyu Ridge) has been formed since then. The Ryukyu Arc may have been nearly connected to the Chinese continent, through Taiwan as a land bridge, sometime during the two major development periods (such as sometime during 1.6–1.0 Ma and 0.2–0.025 Ma). The Paleo-land may have been submerged step by step since 0.03 Ma by both crustal movement and sea level rising after the last Ice Age. Submarine stalactite caves at 10–35 m deep off the Ryukyu Islands were discovered. The caves have subsided since the Würm Ice Age. Stone tools were also recovered inside one of them (Kimura 2000). Watase’s Line, which crosses the Tokara Islands, marks a major biogeographic boundary. The north of the line belongs to the Palearctic region, while the southern portion is the northern limit of the Oriental region. Yakushima in Ôsumi is the southern limit of the Palearctic region. It is featured with millennium-old cedar trees. The island is part of Kirishima-Yaku National Park and was designated as a World Heritage Site by UNESCO in 1993. The south of Watase’s Line is recognized by ecologists as a distinct subtropical moist broadleaf forest ecoregion. The flora and fauna of the islands have much in common with Taiwan, the Philippines, and Southeast Asia and are part of the Indomalayan ecozone. Mammals endemic to the islands include Iriomote cat, the Ryukyu flying fox, the Ryukyu long-tailed giant rat, the Ryukyu mouse, and the Ryukyu shrew. Approximately one half of the amphibian species of the islands are endemic. 7.28 Ryukyu Islands 439 The northern Ryukyu Islands are separated from Kyushu by the so-called Myake Line. Other sources: Dobson (1994), Kimura (2002) Map 7.16 Map of the Ryukyus 7.28.2 Arachnogeography On the Ryukyus are absent the orders Ricinulei, Opilioacarida, and Holothyrida. The study of Haupt (1993) was “based on four groups of ground-living arachnids: whip scorpions and spiders (Hexathelidae, Ctenizidae, Mesothelae). A close zoogeographical connection was found between the Yaeyama Islands and nearby Taiwan. The central and northern parts of the Ryukyu Islands are faunistically closest to Kyushu. No distinct border between an Oriental and a Palearctic fauna can be detected. Instead, only climatic and other ecological factors appear to be responsible for distributional boundaries of various organisms observed in different regions of southern Japan. Similarities to the fauna of continental China can be explained by land connections between the Ryukyus and the continent during the Pleistocene.” Among the Opiliones the “Oriental” elements dominate (Suzuki 1973). 440 7 Regional Arachnogeography Scorpiones Fam. Hormuridae – Liocheles australasiae (Fabricius) (Takashima 1941) Pseudoscorpiones Fam. Olpiidae – Nipponogarypus enoshimaensis okinoerabensis Morikawa, 1960 (endemic for the Ryukyus) (Morikawa 1960) Opiliones According to the paper of Suzuki (1971), “The Opiliones fauna of the Ryukyus is composed of eleven genera and thirteen species (eleven species of Oriental origin and two of Pale- or Holarctic one).” A later paper (Suzuki 1973) listed already 16 genera and 26 species (Phalangiidae, Podoctidae, Epedanidae, Travuniidae, Sclerosomatidae (sub-“Leiobunidae”), and Ischyropsalididae (also Suzuki (1964)). Cyphophthalmi – not recorded Eupnoi Fam. Phalangiidae Fam. Sclerosomatidae – Gagrella Stoliczka, Metagagrella Roewer, Pseudogagrella Redikorzev, Gagrellula Roewer, Leiobunum C.L. Koch, Nelima Roewer) Dyspnoi Fam. Nipponopsalididae (Nipponopsalis Martens et Suzuki) Fam. Travuniidae (Peltonychia) Fam Phalangodidae (Proscotolemon Roewer, Parabeloniscus Suzuki, Tokunosia Suzuki) Fam. Podoctidae (Dongmoa Roewer) Fam. Epedanidae (Zepedanulus Roewer, Epedanellus Roewer, Kilungius Roewer, Pseudobiantes Hirst) Thelyphonida (Uropygi) In Japan live two species of Thelyphonida: Typopeltis crucifer Pocock, 1894, in the southernmost Ryukyu Islands (from Iriomote Shima to Ibeya Shima, also in Taiwan) and T. stimpsonii (Wood, 1862) from the islands Tokunoshima to Amakusa and the south of Kyushu, the only Uropygi from the mainland Japan (Yoshikura 1973) (Map 7.15). According to Karasawa et al. (2015), “…the border of distribution of the two species lies between the Central and Southern Ryukyus, i.e., the Kerama Gap.” Schizomida The order is known only on the smaller islands: Orientzomus sawadai (Kishida, 1930) – endemic sp. on Ogasawara-shotô (= Bonin Islands), Tokyo Pref., and three species on the Ryukyu Islands (Okinawa Pref.) (Cokendolpher 1988, Kishida 1930, Shimojana 1981). There are no endemic genera for Japan. Araneae The spider fauna of Ryukyus has been studied by several arachnologists (Yaginuma, Komatsu, Shimojana, Haupt, and others). A preliminary report of cave spiders (Shimojana 1977) contains 42 species of 36 genera and 23 families. To them has been added the blind spider Coelotes okinawensis Shimojana et Nishihira, 2000. 7.29 Korea (North and South) 441 The conclusion of Shimojana (1977) is that “The cave spider fauna of the Ryukyu Archipelago is much different from the Japanese Islands.” Endemic genera from the Ryukyus: Fam. Liphistiidae: Ryuthela Haupt, 1983 – Ryukyu Islands, Okinawa (seven spp.) Fam. Corinnidae: Humua Ono, 1987 – one sp. 7.29 7.29.1 Korea (North and South) Geography, General Zoogeography, and Paleogeography Korea is a 1100-kilometer-long peninsula located in the easternmost part of the Asian continent. To the northwest, the Amnok River (Yalu) separates Korea from China, and to the northeast, the Duman (Tumen) River separates Korea from China and Russia. Notable island is Jeju. The highest mountain in Korea is Mount Paektu or Paektusan (2744 m), through which runs the border with China. Mountains cover 70 percent of Korea. Most of the 3579 islands off the peninsula are found along the south and the west coasts. Unlike most ancient mountains on the mainland, many important islands in Korea were formed by volcanic activity in the Cenozoic orogeny. Jeju Island, situated off the southern coast, is a large volcanic island whose main mountain Mount Halla or Hallasan (1950 m) is the highest in South Korea. The climate of Korea differs dramatically from north to south. The southern regions experience a relatively warm and wet climate similar to that of Japan, affected by warm ocean waters including the East Korea Warm Current. The northern regions experience a colder and to some extent more inland climate, in common with Manchuria. According to the World Wide Fund for Nature, Korea consists of several ecoregions. The South Korea evergreen forests occupy the southernmost portion of the peninsula, as well as the island of Jeju. The Central Korea deciduous forests occupy the more temperate central portion of the peninsula. Manchurian mixed forests occupy the northern lowlands and low hills of the peninsula and extend north into Manchuria as far as the Amur River on the Russia-China border. The Changbai Mountains mixed forests include the higher elevation mountain region along the North Korea-China border, where forests are dominated by conifers, with alpine meadows and rock slopes on the highest peaks (World Wide Fund for Nature). The sea level of the South Sea area was about 150–160 m. below present sea level about 15,000 years ago. This drop allowed a land bridge to form between the Korea peninsula and Japan, which remained above sea level for about 6000 years. The sea level rose to approximately 60 m. (below present sea level) about 9000 years ago, remained at this depth until about 4000–5000 years ago, and then rose to 10–20 m. below the present level about 3000– 4000 years ago. (Yeon Gyu Lee et al. 2008) 442 7.29.2 7 Regional Arachnogeography Arachnogeography The arachnofauna fauna of North Korea is not well studied. On the peninsula are known the orders Scorpiones, Pseudoscorpiones, Opiliones, Araneae, Uropygi, Ixodida, Mesostigmata, Sarcoptiformes, and Trombidiformes. Lacking are Palpigradi, Ricinulei, Amblypygi, Opilioacarida, Holothyrida, and Cyphophthalmi. To be noted is the presence of the Opilion subfamily Kaolinonychinae (Triaenonychidae, shared with Japan). Our collection of Arachnida from North Korea (1982, 1987) is still in Sofia, unidentified. Scorpiones Only two scorpion species have been recorded so far (Zhu et al. 2004). Fam. Buthidae – Mesobuthus martensii Fam. Hormuridae – Liocheles australasiae Pseudoscorpiones From South Korea have been recorded 19 spp. of 9 genera and the families Chthoniidae, Pseudotyrannochthoniidae, Neobisiidae, Syarinidae, Cheiridiidae, and Chernetidae (Morikawa 1970; Lee 1981, 1982, Harvey 2014). No information about North Korea (material has been collected, but not yet identified). Opiliones The total number of the Opiliones from Korea is 17 spp. of the families Phalangiidae, Sclerosomatidae, Sabaconidae, Nipponopsalididae, and Triaenonychidae (Kharitonow 1957, Roewer 1927, Staręga 1964, 1965, Suzuki 1941b, 1966, 1975, Kim et al. 2006, Ban Kwon and Kim 2010). According to Kury (online), in Korea there are seven endemic species. Cyphophthalmi are not recorded from Korea. Eupnoi Fam. Phalangiidae – Euphalangium Roewer, Mitopus Thorell, Oligolophus C.L. Koch, Opilio Herbst, Himalphalangium Martens Fam. Sclerosomatidae – Metagagrella Roewer, Nelima Roewer Dyspnoi Fam. Sabaconidae – Sabacon Simon Fam. Nipponopsalididae – Nipponopsalis Martens et Suzuki 7.29 Korea (North and South) 443 Laniatores From the described area are known Laniatores from the family Triaenonychidae (according to some authors, the subfamily Kaolinonychinae should belong to Travuniidae). Endemic genera are: Fam. Triaenonychidae Kaolinonychinae – Korea, Japan (two spp.) Kaolinonychus Suzuki, 1975 – Korea (one sp.) Thelyphonida (Uropygi) Fam. Thelyphonidae – according to Harvey (2013d), whip scorpions are not known from Korea, but they certainly live there, as in the neighboring China and Japan (Typopeltis?). Butler (1872) describes from “Korea” the new species Thelyphonus lucanoides (= now considered a dweller of Sarawak and Indonesia). Amblypygi – not known Araneae The list of spider fauna of Korea (Paik 1967) contains 242 spp. of 120 genera and 33 families. Zoogeographically Korean spider fauna “comprises 136 northern species, 61 southern species, 6 cosmopolitan species, and 20 endemic species.” Most species (125) are known also from Japan. Paik Kap-Yong (1914–1996) contributed largely to the study of Korean spiders and also to the publishing of the first pictorial encyclopedia (1978). Namkung Joon published in 2001 even bigger pictorial encyclopedia of Korean spiders with taxonomic accounts on 546 spp. of 220 genera and 43 families. Namkung et al. (2009) increased this number to 681 spp. of 258 genera and 46 families, including 131 endemic species (19.2%). The spider fauna is close to the Japanese, influenced by northern elements, including 35 Holarctic and 67 Palearctic species. Most species are in Linyphiidae (82), Theridiidae (77), Araneidae (70), and Salticidae (65). Endemic genera: Fam. Liphistiidae: Ryuthela Haupt, 1983 Fam. Leptonetidae: Longileptoneta Seo, 2015 (five spp.) Fam. Gnaphosidae: Shiragaia Paik, 1992 444 7 Regional Arachnogeography Map 7.17 Map of Africa (political) 7.30 7.30.1 Africa, North of 20oN Geography, General Zoogeography, and Paleogeography In the narrow sense, here are included the territories of Mauretania, Western Sahara, Morocco, Algeria, Tunisia, Libya, and Egypt, roughly north of 20oN. The Mediterranean coast with its Mediterranean climate, especially in the Maghreb, is followed south by Atlas Mountains, rising in Morocco up to 4167 m (Djebel Toubkal). More to the south start the Saharan sands. The classical schemes in zoogeography include North Africa in the Palearctic region (under different names). We have to notice that 7.30 Africa, North of 20oN 445 Maghreb, by its origin, is a territory of Tirrenis and relates much more to Europe, than to the rest of Africa; its autochthonic elements are Tyrrenic and are in common with the other lands of the Western Mediterranean, and only by the end of the Miocene (in the Pontian) were received various African elements (firstly from the Saharan-Ethiopian area) and Asiatic elements, originating from Palestine and Egypt. (La Greca 1961) Cox (2001) has left the African part of his “Eurasian” (generally called Palearctic) region only a narrow band in Maghreb North of Atlas Mountains. After him, the Sahara is an area where the former tropical flora of North Africa has disappeared and logically is considered a part of the African (usually called “Afrotropical”) region. One zoogeographical subdivision of North Africa was done by Joleau (1928). The Atlas Mountains is a mountain range across the northwestern stretch of Africa extending about 2,500 km through Algeria, Morocco, and Tunisia. The Atlas ranges separate the Mediterranean and Atlantic coastlines from the Sahara Desert. Map 7.18 The location of the Atlas Mountains across North Africa In the Paleogene and Neogene periods (~66 million to ~1.8 million years ago), the mountain chains that today comprise the Atlas were uplifted as the landmasses of Europe and Africa collided at the southern end of the Iberian Peninsula. Such convergent tectonic boundaries occur when two plates slide toward each other forming a subduction zone (if one plate moves underneath the other) and/or a continental collision (when the two plates contain continental crust). In the case of the AfricaEurope collision, it is clear that tectonic convergence is partially responsible for the formation of the High Atlas, as well as for the closure of the Strait of Gibraltar and the formation of the Alps and the Pyrenees. The Sahara is the world’s hottest and largest desert (over 9,400,000 km2). The Sahara stretches from the Red Sea, including parts of the Mediterranean coasts, to the Atlantic Ocean. To the south, it is delimited by the Sahel, a belt of semiarid tropical savanna that composes the northern region of central and western subSaharan Africa. 446 7 Regional Arachnogeography Sahara’s boundaries are the Atlantic Ocean on the west, the Atlas Mountains and the Mediterranean on the north, the Red Sea on the east, and the Sudan (region) and the valley of the Niger River on the south. The Sahara is divided into Western Sahara, the central Ahaggar Mountains, the Tibesti Mountains, the Aïr Mountains (a region of desert mountains and high plateaus), Ténéré Desert, and the Libyan Desert (the most arid region). The highest peak in the Sahara is Emi Koussi (3,415 m) in the Tibesti Mountains in northern Chad. The southern border of the Sahara is marked by a band of semiarid savanna called the Sahel; south of the Sahel lies Southern Sudan and the Congo River Basin. Most of the Sahara consists of rocky hamada; ergs (large areas covered with sand dunes) form only a minor part. Sahara used to be a much wetter place than it is today. Over 30,000 petroglyphs of river animals such as crocodiles survive, with half found in the Tassili n’Ajjer in southeast Algeria. The modern Sahara, though, is not lush in vegetation, except in the Nile Valley, at a few oases, and in the northern highlands, where Mediterranean plants such as the olive tree are found to grow. The region has been this way since about 1600 BCE, after shifts in the Earth’s axis increased temperatures and decreased precipitation. Then, due to a climate change, the savanna changed into the sandy desert. The Sahara covers large parts of Algeria, Chad, Egypt, Libya, Mali, Mauritania, Morocco, Niger, Western Sahara, Sudan, and Tunisia. Several deeply dissected mountains and mountain ranges and many volcanoes rise from the desert, including the Aïr Mountains, Ahaggar Mountains, Saharan Atlas, Tibesti Mountains, Adrar des Iforas, and the Red Sea hills. Most of the rivers and streams in the Sahara are seasonal or intermittent, the chief exception being the Nile River. Underground aquifers sometimes reach the surface, forming oases. The central part of the Sahara is hyperarid, with little vegetation. To the north, the Sahara reaches to the Mediterranean Sea in Egypt and portions of Libya, but in Cyrenaica and the Maghreb, the Sahara borders Mediterranean forest, woodland, and scrub ecoregions of northern Africa, which have a Mediterranean climate characterized by a winter rainy season. According to the botanical criteria of Frank White, the northern limit of the Sahara corresponds to the northern limit of date palm cultivation. The northern limit also corresponds to the 100 mm isohyet of annual precipitation. To the south, the Sahara is bounded by the Sahel, a belt of dry tropical savanna with a summer rainy season that extends across Africa from east to west. According to climatic criteria, the southern limit of the Sahara corresponds to the 150 mm isohyet of annual precipitation (this is a long-term average, since precipitation varies annually). The Sahara comprises several distinct ecoregions, and with their variations in temperature, rainfall, elevation, and soil, they harbor distinct communities of plants and animals. The Atlantic coastal desert is a narrow strip along the Atlantic coast, where fog generated offshore by the cool Canary Current provides sufficient moisture to sus- 7.30 Africa, North of 20oN 447 tain a variety of lichens, succulents, and shrubs. It covers 39,900 km2 in Western Sahara and Mauritania. The North Saharan steppe and woodlands is along the northern desert, next to the Mediterranean forests, woodlands, and scrub ecoregions of the northern Maghreb and Cyrenaica. It covers 1,675,300 km2 of Algeria, Egypt, Libya, Mauritania, Morocco, Tunisia, and Western Sahara. The Sahara Desert ecoregion covers the hyperarid central portion of the Sahara where rainfall is minimal and sporadic. Vegetation is rare, and this ecoregion consists mostly of sand dunes (erg, chech, raoui), stone plateaus (hamadas), gravel plains (reg), dry valleys (wadis), and salt flats. It covers 4,639,900 km2 of Algeria, Chad, Egypt, Libya, Mali, Mauritania, Niger, and Sudan. The South Saharan steppe and woodlands ecoregion is a narrow band running east and west between the hyperarid Sahara and the Sahel savannas to the south. Movements of the equatorial Intertropical Convergence Zone (ITCZ) bring summer rains during July and August which average 100 to 200 mm but vary greatly from year to year. This ecoregion covers 1,101,700 km2 of Algeria, Chad, Mali, Mauritania, and Sudan. In the West Saharan montane xeric woodlands, several volcanic highlands provide a cooler, moister environment that supports Sahara-Mediterranean woodlands and shrublands. The ecoregion covers 258,100 km2, mostly in the Tassili n’Ajjer of Algeria, with smaller enclaves in the Aïr of Niger, the Dhar Adrar of Mauritania, and the Adrar des Iforas of Mali and Algeria. The Tibesti-Jebel Uweinat montane xeric woodlands ecoregion consists of the Tibesti and Jebel Uweinat highlands. Higher and more regular rainfall and cooler temperatures support woodlands and shrublands of palms, acacias, myrtle, oleander, tamarix, and several rare and endemic plants. The ecoregion covers 82,200 km2 in the Tibesti of Chad and Libya and Jebel Uweinat on the border of Egypt, Libya, and Sudan (http://en.wikipedia.org/wiki/Sahara-cite_note-34). The Saharan halophytics is an area of seasonally flooded saline depressions which is home to halophytic (salt-adapted) plant communities. The Saharan halophytics cover 54,000 km2, including the Qattara and Siwa depressions in northern Egypt, the Tunisian salt lakes of central Tunisia, Chott Melghir in Algeria, and smaller areas of Algeria, Mauritania, and Western Sahara. The Tanezrouft is one of the harshest regions on Earth and the driest in the Sahara, with no vegetation and very little life. It is along the borders of Algeria, Niger, and Mali, west of the Ahaggar Mountains. The flora of the Sahara is highly diversified based on the biogeographical characteristics of this vast desert. Floristically, the Sahara has three zones based on the amount of rainfall received – the Northern (Mediterranean), Central, and Southern Zones. There are two transitional zones – the Mediterranean-Sahara transition and the Sahel transition zone. 448 7.30.2 7 Regional Arachnogeography Arachnogeography According to Udvardy (1975), North Africa is divided biogeographically into three parts (from north to south): No 17 (Mediterranean sclerophyll), 28 (Atlas steppe), and 18 (Sahara). The arachnofauna of the first two “provinces” (Udvardy) is very close to the South European fauna, even as the same species (Ptychosoma vitellinum Sør.). Here (especially in the caves) are found such hygrophilous animals as Palpigradi or some pseudoscorpions of Neobisiidae. As expected, the animals of the desert and hot climate as Solifugae and Scorpiones are very numerous and quite different from the European-Mediterranean fauna. The spider fauna is well studied up to the very summit of Atlas (4167 m) (Deltshev 2015, collected by P. Beron). Palpigradi Barranco and Mayoral (2007) described from the cave of Kef Aziza the new species Eukoenenia maroccana – the third Moroccan Palpigradi, after E. mirabilis (Grassi et Calandruccio, 1885) and E. hanseni (Silvestri, 1913) (Remy 1948, 1952). Solifugae In North Africa are distributed Solifugae from 19 genera and 5 families: Galeodidae, Karschiidae, Daesiidae, Solpugidae, and Rhagodidae (Borelli 1924, Lawrence 1966). Endemic genus of Solifugae for North Africa is Barrus Simon, 1880 – Egypt (one sp.). In the North African countries, the number of Solifugae species is as follows (Table 7.3): Scorpiones According to the lists of Vachon (1952) and Dupré (2013a), updated (Lourenço and Duhem 2007), from the North African countries are recorded scorpions of 24 genera and 4 families. From Tassili n’Ajjer, Vachon (1958) identified ten species of scorpions of seven genera and the families Buthidae and Scorpionidae. According to this author, in the “saharo-mountainious” region (Tassili n’Ajjer, Air, Adrar Iforas, and Tibesti) is seen one altitudinal separation of species of Palearctic origin, having “climbed” above 1000 m with the establishment of drought in North Africa, and species, widespread in Sahara. This list has been completed by Lourenço (2009) with the description of Compsobuthus tassili and by Qi Jian-Xin and Lourenço (2007) who described from Mauritania the new genus and species Mauritanobuthus geniezi. From Tunisia Vachon (1951) identified seven spp. of scorpions of the genera Euscorpius, Scorpio, Buthus, Androctonus, Buthacus, and Buthiscus. Among the scorpions of Tunisia, this author distinguishes two species of European origin (“Buthus” occitanus and Euscorpius sicanus) and five North African elements. The scorpions of Egypt (then 12 spp., now they are 28) have been listed as early as 1910 by E. Simon. The list of Moroccan scorpions (50 spp.) is more recent – of Nickel et al. (2009). Teruel (2007) described from Atlas the new genus and species Cicileiurus monticola (relict at 2000 m) (Table 7.4). 7.30 Africa, North of 20oN 449 Table 7.3 Distribution of Solifugae in North Africa Country No. of species Fam. Daesiidae Biton Karsch Blossia Simon Gluviopsilla Roewer Gnosippus Karsch Tarabulida Roewer Fam. Galeodidae Galeodes Olivier Galeodopsis Birula Othoes Hirst Paragaleodes Kraepelin Fam. Karschiidae Barrus Simon Eusimonia Kraepelin Fam. Rhagodidae Rhagodes Pocock Rhagoditta Roewer Rhagodira Roewer Rhagodeya Roewer Fam. Solpugidae Oparba Roewer Oparbella Roewer Solpuga Lichtenstein Zeria Simon Mauritania 4 + – 1 – Morocco 25 + 1 5 – Algeria 30 + 6 1 1 Tunisia 23 + 5 1 – Libya 19 + 5 – – Egypt 27 + 4 2 – – – – – – 1 – – – – 1 – + 1 – + 9 – + 7 – + 9 – + 7 1 + 1 – 1 1 – 1 1 2 – 1 – – – 2 – + + – + + – – – 4 – 3 – – – 1 1 1 – + + + + + – – 1 1 1 3 – – 2 1 – 1 – – 2 1 – 1 – – – – 1 – – – – + 1 2 + – 4 + – 4 + – 1 + – 2 – – 1 – 1 – – 1 1 – – – 450 7 Regional Arachnogeography Table 7.4 Scorpions in northern Africa Country Number of Species Fam. Buthidae Ananteroides Borelli Androctonus Ehrenberg Buthacus Birula Butheoloides Hirst Buthiscus Birula Buthus Leach Cicileiurus Teruel Cicileus Vachon Compsobuthus Vachon Egyptobuthus Lourenço Hottentotta Birula Isometrus Ehrenberg Leiurus Ehrenberg Lissothus Vachon Mauritanobuthus Qi et Lourenço Microbuthus Kraepelin Orthochirus Karsch Parabuthus Pocock Pseudolissothus Lourenço Saharobuthus Lourenço et Duhem Fam. Euscorpiidae Euscorpius Thorell Fam. Scorpionidae Nebo Simon Scorpio L. Fam. Hemiscorpiidae Hemiscorpius Peters Morocco 50 + – Algeria 30 + – Tunisia 17 + – Libya 16 + – Egypt 28 + – 16 + 1 Mauritania 28 + – 7 7 4 3 5 5 5 4 5 – 15 1 – 2 5 1 1 4 – 1 2 2 – 1 4 – – – 3 – 1 3 – – 1 2 – – 6 – – 4 2 1 – 1 – – 1 2 – – 6 – – 4 – – – – 1 – 1 3 – 3 1 – – 2 – – – – – – – – – – – 1 – – – – – 1 – 1 – – – 1 1 1 – – 1 – – – 1 – 1 3 2 1 1 2 1 2 – – – 1 – 1 – – 2 – – – 2 – 1 – – – – – – + + + – – – + 1 + 2 + 3 + – + – + – + – + – 6 – – 2 – – 2 – – 1 – 1 1 + – 1 – 1 1 + – – – – 1 – 1 7.30 Africa, North of 20oN 451 Endemic genera for the area are: Mauritanobuthus Qi et Lourenço, 2007 – Mauritania (M. geniezi) Saharobuthus Lourenço et Duhem, 2009 – Western Sahara Lissothus Vachon, 1948 – Mauritania, Libya, Algeria (three spp.) Cicileiurus Teruel, 2007 – Morocco Pseudoscorpiones From North Africa (Egypt, Libya, Tunisia, Algeria, Morocco, and Mauritania) and Tibesti in northern Chad are known Pseudoscorpiones of nine families (Chthoniidae, Garypidae, Garypinidae, Geogarypidae, Olpiidae, Atemnidae, Chernetidae, Cheliferidae, Withiidae) (Beier 1955, 1961, El-Hennawy 1988, Heurtault 1970a, 1970b, 1970c, 1971, 1990, Leclerc 1989, Mahnert 1983, Vachon 1954). General papers of the pseudoscorpions have been published by Callaini on Algeria (1983) and Morocco (1988). According to Harvey (online), the number of species known from these countries is Egypt, 14; Libya, 15; Tunisia, 26; Algeria, 43; Morocco, 38; Mauritania, 2; and Tibesti in northern Chad (after Heurtault 1970 a, b, c, 1971), 8. Endemic species are: Chthonius kabylicus Callaini, 1983 – Algeria Neobisium (Blothrus) peyerimhoffi Heurtault, 1990 – Algeria N. dumitrescoae Heurtault, 1990 – Algeria Roncus numidicus Callaini, 1983 – Algeria Roncus (Parablothrus) comasi Mahnert, 1985 – Tunisia Roncus (Parablothrus) gardinii Heurtault, 1990 – Algeria Rhacochelifer massylicus Callaini, 1983 – Algeria Geogarypus mirei Heurtault, 1970 – Chad (Borkou) Goniochernes vachoni Heurtault, 1970 – Tibesti Endemic genus for this (mostly desert) area is only Plesiowithius Vachon, 1954 – Mauritania (one sp.) (Withiidae). Opiliones In North Africa are registered Opiliones of the families Phalangiidae, Trogulidae, Nemastomatidae, and Phalangodidae. The Cyphophthalmi, so numerous in South Europe, are not known so far from North Africa. In Egypt (Cokendolpher 1990) are known six spp. of Opiliones: Trogulus gypseus, Metaphalangium cirtanum, M. orientalis, Phalangium aegyptiacum, Ph. savignyi, and Ph. copticum. Eupnoi Fam. Phalangiidae – Metaphalangium Roewer, Phalangium L. Dyspnoi Fam. Nemastomatidae – Nemastomella M.L. (N. maarebensis Simon, 1913, ¿ “Nemastoma” lilliputanum Lucas, Algeria) Fam. Trogulidae – Trogulus Latreille (one sp., Egypt) 452 7 Regional Arachnogeography Laniatores Fam. Phalangodidae: Ptychosoma vitellinum Soerensen, 1873 – Algeria Amblypygi Musicodamon atlanteus Fage, 1939 (Phrynichidae), is an endemic genus and species from Morocco and Algeria. El-Hennawy and Hisham (2002) recorded from Egypt Charinus ioanniticus (Kritscher) (Charinidae), known also from Turkey, Israel, and the Greek islands Kos and Rhodes. Charinidae and Phrynichidae are the only families of Amblypygi, known from the Palearctic region. Araneae Bosmans (many papers on Atlas spiders), Denis (1961, 1967), El-Hennawy (1990 2006), Deltshev (2015), Caporiacco (1936) Some endemic genera of spiders in Africa, north of 20oN: Fam. Linyphiidae Cherserigone Denis, 1954 – Algeria (one sp.) Fam. Lycosidae Phonophilus Ehrenberg, 1831 – Libya Fam. Dictynidae Chaerea Simon, 1884 – Algeria (one sp.) Fam. Agelenidae Pseudotegenaria Caporiacco, 1934 – Libya (one sp.) Fam. Araneidae Poecilarcys Simon, 1895 – Tunisia (one sp.) Fam. Corinnidae Scorteccia Caporiacco, 1936 – Libya (one sp.) Opilioacarida One species (Opilioacarus segmentatus With, 1903) is known from Algeria (also in South Europe) Parasitiformes In Egypt have been registered Ixodida by Hoogstraal and Kaiser (1958). 7.31 Africa Between 20oN and Zambezi-Kunene (Tropical or Intertropical Africa) 7.31 7.31.1 453 Africa Between 20oN and Zambezi-Kunene (Tropical or Intertropical Africa) Geography, General Zoogeography, and Paleogeography Across Sahara around the 20°N, is situated the usually accepted southern border of the Palearctic (Holarctic), a frontier between one kingdom and another (Paleotropica). South of the Sahara follow the belts of semidesert (Sahel) and the different types of savanna. In the described area, there are also high mountains with afroalpine vegetation and mountain rain forest. Tropic rain forests are tropical moist forests of semi-deciduous varieties distributed across nine West African countries. In the first half of the 1980s, an annual forest loss of 7200 square kilometers was noted down along the Gulf of Guinea, a figure equivalent to 4–5% of the total remaining rain forest area. By 1985, 72 percent of West Africa’s rain forests had been transformed into fallow lands, and an additional 9 percent had been opened up by timber exploitation. It is generally believed that firewood provides 75 percent of the energy used in sub-Sahara Africa. With the high demand, the consumption of wood for fuel exceeds the renewal of forest cover. The rain forests which remain in West Africa now merely are how they were hardly 30 years ago. In Guinea, Liberia, and the Ivory Coast, there is almost no primary forest cover left unscathed; in Ghana the situation is much worse, and nearly all the rain forest are cut down. Guinea-Bissau loses 200 to 350 km2 of forest yearly, Senegal 500 km2 of wooded savanna, and Nigeria 6,000,050,000 of both. Liberia exploits 800 km2 of forests each year. Tropical Africa is about 18% of the world total covering 20 million km2 of land in West and Central Africa. Recent estimates show that the annual pace of deforestation in the region can vary from 150 km2 in Gabon to 2900 km2 in Ivory Coast. The remaining tropical forests still cover major areas in Central Africa but are abridged by patches in West Africa. The tropical environment is rich in terms of biodiversity. Tropical African forest is 18 percent of the world total and covers over 3.6 million square kilometers of land in West, East, and Central Africa. This total area can be subdivided to 2.69 million square kilometers (74%) in Central Africa, 680,000 square kilometers (19%) in West Africa, and 250,000 square kilometers (7%) in East Africa. In West Africa, a chain of rain forests up to 350 km long extends from the eastern border of Sierra Leone all the way to Ghana. In Ghana the forest zone gradually dispels near the Volta River, following a 300 km stretch of Dahomey savanna gap. The rain forest of West Africa continues from east of Benin through southern Nigeria and officially ends at the border of Cameroon along the Sanaga River. The variety of the African rain forest flora is also less than the other rain forests. This lack of flora has been credited to several reasons such as the gradual infertility since the Miocene, severe dry periods during Quaternary, or the refuge theory of the cool and dry climate of tropical Africa during the last severe Ice Age of about 18000 years ago. 454 7 Regional Arachnogeography These analyses of the prominent French entomologist are based mostly on the distribution and the affinities of pselaphids and some other groups of Coleoptera. When studying the distribution of Arachnida, it seems useful to compare the conclusions of Jeannel with the data extracted of the recent profound research on many groups of arachnids. It is clear, for example, that such group as Ricinulei in Africa (only in the western part) suggest former very old “Afro-Brazilian” distribution. There are similar examples also by other groups (the family Neogoveidae – Opiliones, Cyphophthalmi). Jeannel (loc. cit.) indicates that the equatorial forest in Central Africa was established in the Pliocene and is only a residue of a much more extensive forest, coming from the Malaysian area and broken by the vast steppic and desert regions of India, Arabia, and East Africa, deprived of forest by climatic changes and human activities. So, when defining a group, family, or genus of arachnids (e.g., in Madagascar) as having an African origin, we should think of a former origin of these (now East African) elements from Southeast Asia. Some of the elements in the present-day intertropical fauna of Africa actually originate of the southern “sudamadian” areas. Usually tropical Africa is divided into two subregions: East Africa and West Africa. Other sources: Hurni (1989), Scott (1958) 7.31.2 Arachnogeography Old and rich fauna, with all orders of Arachnida (Holothyrida only on the islands) Palpigradi – one endemic genus; Solifugae – six families, some endemic genera, no endemic families Ricinulei – one endemic genus in West Africa Thelyphonida (Uropygi) – one endemic (relict?) genus in West Africa Amblypygi – three families; one endemic and relict suborder in West Africa; Schizomida – three endemic genera Scorpiones – three families Pseudoscorpiones – 16 families Opiliones – eight families, many endemic genera Araneae – many endemic genera It is to notice the relict elements of orders, represented only in West Africa (Ricinulei, Thelyphonida, or Uropygi, suborder Palaeoamblypygi, fam. Neogoveidae). Missing are the suborders Dyspnoi and Mesothelae. 7.31 Africa Between 20oN and Zambezi-Kunene (Tropical or Intertropical Africa) 455 Palpigradi Very few Palpigradi have been described from tropical Africa: Eukoenenia pauli Condé, 1979 (Gabon), E. angolensis (Remy, 1956), E. machadoi (Remy, 1950) (Angola), E. hesperia (Remy, 1953) (Ivory Coast), E. kenyana Condé, 1979 (Kenya), Koeneniodes notabilis Silvestri, 1913, Leptokoenenia scurra Monniot, 1966 (Congo), and Allokoenenia afra Silvestri, 1913 (Guinea) (Condé 1979a, 1979b, Monniot 1966, Remy 1950, 1953, 1956, Silvestri 1913). The genus Allokoenenia is endemic for tropical Africa. This number does not reflect the real picture of the distribution of Palpigradi between 20oN and KuneneZambezi. These tiny and fragile creatures are rarely collected, mostly by the few specialists on them in person. Table 7.5 Solifugae in the countries of Afrotropical Region Countries Number of species Fam. Ceromidae Ceroma Karsch Fam. Daesiidae Biton Karsch Bitonota Roewer Blossia Simon Blossiana Roewer Eberlanzia Roewer Gluviopsilla Roewer Gluviopsis Roewer Gnosippus Karsch Hemiblossia Roewer Triditarsula Roewer Fam. Galeodidae Galeodes Olivier Gluviema Caporiacco Paragaleodes Kraepelin Fam. Rhagodidae Rhagodes Pocock Rhagodessa Roewer Rhagodia Roewer Rhagodinus Roewer Rhagodippa Roewer Rhagoditta Roewer Rhagodoca Roewer Rhagoduna Roewer Ethiopia 46 + 1 + 7 1 – 1 – – 1 – 1 1 + 7 – 2 + – – 1 1 – 1 5 Eritrea 5 – – + 2 – – – – – – 1 – – – – – – + – 1 – – – 1 – Somalia 48 – – + 8 – 2 – 1 1 5 – – – + 5 1 – + 6 – – – 1 5 1 Djibouti 8 – – + 2 – – – – – 1 – – – + 1 – – + – – – 1 1 – 1 – (continued) 456 7 Regional Arachnogeography Table 7.5 (continued) Countries Fam. Solpugidae Ferrandia Roewer Solpuga Lichtenstein Solpugassa Roewer Solpugeira Roewer Solpugella Roewer Solpugyla Roewer Zeria Simon Zeriassa Pocock Countries Number of sp. Fam. Daesiidae Biton Karsch Blossia Simon Gluviopsis Roewer Gnosippus Karsch Fam. Galeodidae Galeodes Olivier Othoes Hirst Fam. Rhagodidae Rhagodalma Roewer Rhagodessa Roewer Rhagodeya Roewer Rhagodolus Roewer Rhagoduna Roewer Fam. Solpugidae Oparbella Roewer Ethiopia + – 1 2 1 – 1 8 3 Eritrea – – – – – – – – – Somalia + 2 – 1 – – 1 3 5 Djibouti + – – 2 – – – – – Sudan Senegal 21 1 + – Gambia Guinea 1 1 – – Guinea-Bissau Chad 5 1 + – Mali 2 – Niger 6 + 4 1 – – – – – – – – – – – – – – – – – – – 1 – 1 – – – – 1 – – – + – – + + + + + 5 1 – – 1 1 1 2 1 + – – – + – – – – – – – – – – 1 – – – – – – – 3 – – – – – – – 1 – – – – – – – – – 1 – – – – – 1 – – – – – – – + – – + + – – + – – – – – – – 1 (continued) 7.31 Africa Between 20oN and Zambezi-Kunene (Tropical or Intertropical Africa) 457 Table 7.5 (continued) Countries Solpugassa Roewer Zeria Simon Zeriassa Pocock Sudan Senegal 1 – Gambia Guinea – – Guinea-Bissau Chad – – Mali – Niger – 2 1 – – 2 – – – – – Countries Number of species Fam. Ceromidae Ceroma Karsch Fam. Daesiidae Biton Karsch Bitonupa Roewer Blossia Simon Hemiblossia Roewer Fam. Galeodidae Galeodes Olivier Fam. Rhagodidae Rhagodes Pocock Rhagodoca Roewer Rhagodolus Roewer Fam. Solpugidae Oparba Roewer Oparbella Roewer Solpuga Licht. Solpugassa Roewer Solpugella Roewer Solpugiba Roewer Solpugistella Turk Solpugylla Roewer Zeria Simon Zeriassa Pocock – – 1 – – – Nigeria Togo Cameroon 4 9 2 Malawi 4 Ruanda 1 Kenya 37 Tanzania 33 – – – – – – – – – + 2 – – – – – – – – – – + 2 – – – – – – – – – – – – + 1 + 3 1 2 1 + 4 + – – 3 – + 1 + + 2 – + 1 – – – – – – – + 1 + – – – – – – – – – – – – – – 7 2 – 1 – – – – – – + 1 – – – + 3 1 – – + – – – – + – – – – + – – – – + – – 1 – + – – – 1 – – – – 1 – – – – – – – – – – – – – – – – – 1 1 – – – 4 1 – 1 – 1 – 2 – – – 11 7 15 3 (continued) 458 7 Regional Arachnogeography Table 7.5 (continued) Countries Number of sp. Taxa Fam. Ceromidae Ceroma Karsch Fam. Hexisopodidae Chelypus Purcell Hexisopus Karsch Congo 1 DR Congo 30 Angola 31 Zambia 4 Mozambique 13 – + + – – – – 1 – 1 + – + – – – – 1 1 – – – 1 – – Representatives of six families are known from the described area: Ceromidae, Daesiidae, Galeodidae, Hexisopodidae, Rhagodidae, and Solpugidae (El-Hennawy 2008, Lawrence 1960, Roewer 1934, Simonetta and Delle Cave 1968, Harvey 2013b). None of the families is endemic or typical for tropical Africa. Endemic genera for this area: Blossiana Roewer (Ethiopia) Bitonota Roewer (Ethiopia) Bitonupa Roewer (Kenya) Solpugistella Turk (Kenya) Rhagodalma Roewer (Sudan) Rhagodippa Roewer (Djibouti) Benoit (1960) enumerated 29 species from the former Belgian Congo and Ruanda-Urundi. They all live far from the dense tropical forest of Congo and are concentrated mainly in Katanga. Ricinulei All Ricinulei in Africa (and the Old World) are known from west tropical Africa (11 spp., all of genus Ricinoides Ewing, 1929) (Guerin-Meneville 1838, Hansen 1921, Hansen and Soerensen 1904, Judson and Hardy 2001, Legg 1976, 1978, 1982, Naskrecki 2008, Tuxen 1974). 7.31 Africa Between 20oN and Zambezi-Kunene (Tropical or Intertropical Africa) 459 Map 7.19 Distribution of Ricinulei (Genus Ricinoides Ewing) in Africa Scorpiones In tropical Africa from 20oN to Zambezi-Kunene are known 21 genera of scorpions of the families Buthidae (13), Scorpionidae (three), Hemiscorpiidae (one), and Hormuridae (four) (Dupré 2013, Kovaøik 2001, 2003, Lamoral and Reynders 1975, Lourenço et al. 2010, Probst 1973, Vachon 1968) (Tables 7.6 and 7.7). Gambia 2 + – Guinea 15 + 1 Guinea-Bissau Liberia Ghana 3 3 5 + + 1 – – Burkina Faso 55 + – Mali 8 + – Congo 18 + 1 DR Congo 14 + 1 Gabon 9 + – Centr. AR 4 + – Angola 24 + – 1 – – – – – – 2 4 4 4 1 3 2 2 1 2 – – – – – – 3 – – 1 – – – – – – – – – – – 3 – 1 – 1 – – 1 – 1 – – 1 – 1 1 – 2 1 – – – 1 – – – 1 1 – – – – 2 – 1 – – 1 – – 1 – + 1 – + 1 – + 1 – + – 1 + – – + 1 – ++ 1 – + 2 – – 2 – – – – 1 – – 1 1 5 – 1 1 – – – 1 – – – – 4 – + 2 1 1 – – – – 1 – + – – 1 – – – – 1 1 – – – 2 – – – – + – 2 + 1 + – 3 + – + – 1 + – + – 1 + – + 5 – + 1 – – 2 – – – – – 1 1 2 1 1 Regional Arachnogeography Senegal 11 + – 7 Countries Number of species Fam. Buthidae Ananteroides Borelli Androctonus Ehrenberg Babycurus Karsch Buthacus Birula Butheoloides Hirst Buthus Leach Compsobuthus Vachon Hottentotta Birula Uroplectes Peters Fam. Scorpionidae Pandinus Thorell Scorpio L. Fam. Hormuridae Hormurus Thorell Liocheles Sundevall Opisthacanthus Peters 460 Table 7.6 Scorpions in Western and Central Tropical Africa Benin 4 Nigeria 7 Niger 14 Cameroon 17 Equatorial Guinea 5 + – 1 2 – – – – – – 1 – – – + 1 – – – – + 1 – 1 – – – – – – 1 – – – + 1 – – – – + – – 2 1 1 – – – – 1 – – 1 + 1 – – – – + – 3 1 2 1 1 1 2 – 1 – – – + 1 1 – – – + – – 3 – – 1 – – 1 2 1 1 1 + 1 1 + 1 2 + – – 2 – – – – – – – – – – + 1 – + – 2 Africa Between 20oN and Zambezi-Kunene (Tropical or Intertropical Africa) Togo 5 7.31 Countries Number of sp. Taxa Fam. Buthidae Akentrobuthus Lam. Androctonus Ehrenberg Babycurus Karsch Buthacus Birula Butheoloides Hirst Buthus Leach Cicileus Vachon Congobuthus Lour. Darchenia Vachon Hottentotta Birula Leiurus Ehrenberg Lychasioides Vachon Uroplectes Peters Fam. Scorpionidae Pandinus Thorell Scorpio L. Fam. Hormuridae Hormurus Thorell Opisthacanthus Peters 461 462 7 Regional Arachnogeography Table 7.7 Scorpions in Eastern Tropical Africa Countries Number of sp. Taxa Fam. Buthidae Androctonus Ehr. Babycurus Karsch Buthacus Birula Butheoloides Hirst Butheolus Simon Buthus Leach Compsobuthus Vachon Gint Kovarik et al. Hottentotta Birula Isometrus Ehrenb. Lanzatus Kovařik Lychas C.L. Koch Microbuthus Kraep. Neobuthus Hirst Odonturus Karsch Orthochiroides Kov. Orthochirus Karsch Parabuthus Pocock Sabinebuthus Lour. Somalibuthus Kov. Somalicharmus Kovařik Uroplectes Peters Uroplectoides Lour. Fam. Scorpionidae Pandinus Thorell Fam. Hemiscorpiidae Hemiscorpius Peters Fam. Hormuridae Iomachus Pocock Countries Number of sp. Taxa Fam. Buthidae Babycurus Karsch Hottentotta Birula Isometrus Ehrenb. Lychas C.L. Koch Odonturus Karsch Parabuthus Pocock Sudan 23 Ethiopia 43 Erythraea 17 Djibouti 12 + 1 4 1 2 – 3 3 1 6 1 – 1 1 2 – – – 4 – – – 1 + 1 4 – – – 2 4 2 3 1 1 2 2 1 1 1 1 6 1 1 1 3 + – 1 2 – 1 1 1 – 2 – – – 1 1 – – – 4 – – – – + – – – – – 1 1 – 2 – – – 2 1 – – 1 2 1 – – – + 10 – – + 1 + 14 + 2 – – + 3 + 1 – – + 1 – – – – Kenya 2 Tanzania 29 Uganda 6 Malawi 7 + 4 4 – 2 1 2 + 5 3 – – – – + 1 4 1 3 1 3 + – 1 – 1 – – (continued) 7.31 Africa Between 20 oN and Zambezi-Kunene (Tropical or Intertropical Africa) Table 7.7 (continued) Countries Riftobuthus Lour. et al. Uroplectes Peters Uroplectoides Lour. Fam. Scorpionidae Opistophthalmus C.L. Koch Pandinus Thorell Fam. Hormuridae Chiromachus Pocock Iomachus Pocock Opisthacanthus Peters Kenya 1 1 1 + – Tanzania – 5 – + – Uganda – – – + 2 Malawi – – – + 1 3 + – 1 – 2 + 1 1 1 1 + – 1 – 1 + 2 1 – 463 Some endemic genera of Scorpiones in the area: Fam. Buthidae: Riftobuthus Lourenço, Duhem, et Cloudsley-Thompson (Kenya); Lanzatus Kovařik (Somalia) Pseudoscorpiones In tropical Africa (the described area) have been recorded pseudoscorpions of 16 families: Chthoniidae, Tridenchthoniidae, Feaellidae, Garypidae, Geogarypidae, Olpiidae, Ideoroncidae, Neobisiidae, Syarinidae, Cheiridiidae, Pseudochiridiidae, Sternophoridae, Chernetidae, Cheliferidae, Atemnidae, and Withiidae (Beier 1944, 1955, 1959, 1967, 1972, 1979, Heurtault 1970a, 1970b, 1970c, 1983, Mahnert 1981, 1982a, 1982b, 1982c, 1983a, 1983b, 1985, 1988, Redikorzev 1924). There are no endemic families of pseudoscorpions in tropical Africa. Some endemic genera of pseudoscorpions are: Fam. Chthoniidae Congochthonius Beier, 1959 – DR Congo (one sp.) Fam. Ideoroncidae Afroroncus Mahnert, 1981 – Kenya (two spp.) Nannoroncus Beier, 1955 – Kenya, Uganda (one sp.) Fam. Withiidae Aisthetowithius Beier, 1967 – Kenya, Tanzania (one sp.) Cryptowithius Beier, 1967 – Kenya (one sp.) Pogonowithius Beier, 1979 – DR Congo (one sp.) Trichotowithius Beier, 1944 – Ethiopia, Kenya (two spp.) Fam. Cheliferidae Chamberlinarius Heurtault, 1983 – Ivory Coast (one sp.) Fam. Atemnidae Synatemnus Beier, 1944 – Tanzania (two spp.) 464 7 Regional Arachnogeography Opiliones Giribet and Prieto (2003), Goodnight and Goodnight (1959), Santos and Prieto (2010), Sharma et al. (2011) Map 7.20 Map of Cyphophthalmi of Africa and Madagascar Ogovea – ◊ Parogovia– ♦ Marwe – ▲ Manangotria – ● Purcellia – ► Ankaratra – ◄ Parapurcellia – ■ 7.31 Africa Between 20oN and Zambezi-Kunene (Tropical or Intertropical Africa) 465 Cyphophthalmi From tropical Africa are known three genera: Ogovea Hansen et Soerensen, 1914, from the Gulf of Guinea (Cameroon, Congo, Bioko, three spp.); Parogovia Hansen, 1921, from Bioko, Gabon, and Sierra Leone (three spp.); and the aberrant genus Marwe Shear, 1985, from Kenya (one sp.). The family Ogoveidae is endemic for the Gulf of Guinea. Fam. Neogoveidae has one genus in Africa (Parogovia Hansen); the other genera live in South America. Marwe belongs (?) to Sironidae (Hansen 1921, Hansen and Soerensen 1914, Shear 1985). Eupnoi According to the catalogues of Staręga (1984, 1992, with suppl.), in the described area are recorded Eupnoi of the fam. Phalangiidae. Fam. Phalangiidae (Phalangiinae) – Camerobunus Staręga et Snegovaya (1 sp.), Cristina Loman (13 spp.), Rhampsinitus Simon (47 spp.), Guruia Loman, Coptophalangium Staręga (1 sp.), Odontobunus Roewer, Dacnopilio Roewer, Megistobunus Hansen, ?Hindreus Kauri Endemic (known only from one country) are the genera: Fam. Phalangiidae Camerobunus Staręga et Snegovaya, 2008 – Cameroon Coptophalangium Staręga, 1984 – Ethiopia Dyspnoi – not recorded Laniatores (Kauri 1985, Roewer, 1912, 1927, 1935, 1949a, 1950b, 1951a, 1952, 1954, 1957) According to the catalogues of Staręga (1984, 1992, with suppl.), in the described area are recorded Laniatores from 35 genera and 5 families. Fam. Assamiidae – Aberdereca Goodnight et Goodnight (1 sp.), Bambereca Kauri (1), Bwitonatus Roewer (1), Comereca Roewer (1), Ereca Soerensen (23), Erecella Roewer (9), Erecula Roewer (7), Eusidama Roewer (1), Hypoxestus Loman (10), Metereca Roewer (= Leleupereca Roewer), Lygippulus Roewer (5), Metarhagdopygus Roewer, Randilea Roewer (1), Sesostris Soerensen (5), Sesostrellus Roewer (3), Spinixestus Roewer (6) Fam. Pyramidopidae – Guinea-Bissau, Gambia, Equatorial Guinea, DR Congo, Tanzania, Ghana, Togo, Ivory Coast, São Tomé and Príncipe, Fuerteventura (Canary Islands) (13 genera, ca. 40 spp.) Fam. Podoctidae (incl. Erecananinae and Ibaloniinae) – Erecanana Strand Fam. Biantidae – Hinzuanius Karsch, Metabiantes Roewer, Monobiantes Lawrence, Proconomma Roewer 466 7 Regional Arachnogeography Fam. Samoidae – Microconomma Roewer, Cameroon; Tetebius Roewer, Mozambique Some endemics (known only from one country) are the genera (almost all described by Roewer and monotypic): Microconomma Roewer – Cameroon Tetebius Roewer – Mozambique Zairebiantes Kauri Prolacurbs Roewer – DR Congo Monobiantes Lawrence – Kenya Ivobiantes Lawrence – Ivory Coast Heterolacurbs Roewer – Togo Eubiantes Roewer – Tanzania Clinobiantes Roewer – Cameroon Biantomma Roewer – Bioco, Equ. Guinea Biantella Roewer – Cameroon Vilhena Lawrence – Angola Uviranus Kauri – DR Congo Umbonimba Roewer – Ivory Coast Typhloburista Lawrence – Ivory Coast Typhlobunus Roewer – Kenya Tusipulla Roewer – Tanzania Tubereca Kauri – Rwanda Thomecola Roewer – São Tomé Sokodea Roewer – Togo Simienatus Roewer – Ethiopia Seuthessus Kauri – DR Congo Seuthesplus Roewer – Ivory Coast Selencula Roewer – Bioco (Equatorial Guinea) Scabrosidama Lawrence – Tanzania Santhomea Roewer – São Tomé Roewereca Lawrence – Tanzania Randilea Roewer – Uganda (Elgon) Pulchandrus Kauri – DR Congo Podaucheniellus Roewer – Cameroon Parasesostris Roewer – Tanzania Parakodaika Goodnight et Goodnight – Angola Palmanella Roewer – Sao Tomé Numipedia Kauri – DR Congo Nkogoa Roewer – Congo Orsimonia Roewer – Ethiopia Neocoryphus Lawrence – Ivory Coast Neobaeorix Lawrence – Tanzania Mwenga Kauri – DR Congo Mutadia Kauri – DR Congo 7.31 Africa Between 20oN and Zambezi-Kunene (Tropical or Intertropical Africa) Musola Roewer – Bioko (Equatorial Guinea) Montereca Lawrence – Tanzania Metasidama Roewer – Tanzania Metasesostris Roewer – Tanzania Merucola Roewer – Tanzania Maracandinus Roewer – Ethiopia Mandaria Roewer – Cameroon Machadoessa Lawrence – Angola Lygippus Roewer – Angola Lucandamila Roewer – Tanzania Lubudia Roewer – DR Congo Lossidacola Roewer – Tanzania Lossida Roewer – Tanzania Lisposidama Lawrence – Tanzania Leleupiolus Roewer – DR Congo Kungwea Roewer – Tanzania Kituvia Kauri – DR Congo Karangania Kauri – DR Congo Kasaina Lawrence – DR Congo Kakontwea Roewer – DR Congo Jaundea Roewer – Cameroon Ivocoryphus Lawrence – Ivory Coast Irumua Roewer – DR Congo Irnia Roewer – Ethiopia Humbea Roewer – Angola Henriquea Roewer – Island Principé Gulufia Roewer – Ethiopia Gomezyta Roewer – Ethiopia Fizibius Roewer – DR Congo Fakoa Roewer – Cameroon Eusidama Roewer – Tanzania Eregonda Roewer – DR Congo Erecops Roewer – Tanzania Erecongoa Roewer – DR Congo Erecabia Roewer – Tanzania Erebalda Roewer – Tanzania Ereala Roewer – DR Congo Edeala Roewer – Cameroon Dongila Roewer – Congo Djemia Roewer – Ethiopia Cangonia Roewer – DR Congo Cogonella Roewer – DR Congo Congolla Roewer – DR Congo Comereca Roewer – DR Congo Cereodiscus Roewer – Cameroon 467 468 Cereipes Roewer – Cameroon Celimba Roewer – Tanzania Cassinia Roewer – Guinea-Bissau Bwitonatus Roewer – DR Congo Buniabia Roewer – DR Congo Bundukia Lawrence – Tanzania Buemba Roewer – DR Congo Bueana Roewer – Cameroon Bolama Roewer – Guinea-Bissau Blantyrea Roewer – Malawi Binderia Roewer – Chad Bibundina Roewer – Cameroon Banconyx Lawrence – Ivory Coast Bancoella Lawrence – Ivory Coast Bambereca Kauri – DR Congo Angopygoplus Lawrence – Angola Angolyppa Lawrence – Angola Amhara Pavesi – Ethiopia Allereca Roewer – Rwanda Afroassamia Caporiacco – Ethiopia Adamauna Roewer – Cameroon Acanthophrysella Strand – Tanzania Acaca Roewer – Ethiopia Aburistella Roewer – Ivory Coast Aburista Roewer – Ghana Aberdereca Goodnight et Goodn. – Kenya Abanatus Roewer – DR Congo Tonkouinatus Roewer – Ivory Coast Scufia Roewer – Togo Proconomma Roewer – DR Congo Opconommula Roewer – Cameroon Opconomma Roewer – São Tomé Nimbadus Roewer – Ivory Coast Micronimba Roewer – Ivory Coast Metaconomma Kauri – DR Congo Kwangonia Kauri – DR Congo Aburiplus Roewer – Ghana 7 Regional Arachnogeography 7.31 Africa Between 20oN and Zambezi-Kunene (Tropical or Intertropical Africa) 469 Amblypygi The first list of African Amblypygi (Lawrence 1969a) contained 16 spp. Without the North African Musicodamon and the island Charinus spp. from Socotra, the Seychelles, and Madagascar, 13 spp. remain from tropical Africa (Delle Cave 1986; Fage 1939, 1954; Hansen 1921; Lawrence 1958; Prendini et al. 2005; Weygoldt 1972, 1995, 1999, 2000). Meanwhile several nomenclatorial changes took place. The new genus Euphrynichus Weigoldt, 1995, has been described; Titanodamon Pocock was synonymized with Damon C.L. Koch, Myodalis Simon, and with Phrynichus Karsch. Paracharon caecus Hansen should be added to the list. “Hemiphrynus” machadoi Fage is now Xerophrynus Weigoldt, 1996 (endemic). Now (2017) in tropical Africa are known 26 spp. of Amblypygi of seven genera and three families (Charinidae, Phrynichidae, Paracharontidae). Fam. Charinidae Charinus Simon – widespread in the world, in continental tropical Africa known from Guinea, Equatorial Guinea, São Tomé, and Somalia (six spp.). Fam. Phrynichidae Damon C.L. Koch – widespread and endemic in tropical Africa (11 spp., nine in tropical Africa, north of Zambezi-Kunene). In his revision of the genus, Weygoldt (2006) concluded that there are four species confined to West Africa (which evolved mostly in rain forests) and six species (mostly smaller and capable to survive in dryer habitats) distributed from East Africa to South Africa and Namibia. Euphrynichus Weygoldt – Kenya, Tanzania, Zimbabwe, Angola, Malawi (two spp. – E. amanica (Werner, 1916), E. bacillifer (Gerstaecker, 1873) Phrynichus Karsch – Cameroon, Eritrea, Somalia, Djibouti, Rwanda, Kenya, Tanzania, Congo (seven spp.) Xerophrynus Weigoldt – Angola (one sp. X. machadoi with unclear position), Namibia Fam. Paracharontidae Paracharon Hansen – P. caecus Hansen, 1921 The most interesting amblypygid in tropical Africa is Paracharon Hansen from Guinea-Bissau (one sp., P. caecus Hansen, 1921), the only member of the family Paracharontidae and the suborder Palaeoamblypygi (“living fossil”). 470 Map 7.21 Amblypygi and Uropygi in Africa Charinus – x Paracharon – Musicodamon – ▲ Euphrynichus – ▼ Etienneus (Uropygi) – ♣ Damon – ● Phrynichus – ■ Xerophrynus – ◊ Phrynichodamon – □ 7 Regional Arachnogeography 7.31 Africa Between 20oN and Zambezi-Kunene (Tropical or Intertropical Africa) 471 Thelyphonida (Uropygi) The enigmatic Etienneus africanus (Hentschel, 1899) from Senegal, Gambia, Guinea, and Guinea-Bissau is now considered autochthonous and relict (Huff and Prendini 2009). Nowhere else in Africa live Uropygi (Cooke and Shadab 1973, Harvey 2013d, Hentschel 1899, Heurtault 1984, Huff and Prendini 2009). Schizomida From tropical Africa are known five genera of Schizomida (four endemic) of seven species (all endemic), all of family Hubbardiidae (Hubbardiinae) (Lawrence 1969b, Reddell and Cokendolpher 1995, Harvey 2013g, Armas 2014). Fam. Hubbardiidae Enigmazomus Harvey, 2006 – Somalia, Arabian Peninsula (two spp.) Afrozomus Reddell et Cokendolpher, 1995 – Angola, Congo (one sp.) Artacarus Cook, 1899 – Liberia, Ivory Coast (one sp.) Kenyazomus Armas, 2014 – Kenya (one sp.) Lawrencezomus Armas, 2014 – Cameroon, Liberia (two spp.) 472 7 Regional Arachnogeography Map 7.22 Distribution of Schizomida in Africa, Socotra, Seychelles, and Madagascar Megaschizomus – X Enigmazomus – ▼ Kenyazomus – ▲ Lawrencezomus – Mahezomus – Afrozomus – ► Artacarus – □ Bamazomus – ■ Anepsiozomus – ◊ Ovozomus – Secozomus – 7.31 Africa Between 20oN and Zambezi-Kunene (Tropical or Intertropical Africa) 473 Araneae The spiders of Africa south of Sahara have been (and are) studied by many arachnologists, and among them are Benoit (many papers), Griswold 1991, Holm 1962, Scharff 1992, and others. According to Scharff (1990), in Africa south of Sahara are considered valid 365 species of Linyphiidae of 67 genera. Some endemic genera of spiders in tropical Africa from 20oN to Zambezi-Kunene: Fam. Theridiidae Pycnoepisinus Wunderlich, 2008 – Kenya (one sp.) Fam. Theraphosidae Bacillochilus Gallon, 2010 – Angola (one sp.) Batesiella Pocock, 1903 – Cameroon (one sp.) Encyocratella Strand, 1907 – Tanzania (one sp.) Eucratoscelus Pocock, 1898 – Kenya, Tanzania (two spp.) Loxomphalia Simon, 1889 – Zanzibar (one) Loxoptygus Simon, 1903 – Ethiopia (three spp.) Pelinobius Karsch, 1885 – Kenya, Tanzania (one sp.) Fam. Liocranidae Cteniogaster Bosselaers et Jocqué, 2013 – Tanzania, Kenya (seven spp.) ToxoniellaWarni et Jocqué, 2002 – Kenya (two spp.) Fam. Mimetidae Kratochvilia Strand, 1934 – Principe (one sp.) Fam. Mysmenidae Kilifina Baert et Murphy, 1992 – Kenya (one sp.) Leviola Miller, 1970 – Angola (one sp.) Fam. Oecobiidae Urocteana Roewer, 1961 – Senegal (one sp.) Fam. Oonopidae Anophthalmoonops Benoit, 1976 – Angola Blanioonops Simon et Fage, 1922 – East Africa (one sp.) Caecoonops Benoit, 1964 – Congo (two spp.) Hypnoonops Benoit, 1977 – Congo (one sp.) Kijabe Berland, 1914 – Kenya, East Africa (two spp.) Termitoonops Benoit, 1964 – Congo (five spp.) Zyngoonops Benoit, 1977 – Congo (one sp.) 474 7 Regional Arachnogeography Fam. Palpimanidae Badia Roewer, 1961 – Senegal (one sp.) Fam. Phyxelididae Kulalania Griswold, 1990 – Kenya (one sp.) Fam. Pisauridae Cispinilus Roewer, 1955 – Central Africa (one sp.) Conakrya Schmidt, 1956 – Guinea (one sp.) Tapinothele Simon, 1898 – Zanzibar (one sp.) Vuattouxia Blandin, 1979 – Ivory Coast (one sp.) Fam. Prodidomidae Katumbea Cooke, 1964 – Tanzania (one sp.) Plutonodomus Cooke, 1964 – Tanzania (one sp.) Theumella Strand, 1906 – Ethiopia (two spp.) Fam. Sparassidae Berlandia Lessert, 1921 – East Africa (two spp.) Sarotesius Pocock, 1898 – East Africa (one sp.) Stasinoides Berland, 1922 – Ethiopia (one sp.) Fam. Tetrablemmidae Anansia Lehtinen, 1981 – Angola (one sp.) Cuangoblemma Brignoli, 1974 – Angola (one sp.) Hexablemma Berland, 1920 – Kenya (one sp.) Fam. Tetragnathidae Parazilia Lessert, 1938 – Congo Fam. Anapidae Forsteriola Brignoli, 1981 – Congo, Burundi, Rwanda (two spp.) Fam. Ctenidae Arctenus Polotow et Jocqué, 2014 – Kenya (one sp.) Fam. Dictynidae Hoplolathys Caporiacco, 1947 – Ethiopia (one sp.) Mashimo Lehtinen, 1967 – Zambia (one sp.) Fam. Zodariidae Mastidiores Jocqué, 1987 – Kenya (one sp.) 7.32 Differences Between the Arachnofaunas of East and West Africa 475 Fam. Thomisidae Felsina Simon, 1895 – West Africa (one sp.) Gnoerichia Dahl, 1907 – Cameroon (one sp.) Haedanula Caporiacco, 1941 – Ethiopia (one sp.) Heriaesynaema Caporiacco, 1939 – Ethiopia (one sp.) Hewittia Lessert, 1928 – Congo (one sp.) Mystaria Simon, 1895 – West Africa (two spp.) Ostanes Simon, 1895 – West Africa (one sp.) Parasmodix Jezequel, 1966 – Ivory Coast (one sp.) Pasiasula Roewer, 1942 – Bioco (one sp.) Opilioacarida The known opilioacarids of tropical Africa belong to four species: Phalangiacarus brosseti Coineau et Van der Hammen, 1979, from Gabon; Salfacarus tanzaniensis Van der Hammen, 1977, from Tanzania; and two Panchaetes from Ivory Coast and Angola. The genera Panchaetes Naudo and Phalangiacarus Coineau et Van der Hammen are known only from tropical Africa (Beron 2014). Acariformes: Beron (2008a), Niedbala (2002) Prostigmata Fam. Erythraeidae Lomeustium Haitlinger, 2006 – one sp. (Togo, Ghana, Benin) (larval, endemic genus) Opserythraeus Fain, 1996 – one sp. (Rwanda) (larval, endemic genus) 7.32 Differences Between the Arachnofaunas of East and West Africa In many manuals in zoogeography (Geptner 1936), delimitation of tropical Africa is made between the eastern part (mostly savanna, including also the highest mountains) and western part (mostly rain forest, or what remains from it, and savanna forest). Usually they are considered at level of subregions. Eastern part. Geptner 1936: East Africa plus Sahel and the area between DR Congo and Southern Africa Western part. Geptner 1936: From Senegal to (including) DR Congo and Northern Angola Palpigradi – endemic genus in West Africa, Allokoenenia (Guinea); in East Africa, none. The same family. Ricinulei – in the Old World only in West Africa (end. genus Ricinoides in Gambia, Guinea, Guinea-Bissau, Nigeria, Ghana, Cameroon, Sierra Leone, Equatorial Guinea, Benin, Congo, Togo, Ivory Coast 476 7 Regional Arachnogeography Amblypygi – in West Africa (Guinea-Bissau) endemic suborder (Palaeoamblypygi) with one family, genus, and species Thelyphonida (Uropygi) – in Africa only in the western part (end. genus Etienneus in Senegal, Guinea-Bissau, Gambia, Guinea). Relicts? Schizomida – endemic genera in West Africa: Afrozomus, Artacarus, Lawrencezomus; in East Africa, Enigmazomus, Kenyazomus. No endemic families. Scorpiones – endemic genera in West Africa In East Africa: Somalibuthus, Riftobuthus, Somalicharmus, Lanzaia. No endemic families. Pseudoscorpiones – endemic genera in West Africa: Congochthonius, Pogonowithius, Chamberlinarius; In East Africa: Afroroncus, Nannoroncus, Aisthetowithius, Cryptowithius, Trichotowithius, Synatemnus. No endemic families. Opiliones Cyphophthalmi – endemics in West Africa, Ogoveidae (endemic family), Ogovea, Paragovia; in East Africa, Marwe (Kenya) Opilioacarida – in West Africa two endemic genera (Panchaetes, Angola, Ivory Coast; Phalangiacarus, Gabon); in Tanzania the genus Salfacarus, known also from Southern Africa and Madagascar Holothyrida – not living on the African continent, but found in the Seychelles near East Africa Conclusion In West Africa there is one endemic suborder (Palaeoamblypygi), one order living in Africa only in the western part (Ricinulei), and two endemic families, living only in the western part of Africa. In East Africa there are no orders or suborders living only there (Holothyrida on the Seychelles). In both parts there are many endemic genera of various orders. The differences between both parts are mainly in the landscape and the flora. 7.33 7.33.1 Tropical South America Geography, General Zoogeography, and Paleogeography The South American continent between Panama and the 30oC is the undisputed core of Neotropica in the classical subdivision of the world. Its center is the great Amazonian selva and its backbone are the Andes. It contains also the cold plains of Bolivia (the Altiplano) and many other specific features. The chain of the Andes reaches 6961 m (Aconcagua), and many of the world records of high-altitude arachnids (Solifugae, etc.) are registered in Peruvian and 7.33 Tropical South America 477 other parts of the Andes. In Peru we visited once the highest high forests in the world (Polylepis forest, with trees as tall as 15 meters at an altitude of 5500 m). The tropical zone of South America consists of three fundamentals of vegetal formations: the wet tropical forest; the Serrados et similar formations of savanna type, corresponding to open non xerophytic formations; the Caatingas, open xerophytic formations. The wet tropical forest includes two vast regions: Hileia and Atlantic forest. (Lourenço 1986) What concerns the climatic regions of South America, after Eidt (1968), is that we follow the Köppen system – four major types of climate: A, B, C, and E. The A climates, or those in which average monthly temperatures are all above 18oC, are divided into two primary types: the ever-humid or tropical rain forest variety (Af) and the “winter” dry season or savanna type (Aw). The four zones of Af are the Pacific coast of Colombia and Ecuador, Amazon Basin, Bahia-Victoria coast of Brazil, and Rio de Janeiro-Santos coast of Brazil. Monthly and annual temperature ranges are less than 3oC throughout the entire area with the exception of some of the highest valleys in the southwest. In the Andes this zone coincides with the so-called Tierra Caliente and Tierra Templada. The second type of A climate, the savanna of Aw variety, is with a definite dry season with insufficient moisture to support a true rain forest vegetation. The Neotropical region has been subdivided many times (Cabrera and Willink 1973, 1980, Cabrera and Yepes 1940, Sclater and Sclater 1999, Eidt 1968, Harrington 1962, Martin 1968, Mello-Leitão 1931, 1939, Morrone 2001, 2006, 2010, 2015, Müller 1974, Roig-Juñent et al. 2006). Here is one biogeographic subdivision by Morrone (2006): The Nearctic region, inhabited by Holarctic insect taxa, comprises five provinces: California, Baja California, Sonora, Mexican Plateau, and Tamaulipas. The Mexican transition zone comprises five provinces: Sierra Madre Occidental, Sierra Madre Oriental, Transmexican Volcanic Belt, Balsas Basin, and Sierra Madre del Sur. The Neotropical region, which harbors many insect taxa with close relatives in the tropical areas of the Old World, comprises four subregions: Caribbean, Amazonian, Chacoan, and Parana. The South American transition zone comprises five provinces: North Andean Paramo, Coastal Peruvian Desert, Puna, Atacama, Prepuna, and Monte. The Andean region, which harbors insect taxa with close relatives in the Austral continents, comprises three subregions: Central Chilean, Subantarctic, and Patagonian. The most important of the biogeographic subdivisions of Morrone is the dividing of South America into two regions: Neotropical (mostly lowlands) and Andean (mostly mountainous, but including the southernmost plains of Patagonia). The separation is along the meridian and not across the continent. The “Andean region” includes very long band, from Cape Horn to Venezuela. The “Caribbean” Subregion of Morrone almost separates from the main area of the “Amazonian, Chacoan, and Parana” subregions, which is not very practical. Most classical zoogeographers consider the whole of South America as an entire Neotropical region (some of them separating the far south with its relations to Australian or/and Neozealandian faunas). 478 7.33.2 7 Regional Arachnogeography Arachnogeography South America, one of the richest zoogeographical territories in the world, was longtime considered to represent one (the only) region (Neotropical) of the Neogean Kingdom. However, the difference – climatic and paleogeographical – of Patagonia with the tropical part of the continent has been noticed since longtime. The presence of many elements in the Patagonian fauna close to the fauna of Australia and New Zealand was incited by some authors (Lopatin 1980) to include the south of Chile and Argentina in the Notogean Kingdom. We divided conditionally South America in Patagonian (south of 30oS) and tropical parts, in order to examine their arachnofauna separately. The very active work of Brazilian, Argentinian, and Mexican arachnologists made possible to elucidate many aspects in the arachnofauna of the continent – see Mello-Leitão (1937) and Ringuelet (1956, 1978). In tropical South America are recorded so far most of the orders and suborders of Arachnida: Palpigradi (no endemics above species), Solifugae (2 families, one of them endemic for tropical South America), Ricinulei (17 spp. of 1 genus, only end. species), Scorpiones (3 end. genera; ?1 end. family in caves); Brazil is one of the richest countries in scorpions (165 spp. of 26 genera): Pseudoscorpiones (16 families, 18 endemic genera), Opiliones (Cyphophthalmi, 5 endemic genera; Eupnoi, 1 endemic genus; Laniatores, huge variety, with 15 fam., 13 of them endemic), Amblypygi (many species, 1 endemic genus, the only genus in the family Phrynichidae living in the Western Hemisphere), Uropygi (2 endemic genera), Schizomida (8 genera), and Araneae (spiders of 77 families are known south of Panama (72% of all spider families)). No family is endemic for the Southamerican continent. There are Opilioacarida (two spp.), Holothyrida (two endemic genera), many mites. Absent are the suborders Palaeoamblypygi, Dyspnoi, and Mesothelae. As a whole, the level of endemism is too low to characterize a kingdom. The regions, subregions, and provinces of Morrone (2001) are characterized by many taxa, but very few of them (and not the most typical) are among the arachnids. In their analysis of the biogeographic history of South American arid lands, Roig-Juñent et al. (2006) outlined 21 areas of endemism south of 5oS. Among the taxa used were spiders (Nemesiidae: Acanthogonatus Karsch and Diplothelopsini) and scorpions. Their main conclusion was that “That area of endemism of Patagonia form a natural group, showing that this biota evolved as a unit, as well the biota that occurs in the areas of Central Chile.” 7.33 Tropical South America Map 7.23 Zoogeographical subdivision of South America 479 480 7 Regional Arachnogeography (I) (after Cabrera and Yepes 1940) – 1. Sabánica, 2. Amazonica, 3. Tropical, 4. Subtropical, 5. Tupi, 6. Pampasica, 7. Patagónica, 8. Subandino, 9. Chileno, 10. Andino, 11. Incásico (II) (after Mello-Leitão 1931) – 1. Guayano-Amazonica, 2. Andina, 3. BororôCariri, 4. Tupi-Guarani, 5. Patagónica (III) (after Mello-Leitão 1937) – 1. Caribea, 2. Hyléa, 3. Gê, 4. Bororo, 5. Tupi, 6. Guarani, 7. Andino-Patagonica (IV) (Sclater and Sclater 1999) – 1. Colombiana, 2. Amazonica, 3. Sud-brasileira, 4. Patagónica Map 7.24 Regions of the world, with indication of the transition zones in gray (After Morrone 2015b) 1, Mexican; 2, Sahar-Arabian; 3, Chinese; 4, South American; 5, Indomalayan 7.33 Tropical South America 481 According to the biogeographical subdivision in Morrone (2001), the sierra in Ecuador belongs to the Andean region and the other three areas (Costa, Oriente, and Insular) – to the Neotropical region. Palpigradi Very few Palpigradi have been recorded from South America – see Condé (1979, 1986, 1993), Hansen (1901), and Mello-Leitão and Arlé (1935). Condé (1996) enumerates Eukoenenia improvisa Condé, 1979, and Koeneniodes notabilis Silvestri, 1913, from French Guiana; E. grassii (Silvestri in litt., Hansen, 1901) from Paraguay; and E. janetscheki Condé, 1993, and E. roquettei (MelloLeitão et Arlé, 1935) from Brazil. The first troglobitic Palpigradi from South America is Eukoenenia spelunca Souza et Ferreira, 2010, again from Brazil. Followed several new species from the caves of Brazil (Sousa and Ferreira 2011, 2012a, 2012b) and two Leptokoenenia (the first in the New World) have been described from an iron ore mine in Brazil (Sousa and Ferreira 2013). The widespread E. florenciae is known also from Colombia. Solifugae In tropical (north of Chile and Argentina and south of Panama) South America are known representatives of two families of Solifugae: Ammotrechidae and Mummuciidae (?14) (Kraepelin 1900, Maury 1982, Mello-Leitão 1937, Muma 1976, Roewer 1934, 1957). Endemic genera are: Fam. Mummuciidae (and five genera) Gauchella Mello-Leitão, 1937 – Bolivia (one sp.) Metacleobis Roewer, 1934 – Brazil (one sp.) Mummucina Roewer, 1934 – Chile, (five spp.) Mummuciona Roewer, 1934 – Venezuela (one sp.) Mummucipes Roewer, 1934 – Paraguay (one sp.) Fam. Ammotrechidae Ammotrechinae Campostrecha Mello-Leitão, 1937 – Ecuador (one sp.) 482 7 Regional Arachnogeography Saronominae Saronomus Kraepelin, 1900 – Colombia, Venezuela (one sp.) Subfamily not assigned Eutrecha Maury, 1982 – Venezuela (one sp.) Xenotrecha Maury, 1982 — Venezuela (one sp.) Ricinulei So far 17 spp. have been recorded from South America, the southernmost localities being in Peru and Brazil. They belong to the single genus Cryptocellus Westwood (Ricinoididae), which is found also in Central America and Cuba (Cokendolpher 2000, Cooke 1967, Cooreman 1977, Dumitresco and Juvara-Balş 1977, Ewing 1929, González-Sponga 1998, Platnick 1977, 1980, 1981, 1988, Platnick and Paz 1979, Roewer 1952, Tourinho and Salette de Azevedo 2007, Tourinho et al. 2010, 2014, Pinto-da-Rocha and Andrade 2012, Pinto-da-Rocha and Bonaldo 2007). Scorpiones In South America are represented eight families of scorpions: Buthidae, Chactidae, Hormuridae, Bothriuridae, Troglotayosicidae, Scorpionidae, Caraboctonidae, and Euscorpiidae (Mello-Leitão 1942, 1945, 1949, Lourenço 1981, 1986, 1987, Lourenço et al. 2004, Lourenço and Duhem 2009, Lourenço and Monod 2000, Ringuelet 1953) In Paraguay, according to Maury (1984), have been found until then seven spp. of Buthidae and five spp. of Bothriuridae. In Bolivia Acosta and Ochoa (2002) have listed 24 spp. of scorpions from the families Bothriuridae, Buthidae, and Iuridae. The Scorpiones of Ecuador, according to Lourenço (1995a), are 36 spp. of 8 genera and 4 families (Bothriuridae, Buthidae, Chactidae, and Iuridae). According to him, “Predominant distributional patterns show an Amazonian origin for some Buthidae (Ananteris and Tityus) and Chactidae (genus Chactas), whereas several other Buthidae (genus Tityus), Chactidae (genus Teuthraustes) and Iuridae (genus Hadruroides) represent a typical andine fauna.” Now the genus Hadruroides is considered a member of fam. Caraboctonidae (Soleglad and Fet 2003), and the number of Scorpiones in Ecuador is already 48 (or 47 on the continent and two on Galapagos, according to Brito and Borges 2015). Acosta (online) compiled a list of Argentinian scorpions, based on Acosta and Maury (1998). It contains the families Buthidae and Bothriuridae (Table 7.8). 7.33 Tropical South America 483 So far the following genera of scorpions are known only from tropical South America: Brazilobothriurus Lourenço et Monod, 2000; Troglorhopalurus Lourenço, Baptista, et Giupponi, 2004; Troglotayosicus Lourenço, 1981 Fam. Troglotayosicidae – if the European Belisarius Simon does not belong to this family, the family would be endemic for tropical South America. Pseudoscorpiones The checklist of Ceballos and Florez (2007) lists for Colombia only 22 spp. of Pseudoscorpiones of 18 genera and 6 families. Mahnert and Adis (2002) enumerate from Amazonia 34 genera of 12 families. According to the catalogue of Harvey (1990) and Harvey (2011, 2013f) and the supplementary publications from tropical South America (between Panama and 30oS) are recorded pseudoscorpions from 17 families: Chthoniidae, Lechytiidae, Tridenchthoniidae, Cheiridiidae, Feaellidae, Bochicidae, Syarinidae, Ideoroncidae, Geogarypidae, Garypidae, Garypinidae, Gymnobisiidae, Olpiidae, Atemnidae, Cheliferidae, Chernetidae, and Withiidae (Beier 1959a, Heurtault 1986c, Andrade and Mahnert, 2003, Mahnert 1979, 1984, 1985b, 1985c, 1994, 2001, 2009, Mahnert and Adis 1985, Mahnert et al. 1986, Harvey et al. 2016). The following are the number of pseudoscorpion species in different countries: Brazil, 164; Ecuador, 59; Peru, 42; Colombia, 22; Venezuela, 62; Guyana, 9; French Guiana, 1; Suriname, 5; and Bolivia, 6. Some of these countries are clearly understudied. From this area endemic genera are: Fam. Tridenchthoniidae Cryptoditha J.C. Chamberlin et R.V. Chamberlin, 1945 – Brazil (one sp.) Haploditha Caporiacco, 1951 – Venezuela (one sp.) Neoditha Feio, 1945 – Brazil (one sp.) Sororoditha J.C. Chamberlin et R.V. Chamberlin, 1945 – Brazil (one sp.) Spelaeobochica Mahnert, 2001 – Brazil (one sp.) Fam. Feaellidae Iporangella Harvey, Andrade, et Pinto-da-Rocha, 2016 – Brazil (one sp.) Fam. Ideoroncidae Ideoroncus Balzan, 1887 – Brazil, Paraguay (eight spp.) Fam. Cheliferidae Lophodactylus J.C. Chamberlin, 1932 – Brazil (one sp.) Colom. 80 Venezuela Guyana 230 26 Suriname 14 French Guiana Uruguay 25 6 + 23 1 1 – – 1 1 10 60 1 1 + 3 4 10 2 5 1 3 4 – 6 + 13 4 – – – 2 – 2 32 – – + – – 1 15 – – – – – 1 + 31 3 1 1 – – – 2 81 – – + 1 10 15 26 3 3 – 3 2 27 + 2 – 1 – + 8 – – 1 – 1 – 1 4 – – + 5 1 1 – – – 1 1 – 6 + 8 – – 1 – 9 – 3 8 – – + – 1 2 – – – – 1 – 1 – – – 5 – – + 1 – 3 – – – – 1 – 1 + 1 1 – – – – – 1 – 1 – – – – – – – – – – – Paraguay 14 + – – – – – – – – 1 – – + – – – – – – – – – – + 1 1 – – – – – 5 – 1 – – – – – – – – – – – Regional Arachnogeography Brazil 165 7 Country Number of species Таxa Fam. Buthidae Ananteris Thorell Centruroides Marx Isometrus Ehrenberg Mesotityus G.-Sponga Microananteris Lour. Microtityus K.-Waer. Physoctonus M.-Leitão Rhopalurus Thorell Tityus C.L. Koch Troglorhopalurus Lour. et al. Zabius Thorell Fam. Chactidae Auyantepuia G.-Sponga Broteochactas Pocock Brotheas C.L. Koch Chactas Gervais Chactopsis Kraepelin Chactopsoides Ochoa et al. Guyanochactas Lour. Hadrurochactas Pocock Megachactops Ochoa et al. Neochactas Sol. et Fet 484 Table 7.8 Scorpions in South America Country Number of species Тaxa Fam. Buthidae Ananteris Thorell Centruroides Marx Brazil 4 2 + 3 + 13 1 1 3 – 1 – – – – (+) 1 Colom. 2 1 + 1 – – – – – – + 2 + 3 – - Venezuela 5 3 + 2 – – – – – – – – – + 3 – – Guyana – 1 – – – – – – – – – – – – – – – Suriname – – – – – – – – – – – – – – – – – French Guiana – – – 1 – – – – – – – – – – – – – Uruguay – – – – + 3 1 – – 1 – – – – – – (+) Paraguay – – – – + 3 1 – – 2 – – – – – (+) 1 – – – – + 3 – – – – 1 – – – – – – Аrgentina 66 Bolivia 27 Chile 53 Peru 62 Еcuador (cont.) 47 + 1 1 + 3 1 + – 1 + 1 1 + 3 2 7.33 Tropical South America Country Teuthraustes Simon Vachoniochactas G.–Sponga Fam. Hormuridae Opisthacanthus Peters Fam. Bothriuridae Bothriurus Peters Brachistosternus Pocock Brazilobothrius Lour. et Monod Thestylus Simon Timogenes Simon Urophonius Pocock Fam. Troglotayosicidae Troglotayosicus Lour. Fam. Scorpionidae Tarsoporosus Francke Fam. Euscorpiidae Euscorpius Thorell (introduced) (continued) 485 Country Isometrus Ehrenb. Tityus C.L. Koch Zabius Thorell Fam. Chactidae Brotheas C.L. Koch Chactas Gervais Chactopsis Kraepelin Teuthraustes Simon Fam. Caraboctonidae Caraboctonus Pocock Hadruroides Pocock Fam. Hormuridae Opisthacanthus Peters Fam. Troglotayosicidae Troglotayosicus Lour. 486 Table 7.8 (continued) Аrgentina 1 8 2 – – – – – – – – – – – – Bolivia – 9 – + 1 – – – – – – – – – – Chile – 1 – – – – – – + 1 – – – – – Peru 1 12 – + – 2 2 – + 1 16 + 1 – – Еcuador (cont.) – 16 – + – 3 – 12 + – 6 – – + 1 7 Regional Arachnogeography 7.33 Tropical South America 487 Fam. Atemnidae Brazilatemnus Muchmore, 1975 – Brazil (one sp.) Caecatemnus Mahnert, 1985 – Brazil (one sp.) Fam. Chernetidae Attaleachernes Mahnert, 2009 – Brazil (one sp.) Atherochernes Beier, 1954 – Venezuela (one sp.) Calidiochernes Beier, 1954 – Venezuela (one sp.) Ceratochernes Mahnert, 1994 – Venezuela (two spp.) Corosoma Karsch, 1879 – Brazil (one sp.) Dasychernes J.C. Chamberlin, 1929 – Colombia, Panama (four spp.) Spelaeochernes Mahnert, 2001 – Brazil (eight spp.) Attaleachernes Mahnert, 2009 – Brazil (one sp.) Fam. Withiidae Tropidowithius Beier, 1955 – Peru (one sp.) Opiliones South America is extremely rich in Opilions, mostly Laniatores (Acosta 2002, 2006, Benavides and Giribet 2007, 2013, DaSilva et al. 2010, Pinto-da-Rocha et al. 2012, González-Sponga – 19 papers from 1981 to 2005; 1987, 1992, Goodnight and Goodnight 1943, 1980, Hinton 1938, Kury 2003, Kury and Maury 1998, Kury and Pérez 2002, Kury and Pinto-da-Rocha 2002, Martens 1969, 1988, Mello-Leitão 1932, 1935, 1941, 1945, Pinto-da-Rocha 1997, Rambla 1976, Ringuelet 1959, Roewer 1913, 1932, 1943, 1949a, 1949b, 1949c, 1956, 1957, 1961, 1963, Rosas Costa 1950, Shear 1979, 1993b, H. E. M. Soares and Avram 1981, H. E. M. Soares 1979, Soares B.A.M. and H.E.M. Soares 1948). According to the statistics of Kury (2002 and onward), only in Brazil have been recorded 987 spp. of Opiliones (the richest country in the world). In the Neotropical region are known 20 fam. of Opilions, including 13 families endemic to the Neotropics (all Laniatores). Cyphophthalmi Typical for tropical South America is the family Neogoveidae (Colombia, Brazil, Ecuador, Guyana, French Guiana, Peru, Suriname, Trinidad and Tobago, Venezuela), which has also one genus in tropical Africa (Parogovia Hansen, 1921) and to the north goes as far as Southern USA (Metasiro). Five genera in South America: Brasilogovea Martens, 1969 – Brazil, Colombia (two spp.) (endemic) Canga DaSilva, Pinto-da-Rochaet Giribet, 2010 – Brazil (one sp., caves) (endemic) Huitaka Shear, 1979 – Colombia (seven spp.) (+ two undescr.) (endemic to Colombia) Metagovea Rosas Costa, 1950 – Colombia, Ecuador, Brazil (Manaus) (four spp.) (+17 undescr.) (endemic) Neogovea Hinton, 1938 – Brazil, Guyana (five spp.) (+12 undescr.) (endemic) 488 7 Regional Arachnogeography One genus of the family Pettalidae (Chileogovea Roewer, 1961) is known from Chile (two spp.); Ch. oedipus Roewer, 1961, from Concepción to Chiloé; and Ch. jocasta Shear, 1993, from Malleco and Arauco Provinces. Eupnoi Fam. Phalangiidae (Dicranopalpinae – Dicranopalpus Doleschall, 1852) – Venezuela Fam. Neopilionidae (Thrasychiroides Soares et Soares, 1947 – Brazil, one sp., the only endemic genus among the Eupnoi of tropical South America) Dyspnoi Fam. Acropsopilionidae (Acropsopilio Silvestri, 1904 – Venezuela) Laniatores According to the catalogue of Kury (2003), in tropical South America (south to 30oS), this suborder contains genera from the families Agoristenidae, Cosmetidae, Cranaidae, Escadabiidae, Fissiphaliidae, Gonyleptidae, Guasiniidae, Icaleptidae, Manaosbiidae, Kimulidae (Minuidae), Samoidae, Stygnidae, Stygnommatidae, Stygnopsidae, and Zalmoxidae. The 13 underlined families are entirely Neotropical. Many of the genera are monotypic and have been partly revised and synonymized by Pinto-da-Rocha et al. (2012). According to Kury (2003), up to 2002 in the New World have been recorded 2372 spp. of 746 genera and 21 families. Of course, with the activities of the South American specialists in the last 15 years, many of these figures are altered, the number of species is increased, but still the most numerous families remain: Gonyleptidae (in 2002 823 spp.) and Cosmetidae (710 spp.). In South America N of 30oN, there are at least 470 endemic genera of Laniatores (including 226 genera of Gonyleptidae). Endemic genera (from one or few countries, Argentina, Uruguay, and Chile excluded) in the described area are: Fam. Agoristenidae Leiosteninae Barinas González-Sponga, 1987; Barlovento González-Sponga, 1987; Ocoita González-Sponga, 1987; Paravima Caporiacco, 1951; Vimina González-Sponga, 1987 – Venezuela Leptostygnus Mello-Leitão, 1940 – Colombia, Venezuela Sabanilla Roewer, 1913 – Colombia?, Venezuela Vima Hirst, 1912 – Guyana Zamorinae Palcabius Roewer, 1956, Ramonus Roewer, 1956 – Peru Zamora Roewer, 1927 – Ecuador Fam. Cosmetidae Acantholibitia Mello-Leitão, 1928; Elleriana Kury, 2003; Metavononoides Roewer, 1927; Bodunius Mello-Leitão, 1935; Fortalezius Roewer, 1947; Paragryne Roewer, 1912; Roquettea Mello-Leitão, 1931 – Brazil Cocholla Roewer, 1927 – Brazil, Peru 7.33 Tropical South America 489 Protus Simon, 1879 – Brazil, Ecuador Neocynorta Roewer, 1915 – Brazil, Venezuela Acrita Sorensen, 1932; Anduzeia González-Sponga, 1992; Baria GonzálezSponga, 1993; Caurimare González-Sponga, 1992; Corosalia González-Sponga, 1998; Cynortetta Roewer, 1947; Forfexia González-Sponga, 1992; Gnaricia González-Sponga, 1992; Gnatopia González-Sponga, 1992; Litoralia González-Sponga, 1992; Oligovonones di Caporiacco, 1951; Puerilia González-Sponga, 1992; Szczurehia GonzálezSponga, 1992 – Venezuela Ambatoiella Mello-Leitão, 1943; Cynortellina Roewer, 1915; Pygocynorta Roewer, 1925; Rhaucoides Roewer, 1912; Vononella Roewer, 1925; Sibambea Roewer, 1917 – Ecuador Cynortellula Roewer, 1925 – Ecuador, Peru Brachylibitia Mello-Leitão, 1941; Cumbalia Roewer, 1963; Cynortosoma Roewer, 1947; Libitiella Roewer, 1947; Megarhaucus Mello-Leitão, 1941; Messatana Strand, 1942; Neorhaucus Pic.-Cambridge, 1905; Zaraxes Roewer, 1947 – Colombia Caracarana Roewer, 1956; Chinchipea Roewer, 1952; Chirinosbius Roewer, 1952; Chusgonobius Roewer, 1952; Frizellia Mello-Leitão, 1941; Gnidiella Roewer, 1957; Moselabius Roewer, 1956; Opisthopristis Roewer, 1952; Pebasia Roewer, 1947; Praelibitia Roewer, 1956; Tobotanus Roewer, 1957; Vononissus Roewer, 1956 – Peru Eulibitia Roewer, 1912 – Ecuador, Colombia Pelechucia Roewer, 1947; Syncynorta Roewer, 1947 – Bolivia Ferkeria Roewer, 1947 – Bolivia, Peru Maniapure González-Sponga, 1992 – Bolivia, Venezuela Rhaucus Simon, 1879 – Colombia, Venezuela Fam. Cranaidae Cranainae Acanthocranaus Roewer, 1913; Carsevennia Roewer, 1913 – French Guiana Aguaytiella Goodnight et Goodnight, 1943; Aucayacuella S. Avram et H. E. M. Soares, 1983; Cenipa Goodnight et Goodnight, 1943 – Peru Alausius Roewer, 1932; Angistrisoma Roewer, 1932; Angistrius Roewer, 1932; Balzabamba Mello-Leitão, 1945; Baustomus Roewer, 1932; Bucayana Mello-Leitão, 1942; Bunicranaus Roewer, 1913; Callcosma Roewer, 1932; Cayabeus Roewer, 1932; Chetronus Roewer, 1932; Clinocippus Roewer, 1932; Diptyonius Roewer, 1932; Guayaquiliana Mello-Leitão, 1935; Licornus Roewer, 1932; Puna Roewer, 1925; Spinicranaus Roewer, 1913; Spirunius Roewer, 1932; Thaumatocranaus Roewer, 1932; Ventripila Roewer, 1917 – Ecuador 490 7 Regional Arachnogeography Allocranaus Roewer, 1915; Homocranaus Roewer, 1915; Megacranaus Roewer; Metacranaus Roewer, 1913; Panalus Goodnight et Goodnight, 1947; Paracranaus Roewer, 1913; Quindina Roewer, 1914; Sibundoxia Roewer, 1963; Tetracranaus Roewer, 1963; Ventrifurca Roewer, 1913; Ventrisudis Roewer, 1963 – Colombia Cranaostygnus Caporiacco, 1951; Stygnicranella Caporiacco, 1951; Timotesa Roewer, 1943 – Venezuela Deriacrus Roewer, 1932; Isocranaus Roewer, 1915; Microcranaus Roewer, 1913; Peripa Roewer, 1925 – Ecuador, Colombia Digalistes Roewer, 1932; Idomenta Roewer, 1932; Ladantola Roewer, 1932; Mecritta Roewer, 1932; Parkocranaus Mello-Leitão, 1949 – Brazil Neocranaus Roewer, 1913 – Colombia, Venezuela Tripilatus Roewer, 1932 – Bolivia Heterocranainae – Ecuador, Colombia Heterocranaus Roewer, 1913 – Ecuador, Colombia Prostygninae Binamballeus Roewer, 1952; Cutervolus Roewer, 1957 – Peru Chiriboga Roewer, 1959; Globibunus Roewer, 1912; Lisarea Roewer, 1943; Peladoius Roewer, 1919; Prostygnellus Roewer, 1919; Troya Roewer, 1919; Yania Roewer, 1919 – Ecuador Globitarsus Roewer, 1913; Micropachylus Roewer, 1913; Prostignidius Roewer, 1915; Prostygnus Roewer, 1913; Sclerostygnellus Roewer, 1943 – Colombia Meridanatus Roewer, 1943 – Venezuela Stygnicranainae – Colombia, Ecuador Stygnicranaus Roewer, 1913 – Colombia Tryferos Roewer, 1931 – Ecuador Fam. Escadabiidae – Brazil (six spp., endemic family) Baculigerus H. E. M. Soares, 1979; Escadabius Roewer, 1949; Jim H. E. M. Soares, 1979; Recifesius H. E. M. Soares, 1978 – Brazil Fam. Fissiphalliidae – Colombia (endemic family) Fissiphallius Martens, 1988 – Colombia Fam. Gonyleptidae Ampycella Roewer, 1931; Hexabunus Roewer, 1913; Neopachyloides Roewer, 1913; Napostygnus Roewer, 1929 – Ecuador Thaumatopachylus Roewer, 1929; Ampycus Simon, 1879; Asarcus C.L. Koch, 1839; Bogdana Mello-Leitão, 1940; Bourgnyia Mello-Leitão, 1923; Caldasius Roewer, 1930; Cnemoleptes Mello-Leitão, 1941; Opisthoplites Sørensen, 1884; Styloleptes Piza, 1943; Stylopisthos Roewer, 1930; Ampheres C.L. Koch, 1839; Arthrodes C.L. Koch, 1839; Caelopygus C.L. Koch, 1839; 7.33 Tropical South America 491 Garatiba Mello-Leitão, 1940; Metampheres Roewer, 1913; Metarthrodes Roewer, 1913; Pristocnemis C.L. Koch, 1839; Proampheres Roewer, 1913; Thereza Roewer, 1943; Adhynastes Roewer, 1930; Clarinus Roewer, 1929; Centroleptes Roewer, 1943; Collonidium Bertkau, 1880; Currala Roewer, 1927; Deltaspidium Roewer, 1927; Friburgoia Mello-Leitão, 1926; Geraecormobius Holmberg, 1887; Gonazula Roewer, 1930; Gonyleptellus Roewer, 1930; Gonyleptes Kirby, 1818; Guatubesia H. Soares; Gyndesops Roewer, 1943; Holoversia Mello-Leitão, 1940; Inhuma Piza, 1938; Liogonyloptoides Mello-Leitão, 1925; Megapachylus Roewer, 1913; Metagonyleptes Roewer, 1913; Mischonyx Bertkau, 1880; Multumbo Roewer, 1927; Neosadocus Mello-Leitão, 1926; Piassagera Roewer, 1928; Proctobunoides Mello-Leitão, 1944; Progonyleptoides Roewer, 1917; Pseudotrogulus Roewer, 1932; Sphaerobunus Roewer, 1917; Stefanasia Soares et Soares, 1988; Tupacarana Mello-Leitão, 1939; Uracantholeptes Mello-Leitão, 1926; Urodiabunus Mello-Leitão, 1935; Aerogonyleptes Roewer, 1917; Ariaeus Sørensen, 1932; Despirus Roewer, 1929; Discocyrtoides Mello-Leitão, 1923; Encheiridium Kury, 2003; Ischnotherus Kury, 1991; Longiperna Roewer, 1929; Metanitobates Roewer, 1913; Mitobates Sundevall, 1833; Mitobatula Roewer, 1931; Neoancistrotus MelloLeitão, 1929; Promitobates Roewer, 1913; Ruschia Mello-Leitão, 1940 – Brazil Parampherus Roewer, 1913 – Bolivia, Brazil Chacoikeontus Roewer, 1929; Metasarcus Roewer, 1913; Nemastygnus Roewer, 1929 – Bolivia Sibillus Roewer, 1929; Oxapampeus Roewer, 1963; Ayacucho Roewer, 1949; Cajacayba Roewer, 1957; Cajamarca Roewer, 1952; Cargaruaya Roewer, 1956; Incasarcus Kury et Maury, 1998; Palcares Roewer; Tapacochana Roewer, 1957; Taurisa Roewer, 1956; Tschaidicancha Roewer, 1957 – Peru Heteropachylinae – endemic subfamily for Brazil Aesotrinoma H. Soares, 1977; Chavesincola Soares et Soares, 1946; Heteropachylus Roewer, 1913; Mangaratiba Mello-Leitão, 1940; Melloa Roewer, 1930; Pseudopulcrolia Roewer, 1912; Thaumatoleptes Roewer, 1930; Tribunosoma Roewer, 1943 – Brazil Cobaniinae – endemic subfamily for Brazil Cobania Roewer, 1913 – Brazil Gonyassamiinae – endemic subfamily for Brazil Acutisoma Roewer, 1913; Goniosoma Perty, 1833; Goniosomoides MelloLeitão, 1932; Lyogoniosoma Mello-Leitão, 1926; Metalyogoniosoma Soares et Soares, 1946; Phalangochilus Mello-Leitão, 1938; Trichominua MelloLeitão, 1938 – Brazil 492 7 Regional Arachnogeography Pachylinae Acanthopachylopsis Soares et Soares, 1949; Acrographinotus Roewer, 1929; Allogonyleptes Roewer, 1917; Anoplogynopsis H. Soares, 1966; Antetriceras Roewer, 1949; Beckeresia H. Soares, 1970; Berlaia Mello-Leitão, 1940; Biconisoma Roewer, 1936; Bristoweia Mello-Leitão, 1924; Bunoplus Roewer, 1927; Caldanatus Roewer, 1943; Camposicola Mello-Leitão, 1924; Camposicoloides B. Soares, 1944; Capichabesia B. Soares, 1944; Carlotta Roewer, 1943; Ceratoleptes Soares et Soares, 1979; Discocyrtulus Roewer, 1927; Discocyrtus Holmberg, 1878; Eopachyloides H. Soares, 1970; Eopachylus MelloLeitão, 1931; Ergastria Mello-Leitão, 1941; Eugyndes Roewer, 1923; Eugyndopsiella H. Soares, 1972; Goodnightiella Soares et Soares, 1945; Graphinotus C.L. Koch, 1839; Guaraniticus Mello-Leitão, 1933; Gyndesoides Mello-Leitão, 1933; Gyndoides Mello-Leitão, 1927; Harpachylus Roewer, 1943; Huralvioides H. Soares, 1970; Hypophyllonomus Giltay, 1928; Iandumoema Pinto-da-Rocha, 1996; Iguassua Mello-Leitão, 1935; Iguassuoides Soares et Soares, 1954; Itatiaincola Soares et Soares, 1948; Izecksohnopilio H. Soares, 1977; Juticus Roewer, 1943; Lacronia Strand, 1942; Maromba Soares et Soares, 1954; Metadiscocyrtus Roewer, 1929; Metagraphinotus Mello-Leitão, 1927; Metagyndoides Mello-Leitão, 1931; Metalycomedes Mello-Leitão, 1927; Metapachyloides Roewer, 1917; Meteusarcoides Mello-Leitão, 1922; Meteusarcus Roewer, 1913; Neopachylus Roewer, 1913; Pachylibunus Roewer, 1913; Pachylusius Mello-Leitão, 1949; Paradiscocyrtus Mello-Leitão, 1927; Paraluederwaldtia Mello-Leitão, 1927; Paranaleptes Soares et Soares, 1947; Paraprosontes Soares et Soares, 1947; Pareusarcus Roewer, 1929; Passosa Roewer, 1927; Pherania Strand, 1942; Planiphalangodus Roewer, 1929; Pseudoacrographinotus H. Soares, 1966; Pseudogyndesoides B. Soares, 1944; Rhioxyna Soares et Bauab-Vianna, 1970; Roeweria Mello-Leitão, 1943; Schubartesia B. Soares, 1944; Singram Mello-Leitão, 1937; Triglochinura MelloLeitão, 1924; Ubatubesia B. Soares, 1945; Uropachylus Mello-Leitão, 1922; Victoriaincola Soares et Soares, 1946; Yraguara Mello-Leitão, 1937 – Brazil Paraphalangodus Roewer, 1915 – Colombia Bullaepus Roewer, 1930; Gerdesius Roewer, 1952; Huadquina Roewer, 1930; Huasampillia Roewer, 1913; Hyperpachylus Roewer, 1957; Junicus Goodnight et Goodnight, 1947; Palcapachylus Roewer, 1952; Pichitus Roewer, 1959; Pirunipygus Roewer, 1936; Platygyndes Roewer, 1943; Polyacanthoprocta Mello-Leitão, 1927; Punagraphinotus Soares et Bauab-Vianna, 1972; Tarmapachylus Roewer, 1956; Tingomaria Mello-Leitão, 1948 – Peru Punrunata Roewer, 1952 – Brazil, Peru Acrographinotus Roewer, 1929 – Peru, Bolivia Unduavius Roewer, 1929 – Bolivia Ceropachylinus Mello-Leitão, 1943; Ceropachylus Mello-Leitão, 1942; Ibarra Roewer, 1925 – Ecuador Pachylospeleinae – endemic subfamily for Brazil Pachylospeleus Šilhavý, 1974 – Brazil 7.33 Tropical South America 493 Progonyleptoidellinae Cadeadoius Mello-Leitão, 1936; Detigalus Roewer, 1931; Gonyleptoides Roewer, 1913; Heliella B. Soares, 1945; Iguapeia Mello-Leitão, 1935; Iporangia Mello-Leitão, 1935; Leptocnema C.L. Koch, 1839; Mitopernoides B. Soares, 1945; Moreiranula Roewer, 1930; Progonyleptoidellus Piza, 1940 – Brazil Sodreaninae – endemic subfamily for Brazil Gertia Soares et Soares, 1946; Sodreana Mello-Leitão, 1922; Stygnobates Mello-Leitão, 1927; Zortalia Mello-Leitão, 1942 – Brazil Tricommatinae Arminda Roewer, 1949; Arruda Mello-Leitão, 1940; Bebedoura Roewer, 1949; Bissulla Roewer, 1929; Bresslauius Mello-Leitão, 1935; Bunostigma MelloLeitão, 1935; Camarana Mello-Leitão, 1935; Cryptogeobius Mello-Leitão, 1942; Heteromeloleptes Mello-Leitão, 1931; Lanesoares Roewer, 1949; Liops Mello-Leitão, 1940; Lyssanvira Mello-Leitão, 1935; Pararezendesius H. Soares, 1972; Poecilosophus Mello-Leitão, 1948; Pseudopachylus Roewer, 1912; Pseudophalangodes Roewer, 1912; Rezendezius H. Soares, 1945; Segundolus Roewer, 1949; Spinopilar Mello-Leitão, 1940; Tachusina Strand, 1942; Tibangara Mello-Leitão, 1940; Tricommatus Roewer, 1912; Zalanodicus MelloLeitão, 1936 – Brazil Otuquisa Roewer, 1927 – Bolivia Paratricommatus Piza, 1943 – Brazil, Paraguay Fam. Guasiniidae – Venezuela, Brazil Guaiquinimia González-Sponga, 1997 – Venezuela Guasinia González-Sponga, 1997 – Venezuela, Brazil Fam. Icaleptidae – Colombia, Ecuador Icaleptes Kury et Pérez, 2002 – Colombia Zalmopsylla Kury et Pérez, 2002 – Ecuador Fam. Manaosbiidae Azulamus Roewer, 1957 – Peru Belemnodes Strand, 1942; Belemulus Roewer, 1932; Manaosbia Roewer, 1943; Paramicrocranaus H. Soares, 1970; Syncranaus Roewer, 1913 – Brazil Camelianus Roewer, 1912; Gonogotus Roewer, 1943; Cucutacola Mello-Leitão, 1940; Semostrus Roewer, 1943 – Colombia Clavicranaus Roewer, 1915; Saramacia Roewer, 1913 – Suriname, Brazil Dibunostra Roewer, 1943; Poecilocranaus Roewer, 1943 – Venezuela Mazarunius Roewer, 1943 – Guyana Meridia Roewer, 1913 – Venezuela, Suriname Pentacranaus Roewer, 1963; Tegyra Sørensen, 1932 – Peru Rhopalocranellus Roewer, 1925 – Ecuador unknown genus calcar (Roewer, 1943) – Venezuela 494 7 Regional Arachnogeography unknown genus albituberculatus (Roewer, 1943) – Guyana unknown genus strinatii (Šilhavý, 1979) – Venezuela Fam. Kimulidae (= Minuidae) Acanthominua Sørensen, 1932; Euminua Sørensen, 1932; Euminuoides MelloLeitão, 1935; Fudeci González-Sponga, 1997; Microminua Sørensen, 1932; Minua Sørensen, 1932 – Venezuela Tegipiolus Roewer, 1949 – Brazil Pseudominua Mello-Leitão, 1935 – Venezuela, Peru Fam. Samoidae Cornigera González-Sponga, 1987 – Venezuela Fam. Stygnidae Heterostygninae Eutimesius Roewer, 1913 – Colombia, Venezuela, Ecuador Innoxius Pinto-da-Rocha, 1997; Minax Pinto-da-Rocha, 1997; Stenostygnellus Roewer, 1913; Yapacana Pinto-da-Rocha, 1997 – Venezuela Stygnidius Simon, 1879 – Brazil, French Guiana Timesius Simon, 1879 – Colombia Nomoclastinae – Colombia (endemic subfamily) Nomoclastes Sørensen, 1932 – Colombia Stygninae Actinostygnoides Goodnight et Goodnight; Auranus Mello-Leitão, 1941 Iguarassua Roewer, 1943; Kaapora Pinto-da-Rocha, 1997; Paraphareus Goodnight et Goodnight, 1943; Pickeliana Mello-Leitão, 1932; Planophareus Goodnight et Goodnight, 1943; Sickesia H.E.M. Soares, 1979; Stenophareus Goodnight et Goodnight, 1943; Verrucastygnus Pinto-da-Rocha, 1997 – Brazil Metaphareus Roewer, 1912; Phareus Simon, 1879 – Colombia, Venezuela Niceforoiellus Mello-Leitão, 1941 – Colombia Ortonia Wood, 1869 – Ecuador Protimesius Roewer, 1913 – Peru, Brazil, Ecuador, French Guiana, Suriname Stenostygnoides Roewer, 1913 – Suriname Stygnus Perty, 1833 – Venezuela, Brazil, Peru, Colombia, Suriname Subfam. uncertain – Brazil Gaibulus Roewer, 1943 – Brazil Fam. Zalmoxidae Absonus González-Sponga, 1987; Araguita González-Sponga, 1987; Avilaia González-Sponga, 1998; Azulitaia González-Sponga, 1987; Buruquelia González-Sponga, 1999; Cea González-Sponga, 1987; Chamaia GonzálezSponga, 1987; Cubiria González-Sponga, 1987; Curimaguanus GonzálezSponga, in Kury 2003; Granulaia González-Sponga, 1997; Guagonia 7.33 Tropical South America 495 González-Sponga, 1987; Haitonia González-Sponga, 1987; Jajinia GonzálezSponga, 1987; Lara M. A. González-Sponga, 1987; Micro González-Sponga, 1987; Orituco González-Sponga, 1987; Panaquire M. A. González-Sponga, 1987; Paraminuella Caporiacco, 1951; Paramo González-Sponga, 1987; Phalangodinella Caporiacco, 1951; Retropedis González-Sponga, 1987; Soledadiella González-Sponga, 1987; Sphoeroforma González-Sponga, 1987; Spiniella González-Sponga, 1987; Taguaza González-Sponga, 1998; Tiara González-Sponga, 1987; Traiania H. E. M. Soares et S. Avram, 1981; Unare González-Sponga, 1987; Unicornia González-Sponga, 1987; Urachiche González-Sponga, 1987; Yacambuia González-Sponga, 1987; Carayaca González-Sponga, 1998; Chirimena González-Sponga, 1999; Guayania González-Sponga, 1999; Junquito González-Sponga, 1999; Niquitaia González-Sponga, 1999; Ovalia González-Sponga, 1999; Pijiguaia González-Sponga, 1998; Pilosa González-Sponga, 1999 – Venezuela Cochirapha Roewer, 1949; Exlineia Mello-Leitão, 1942; Galanomma Juberthie, 1970 – Ecuador Garanhunsa Roewer, 1949; Tegipiolus Roewer, 1949 – Brazil Viacha Roewer, 1949 – Bolivia Weyrauchiana Roewer, 1952; Sivianus Roewer, 1949 – Peru Pirassunungoleptes H. Soares, 1966 – Brazil, Bolivia Amblypygi From South America are recorded ca. 60 spp. of Amblypygi, belonging to the genera Charinus Simon, Trichodamon Mello-Leitão, Paraphrynus Moreno, Heterophrynus Pocock, and Phrynus Lamarck and the families Charinidae, Phrynichidae, and Phrynidae (Pocock 1894, Baptista and Giupponi 2002ª, 2002b, 2003, Caporiacco 1947, Giupponi and Kury 2002, 2013, Gonzalez-Sponga 1998, Goodnight and Goodnight 1946, Harvey 2013c, Jocqué and Giupponi 2012, MelloLeitão 1931, Pinto-da-Rocha et al. 2002, Miranda and Giupponi, 2002, Ravelo 1975a, 1975b, Quintero 1981, 1983, Vasconcelos et al. 2013, Weygoldt 1972a, 1972b). The genus Charinus is the most widespread, including on islands like Galapagos. From continental South America (Guyana, Brazil, Suriname, Venezuela, Peru) are known more than 20 species, including 12 from Brazil. Trichodamon Mello-Leitão (two spp. from Brazil) is the only genus in the family Phrynichidae living in the Western Hemisphere (the other six genera are represented in Africa and the outlying islands). This is the only one endemic amblypygid genus in South America. Paraphrynus Moreno (18 spp. from Colombia to the Southern USA) is typical mostly for Central America. In South America has been registered one sp. (Colombia). Phrynus Lamarck (five spp. in continental South America – Colombia, Ecuador, Guyana, Venezuela, Brazil) is widespread from Brazil to the southern states of USA (Texas, Florida), mostly in Mexico, Central America, and the Antilles. Recently an aberrant species was described from a cave on the Flores Island in 496 7 Regional Arachnogeography Indonesia (Phrynus exsul Harvey, 2002), the only representative of the family Phrynidae in the Old World. Heterophrynus Pocock, 1894 (the only genus in the subfamily Heterophryninae) – [Mexico] Brazil, Peru, Colombia, French Guiana, Guyana, Venezuela, Ecuador, Suriname (14 spp.) Map 7.25 Amblypygi in South America Charinus – ■ Heterophrynus – ◄ Trichodamon – ● Paraphrynus – ▼ Phrynus – ▲ 7.33 Tropical South America 497 Thelyphonida (Uropygi) The whip scorpions in tropical South America belong to the subfamily Hypoctoninae and to the genera Thelyphonellus Pocock, 1894 (Brazil, Guyana, Suriname, Venezuela, two spp.) and Mastigoproctus Pocock, 1894 (= Amauromastigon Mello-Leitão, 1931) – widespread and reaches the Antilles, Central America, Mexico, and the Southern USA (Haupt 2009b, Mello-Leitão 1931, Pocock 1894, Rowland 2002, Villarreal Manzanilla and Giupponi 2009). In the subfamily Hypoctoninae, only the genus Thelyphonellus is South American; Ravilops lives in the Dominican Republic, Valeriophonus in Costa Rica, and the other three genera in West Africa (Etienneus), India, and Sri Lanka (Labochirus) or Southeast Asia (Hypoctonus). In the subfamily Uroproctinae, two monotypic genera live in Asia; all the other species belong to Mastigoproctus and are widespread. One enigmatic species was described from Hong Kong (Mastigoproctus transoceanicus Lazell, 2000) but is clearly introduced. Thelyphonellus, Valeriophonus, Ravilops, and Mastigoproctus could be termed as Neotropical. 498 7 Regional Arachnogeography Map 7.26 Thelyphonida in South America, Central America, and the Caribbean Thelyphonellus – ■ Mastigoproctus – ▼ Ravilops – ● Mayacentrum – Valeriophonus – ▲ Mimoscorpius– ◄ Schizomida Armas (2010) made a review of Schizomida known from South America. With some supplements, there are species of the genera Adisomus, Calima, Surazomus, Rowlandius, Stenochrus, Tayos, Hansenochrus, Stenoschizomus, and Wayuuzomus in seven countries: Colombia, Venezuela, Ecuador, Peru, Brazil, Guiana, and Suriname. The southernmost finding of Schizomida on the continent is Stenochrus portoricensis Chamberlin in Rio de Janeiro (probably introduced). Otherwise the order does not go more southern than the Amazonia – Armas et al. 7.33 Tropical South America 499 (2009), Bonaldo and Pinto-da-Rocha (2007), Giupponi et al. (2016), Kraus (1957), Kraus and Beck (1967), Pinto-da-Rocha (1996, 2007), Reddell and Cokendolpher (1995), Rowland and Reddell (1979), Villarreal Manzanilla et al. (2008), MorenoGonzález and Manzanilla (2012), Santos and Pinto-da-Rocha (2009), Santos et al. (2013), Moreno-González et al. (2014) (Table 7.9). Endemic genera of Schizomida for South America are: Fam. Hubbardiidae Adisomus Cokendolpher et Reddell, 2000 – Brazil (one sp.) Calima Moreno-González et Manzanilla, 2012 – Colombia (two spp.) Colombiazomus Armas et Delgado-Santa, 2012 – Colombia (one sp.) Stenoschizomus Gonzalez-Sponga, 1997 – Venezuela (one sp.) Tayos Reddell et Cokendolpher, 1995 – Ecuador (one sp.) Wayuuzomus Armas et Colmenares, 2006 – Venezuela (one sp.) Table 7.9 Distribution of Schizomida in South America Country Taxa Fam. Hubbardiidae Adisomus Cok. et Reddell Calima Mor.-Gon. et Manz. Colombiazomus Armas et Del. Hansenochrus Red.et Cok. Piaroa Manz. et al. Rowlandius Red. et Cok. Stenochrus Chamberlin Stenoschizomus Gonz.-Sponga Surazomus Red. et Cok. Tayos Red. et Cokendolpher Wayuuzomus Armas et Cok. Brazil Colombia Venezuela Guiana Suriname Bolivia Ecuador Peru 11 5 8 2 3 1 3 1 + + + + + + + + 1 – – – – – – – – 2 – – – – – – – 2 – – – – – – – – 1 2 – – – – – 1 1 – – – – – 2 – – – – – – 1 1 1 – – – – – – – 1 – – – – – 7 3 – – – 1 1 1 – – – – – – 1 – – – 1 – – – – – 500 7 Regional Arachnogeography Map 7.27 Distribution of Schizomida in South America ■ – Colombiazomus X – Piaroa ● – Stenochrus ▼ – Surazomus ▲ – Wayuuzomus ◊ – Hansenochrus □ – Rowlandius ► – Adisomus ▽– Stenoschizomus ◄ – Calima – Tayos Araneae From tropical South America (without the southernmost end) are recorded spiders of at least 70 families. In the south (Patagonia, Chile) occur 7 more families, or spiders of 77 families are known south of Panama (72 % of all spider families). 7.33 Tropical South America 501 In the entire territory of Chile are recorded 55 families (Aguilera and Casanueva 2005). No family is endemic for the South American continent. This low level of endemism is remarkable for the Neotropic (Huber 2000). On the small islands Trinidad and Tobago have been recorded 51 families of spiders (plus 12 presumed by Sewlal and Cutler 2003). Endemic suprageneric taxa of Laniatores for the Neotropical Kingdom are: Opiliones, Laniatores Fam. Stygnidae – tropical South America, Lesser Antilles Subfam. Nomoclastinae – Colombia Fam. Agoristenidae – South America and the Caribbean Fam. Cosmetidae – South and Central America and Mexico Fam. Cranaidae – South and Central America and the Caribbean Fam. Escadabiidae – Brazil Fam. Fissiphaliidae – Colombia, Brazil Fam. Gonyleptidae – from Patagonia to Guatemala Fam. Guasiniidae – Brazil, Venezuela Fam. Icaleptidae – Colombia, Ecuador Fam. Manaosbiidae – tropical South and Central America, West Indies Fam. Kimulidae (Minuidae) – tropical South America, West Indies The wealth of Opiliones and Laniatores in the Neotropics is amazing and much bigger than in any other kingdom. We have also to take into account that large areas in South America are still almost unexplored (Tourinho and Pérez González 2006). It is also to notice that there is no such situation in any other order in South America. Opilioacarida From tropical South America was described Neocarus ojastii Lehtinen, 1980 (Venezuela). N. platensis (Silvestri) is recorded also from Brazil. Acariformes Fam. Erythraeidae: Iguatonia Haitlinger, 2004 – Brazil (larval) Parasitiformes (Mesostigmata and Ixodida) The very peculiar family of Spelaeorhynchidae is confined to the New World (six spp., parasites on bats of the families Phyllostomatidae and Mormoopidae). These mites have been found so far in Belize, Brazil, Colombia, Cuba, Dominican Republic, Guatemala, Jamaica, Panama, Peru, Puerto Rico, and Venezuela (Beron 2014). Holothyrida From South America have been recorded only two species of Holothyrida, belonging to two endemic genera of Neothyridae – one from Brazilian Amazonas (Diplothyrus schubarti Lehtinen, 1999) and the other from the region of Iquitos (Neothyrus ana Lehtinen, 1999), again from Amazonia, but in Peru. Neothyrus sp. is known from Venezuela (Beron 2014). 502 7.34 7.34.1 7 Regional Arachnogeography Galapagos Geography, General Zoogeography, and Paleogeography The Galapagos Islands are an archipelago in the Pacific Ocean, part of Ecuador, about 966 km far from the continent. The total area of the archipelago is 7845 km2, the 13 main islands being Isabela (5827 km2), Fernandina (642 km2), Santa Cruz (986 km2), Floreana (173 km2), Santiago (585 km2), Española (Hood) (60 km2), San Cristóbal (Chatham) (558 km2), and some smaller islands. There are also six smaller islands and 107 rocks. The highest point is Volcan Wolf in Isabela (1707 m). The archipelago is situated on Nazca Plate which is moving with a speed of 6.4 cm/year and is being subducted under the South American Plate. Underneath is formed the so-called Galapagos hot spot, where the core is melting and forming volcanoes. The first islands are formed at least 8 million years ago and had no connection with the mainland. At present the oldest (and most southern) island is Fernandina, 3.5 million years ago. The colonization of the islands could have commenced earlier on lands, having been dry land 14 Ìa, and now are under water. The line divergence of the terrestrial and the sea iguanas occurred 10 Ìa ago, so it happened on the older, now submerged islands (Chubb 1933, Colantoni et al. 1979, Kuschel 1961, Lanteri 2001, Parent et al. 2008, Simkin 1984, Snell et al. 1995). The volcanic oceanic archipelago has 749 species of higher plants, including 216 endemic. There are two species of bats, and it seems a good question why more bats from the rich South American fauna did not fly over that distance of 966 km. On the archipelago have radiated also 10 spp. of rice rats (Oryzomys, Nesoryzomys, Megaoryzomys), but now only three remained. There are no amphibians, but there are 37 species of reptiles (three snakes, geckos, tortoises, terrestrial and marine iguanas). From (only) 30 species of birds, 25 are endemic, including the 14 spp. of the famous Darwin’s finches (Geospizinae). About 1500 species of 1000 genera of native insects form the entomofauna of the islands. Among the other invertebrate, very impressive is the number of oribatid mites (202 spp., according to Schatz 1989). Morrone (1999, 2001) includes the Galapagos as a province in the Caribbean Subregion of the Neotropical Region. The interesting cave fauna, found on Galapagos (Peck and Kukalova-Peck 1986, Peck and Finstone 1993), contributes to the formulating of new ideas and bioclimatic model of the troglomorphosis. 7.34 Galapagos 7.34.2 503 Arachnogeography On the archipelago have been recorded representatives of Solifugae, Scorpiones, Pseudoscorpiones, Opiliones (Laniatores), Amblypygi, Schizomida, Araneae, Acariformes, and Parasitiformes (Baert 2011, 2014, Baert et al. 1995, Coolidge 1909, Roth and Craig 1970). After Baert (version 2014), on Galapagos are recorded 434 species of Arachnida. Schatz (1998) analyzed in details the possible ways of dispersal, colonization, and speciation of the Oribatida on these islands. The main way is the hydrochorous (directly and on/in floating logs or plant debris). Other ways are zoochory (birds) and anthropochory. Some of the other Arachnida could have followed the same pattern of dispersal; for some others different explanation is required. Peck and Finstone (1993) summarized the amazing cave fauna of Galapagos and proposed an explanation of its origin by parapatric speciation. Solifugae Endemic genus and species is Neocleobis solitarius Roewer, 1934 (Ammotrechidae) (Baert et al. 1995, Banks 1902, Roewer 1934). Scorpiones From Galapagos have been registered the species Centruroides exsul Meise, 1933, and Hadruroides galapagoensis Maury, 1974 (Baert et al. 1995, Banks 1902, Kinzelbach 1973, Maury 1974). Pseudoscorpiones From Galapagos are known 25 spp. of 9 genera (Tyrannochthonius, Lechytia, Apolpium, Stenolpium, Serianus, Galapagodinus, Ideobisium, Parachernes, Rhopalochernes) and 10 families (Chthoniidae, Lechytiidae, Syarinidae, Olpiidae, Garypinidae, Garypidae, Cheiridiidae, Atemnidae, Chernetidae, Withiidae) (Baert et al. 1995, Beier 1940, 1976d, 1978, Mahnert 2014). Endemic genus for Galapagos: Fam. Garypinidae: Galapagodinus Beier, 1978 The species are almost all endemic. Opiliones Cyphophthalmi, Eupnoi, and Dyspnoi – not recorded Laniatores (Baert et al. 1995, Juberthie 1970c) Endemic genus and species is Galanomma microphthalma Juberthie, 1970 (Gonyleptidae). Amblypygi From Galapagos is described only the endemic species Charinus insularis Banks, 1902 (Charinidae). The genus Charinus is widespread. Schizomida One widespread sp., Stenochrus portoricensis (Chamberlin, 1922), known from Central America and West Indies (Baert et al. 1995) 504 7 Regional Arachnogeography Araneae According to Baert (2011, 2014), on Galapagos so far have been recorded 149 spp. of spiders. “Of the 124 identified species, 64 (almost 50%) are known only from the archipelago and hence may be endemic, 35 species have a New World distribution, 16 are cosmopolitan species, six pantropical, and three are cosmotropical species” (Baert, loc. cit.; Baert 1987, 1990, 2013, Baert and Maelfait 1986, 1997, Baert et al. 1989, 2008a, 2008b, Banks 1902, Levi 2009, Peck and Shear 1987a, b). A blind Lygromma (Gnaphosidae) and a new eyeless stridulating Theridion have been described from the lava tubes. Endemic genera: Fam. Mysmenidae: Calomyspoena Baert et Maelfait, 1983 Fam. Pholcidae: Galapa Huber, 2000 Fam. Araneidae: Galaporella Levi, 2009 (one sp.) Fam. Salticidae: Darwinneon Cutler, 1971 Acariformes Sarcoptiformes Oribatida According to Schatz (1998), on the archipelago have been registered 202 spp. of oribatid mites belonging to 64 families (during the 10-year study, 81 new species have been described). The species richness of an island depends on the altitude and number of available habitats rather than the area of the island. Many oribatid species on the Galápagos Islands have a wide biogeographical distribution. The majority originate from the Central and South American mainland, but several Pacific and even Holarctic elements were also found. In comparison with the species composition of the adjacent mainland, the oribatid mite fauna of the Galápagos Islands can be regarded as disharmonic. Sea surface transportation has been proved at least between the islands, which also applies to oribatid species living at higher elevations. Long distance dispersal can be assumed to be mainly hydrochorous. Approximately 40% of all oribatid species from the Galápagos Islands have a known distribution restricted to the islands and can be considered to be endemic. (Schatz 1998) Endemic genus and species is Galapagacarus schatzi Balogh, 1985. Parasitiformes Ixodida Fam. Ixodidae – several species (endemic) are described from lizards and tortoises. Ixodes galapagoensis Clifford et Hoogstraal, 1980; Amblyomma boulengeri Hirst et Hirst, 1910; A. darwini Hirst et Hirst, 1910; A. usingeri Keirans, Hoogstraal, et Clifford, 1973; A. williamsi Banks, 1924 7.35 Juan Fernandez Islands 7.35 7.35.1 505 Juan Fernandez Islands Geography, General Zoogeography, and Paleogeography The volcanic archipelago Juan Fernandez, 674 km west of South America, has an area of 99.6 km2, of which 47.9 km2 is of Robinson Crusoe Island, or Isla Más a Tierra (together with Santa Clara) and 49.5 km2 of Alejandro Selkirk Island (Isla Más Afuera). The two islands are 181 km far from each other. The altitudes are El Yunque (916 m on Robinson Crusoe) and Cerro de Los Inocentes (1319 m on Alejandro Selkirk). The islot Santa Clara (2.2 km2) has 350 m height and is the oldest island of the archipelago (5.8 million years old), next is Robinson Crusoe (3.8– 4.2 million years old) and Alexander Selkirk (1.0–2.4 million years old) (Kuschel 1961, Scottsberg 1954). The climate of the islands is Mediterranean. According to UNESCO World Netwrok of Biosphere Reserves, since 1977 these islands have been considered of scientific importance as an ecoregion, because of the endemic species of flora and fauna There are 126 endemic species of plants (62%), with 12 endemic genera and 1 endemic family, Lactoridaceae. According to Morrone (2000), biogeographically Juan Fernandez Islands are a province within the Subantarctic Subregion of the Andean region. 7.35.2 Arachnogeography From the islands are known representatives (with low-level endemism) of the orders Pseudoscorpiones (15 spp., almost all endemic, with 2 endemic genera), Opiliones, and Araneae (2 endemic genera) and some mites. Pseudoscorpiones The pseudoscorpions recorded on Juan Fernandez Islands belong to the families Lechytiidae, Geogarypidae, Cheiridiidae, Chernetidae, and Withiidae; the genera Lechytia, Geogarypus, Neocheiridium, Parazaona, Asterochernes, Chelanops, Pseudopilanus, Asterochernes, Selachochernes, and Protowithius; and 15 species, 14 of which endemic and 1 known from Argentina (Beier 1955, 1957b, Mahnert 1993, 1997, 2011). Endemic genera: Fam. Chernetidae: Selachochernes Mahnert, 2011 (one sp.) Fam. Withiidae: Protowithius Beier, 1955 (two spp.). Opiliones Eupnoi Fam. Acropsopilionidae 506 7 Regional Arachnogeography Acropsopilio chilensis Silvestri, 1904 – the first Opilion discovered on the archipelago, known also from Chile, including Tierra del Fuego (Pérez-González et al. 2014) Araneae Bäckstrom from the expedition of Skottsberg in 1916–1917 has collected spiders, which were studied by Berland (1924 and 1935). The list of Berland, 1924b (including the spiders recorded by F.O.P. –Cambridge and E. Simon), contains 24 spp., among them are 9 new spp. and the new genus Selkirkiella (Theridiidae). Endemic genera are: Fam. Linyphiidae Schistogyna Millidge, 1991 (one sp.) Malkinola Miller, 2007 Juanfernandezia Koçak et Kemal, 2008 7.36 7.36.1 North America, North of Rio Grande (incl. Greenland) Geography, General Zoogeography, and Paleogeography North America occupies the northern portion of the landmass generally referred to as the New World. North America’s only land connection to South America is at the Isthmus of Panama. The continent is delimited on the southeast by most geographers at the Darién watershed along the Colombia-Panama border, placing all of Panama within North America. The Caribbean Islands, or West Indies, are considered part of North America. Before the Central American isthmus was raised, the region had been underwater. The islands of the West Indies delineate a submerged former land bridge, which had connected North America and South America via what are now Florida and Venezuela (Kaiser et al.1972 Martin and Harrell 1957, Rogers et al. 1999, Williams et al. 2000). Greenland, a self-governing Danish island, and the world’s largest, is on the same tectonic plate (the North American Plate) and is part of North America geographically. In a geologic sense, Bermuda is not part of the Americas, but an oceanic island which was formed on the fissure of the Mid-Atlantic Ridge over 100 million years ago. The vast majority of North America is on the North American Plate. Parts of western Mexico, including Baja California, and of California, including the cities of San Diego, Los Angeles and part of San Francisco, lie on the eastern edge of the Pacific Plate, with the two plates meeting along the San Andreas fault. The southernmost portion of the continent and much of the West Indies lie on the Caribbean Plate. 7.36 North America, North of Rio Grande (incl. Greenland) 507 The western mountains are split in the middle into the main range of the Rockies and the coast ranges in California, Oregon, Washington, and British Columbia, with the Great Basin – a lower area containing smaller ranges and low-lying deserts – in between. The highest peak is Denali in Alaska (6168 m). North America covers an area of about 24,709,000 square kilometers, about 4.8% of the planet’s surface or about 16.5% of its land area. Laurentia is an ancient craton which forms the geologic core of North America; it formed between 1.5 to 1.0 billion years ago during the Proterozoic eon. From the Late Paleozoic to Early Mesozoic eras, North America was joined with the other modern-day continents as part of the supercontinent Pangaea. One of the results of the formation of Pangaea was the Appalachian Mountains, which formed some 480 million years ago, making it among the oldest mountain ranges in the world. When Pangaea began to rift around 200 million years ago, North America became part of Laurasia, before it separated from Eurasia as its own continent during the midCretaceous period. The Rockies and other western mountain ranges began forming around this time from a period of mountain building called the Laramide orogeny, between 80 and 55 million years ago. The formation of the Isthmus of Panama connected the continent to South America about three million years ago. 7.36.2 Arachnogeography In North America, north of Mexico, are found 13 orders of Arachnida, but the “southern” orders (Amblypygi, Ricinulei, Schizomida, Uropygi, Opilioacarida, and to big extent Solifugae and Scorpiones) are confined mostly to Texas, Arizona, California, Florida, Louisiana, and New Mexico, some of them being on the edge of their areals (Dice 1943, Harvey, online, Rowland and Reddell 1976, Sørensen 1898). In Canada the southern groups are either lacking (Ricinulei, Schizomida) or present with one cold resisting species (Scorpiones). One family of spiders (Trogloraptoridae) is endemic to the caves of the USA. In Greenland have been recorded 64 spp. of spiders, 1 opilion (Mitopus morio), and 127 spp. of Acari (Jackson 1930; Sørensen 1898, Jensen and Christensen, Eds. 2003). A remark by Shelley and Golovatch (2011) concerning Diplopoda is to be noticed: “All South American taxa that extend northward to North America/US through Central America also inhabit the Antillean Arc with the exception of Epinannolenidea, which is primarily Antillean and only ranges to central Costa Rica. Conversely, no North American taxon spreads southward completely to South America, and those that go even part way do so via Central America, as the Antillean paleogeographic origin excludes them as a northsouth pathway. No primarily North American taxon inhabits even part of the Antilles”. 508 7 Regional Arachnogeography Palpigradi According to Condé (1996) and suppl., in the USA (Texas, Louisiana) are recorded Eukoenenia florenciae (Rucker, 1903), E. hanseni (Silvestri, 1913), Prokoenenia californica Silvestri, 1913 (California), P. wheeleri (Rucker, 1901) (Texas), and many unidentified (Rucker 1901, 1903, Silvestri 1813, Wheeler 1900). In North America the order is known from the area up to 44oN (Condé 1996). Ricinulei The only recent species of Ricinulei from the USA (Pseudocellus dorotheae) has been described from Edinburg, Texas, Hidalgo County, and on the border with Mexico (Gertsch and Mulaik 1939). In this country are registered also nine fossil species of Ricinulei (Curculioididae and Poliocheridae). Solifugae According to the checklist of Brookhart and Brookhart (2006), in continental North America have been recorded 196 species of Solifugae, from Eremobatidae (genera Eremothera Muma, Eremorhax Roewer, Eremobates Banks, Eremocosta Roewer, Horribates Muma, Chanbria Muma, Eremochelis Roewer, Hemerotrecha Banks, 174 spp.) and Ammotrechidae (Branchia Muma, Ammotrechella Roewer, Ammotrechula Roewer, Ammotrecha Banks, 22 spp.). From these 196 spp., 77 are known from Mexico; for the area north of Rio Grande remain ca. 119 spp. (Brookhart 1972, Muma 1951, 1962, 1970, 1974, 1976, 1986). The deserts Chihuahuan and Sonoran contain the majority of solifugid species. From the southern provinces of Canada are known two species: Eremobates docolora Brookhart et Muma, 1981, and Hemerotrecha denticulata Muma, 1951. They mark the northern limit of Solifugae in the Western Hemisphere. Scorpiones In North America, north of Mexico (30 states of the USA), are distributed 111 spp. of scorpions of 20 genera and 6 families: Buthidae, Caraboctonidae, Chactidae, Scorpionidae (incl. Diplocentridae), Superstitionidae, and Vaejovidae (see Table 8.1). Paruroctonus boreus (Vaejovidae) is the only species that occurs in Canada (southern Saskatchewan, Alberta, and British Columbia) (Ewing 1928, Gertsch and Soleglad 1966, Stahnke 1956, Soleglad and Fet 2005, 2006, 2008, 2010, Soleglad et al. 2014). The states richest in scorpions are California (51 spp.), Arizona (41 spp.), New Mexico (20 spp.), Texas (20 spp.), Nevada (16 spp.), and Utah (12 spp.). Pseudoscorpiones Many data have been published by Beier (1931, 1932), Chamberlin (1925, 1929, 1930, 1931, 1949), Hoff (1946, 1956), and others. Longtime ago Hoff (1958) made a list of the pseudoscorpions of North America, north of Mexico. By this time the list contained 87 genera of 18 modern families. Meanwhile, many new data were published (to mention only a few): Benedict and Malcolm (1978), Buddle (2005), Ćurčić (1978, 1981, 1982, 1984), Harvey (online), and Muchmore (1967, 1974). Rowland and Reddell (1976) made a list of the (then) 30 species in Texas. Now (2017) we may assume that in North America, north of Rio Grande, are recorded about 400 species of pseudoscorpions, belonging to 105 genera and 19 families (Harvey, 2013f). From these families 13 contain 1–2 genera and six, 1–2 species each. 7.36 North America, North of Rio Grande (incl. Greenland) 509 Genera living only in this (quite large) area (Nearctic) are: Fam. Chthoniidae Neochthonius Chamberlin, 1929 – California (four spp.) Fam. Tridenchthoniidae Verrucaditha J.C. Chamberlin, 1929 – USA (one sp.) Fam. Pseudogarypidae Pseudogarypus Ellingsen, 1909 – Canada, USA (six spp.) Fam. Neobisiidae Alabamocreagris Ćurčić, 1984, Alabama (two spp.); Americocreagris Ćurčić, 1982, Oregon, Washington (one sp.); Australinocreagris Ćurčić, 1984, California (one sp.); Cryptocreagris Ćurčić, 1982, New Mexico, California (two spp.); Fissilicreagris Ćurčić, 1984, California (four spp.); Globocreagris Ćurčić, 1984, California (two spp.); Lissocreagris Ćurčić, 1981, USA (seven spp.); Minicreagris Ćurčić, 1981, USA (one sp.); Novobisium Muchmore, 1967, USA (three spp.); Saetigerocreagris Ćurčić, 1984, USA (two spp.); Tartarocreagris Ćurčić, 1984, Texas, Arkansas (six spp.); Trisetobisium Ćurčić, 1982, USA (one sp.); Tuberocreagris Ćurčić, 1978 – USA (two spp.) Fam. Syarinidae Aglaochitra J.C. Chamberlin, 1952 – USA (California) (one sp.) Fam. Cheliferidae Aspurochelifer Benedict et Malcolm, 1979 (one sp.); Florichelifer Hoff, 1964 (Florida) (one sp.); Haplochelifer J.C. Chamberlin, 1932 (one sp.); Idiochelifer J.C. Chamberlin, 1932 (one sp.); Juxtachelifer Hoff, 1956 (New Mexico)(one sp.); Levichelifer Hoff, 1946, Mexico, USA (two spp.); Pachychelifer Beier, 1962, Georgia (one sp.); Paisochelifer Hoff, 1946, USA (one sp.); Phorochelifer Hoff, 1956, USA (one sp.); Pugnochelifer Hoff, 1964, USA (one sp.); Xenochelifer J.C. Chamberlin, 1949, California (one sp.) Fam. Chernetidae Acuminochernes Hoff, 1949 (two spp.); Chrysochernes Hoff, 1956 (New Mexico) (one sp.); Illinichernes Hoff, 1949 (one sp.); Mirochernes Beier, 1930 (one sp.); Mucrochernes Muchmore, 1973 (one sp.) In Alaska are registered three species (Neobisiidae, Globocreagris, Halobisium; Syarinidae, Syarinus). From Canada are registered 24 species of 19 genera and 7 families (Buddle 2005). Opiliones The catalogue of Cokendolpher and Lee (1993) facilitated the overview of Cyphophthalmi and “Palpatores” of Greenland, Canada, the USA, and Mexico, together with bibliography on the Opiliones of this area. Kury (2003) analyzed the 510 7 Regional Arachnogeography American Laniatores. Concerning Canada and the USA, we find (with supplements) the following supraspecific taxa, based on Briggs (1968, 1969, 1971a, 1971b, 1974), Briggs and Hom (1966, 1967), Briggs and Ubick (1981, 1989), Ewing (1923), Giribet and Kury (2003), Giribet and Shear (2010), Goodnight and Goodnight (1942), Hedin and Thomas (2010), Newell (1943), Richart and Hedin (2013), Shear (1975b, 1977, 1980, 2010a), Ubick and Briggs (1989, 1992, 2002, 2004), and others. Cyphophthalmi Fam. Sironidae Siro Latreille, 1804 – western USA (Oregon, California, Washington), Maryland (nine spp., endemic) Fam. Neogoveidae Metasiro Juberthie, 1960 – USA (Florida, Georgia, South Carolina, endemic) (three spp.) Eupnoi Fam. Phalangiidae – North America (Oligolophinae, Phalangiinae, Opilioninae) Fam. Sclerosomatidae – from the four subfamilies, only Leiobuninae lives in North America. Cosmobunus Simon, 1879; Eumesosoma Cokendolpher, 1980; HadrobunusBanks, 1900; Paranelima Caporiacco, 1938; Leuronychus Banks, 1900; Leiobunum C.L. Koch, 1839; Schenkeliobunum Staręga, 1964; Togwoteeus Roewer, 1952 Fam. Protolophidae (endemic family to the USA and northern Mexico) Protolophus Banks, 1893 – USA (California, Arizona, Texas, Oregon, Washington, Utah), Mexico (eight spp.) All families are shared with the Palearctic region, except for Protolophidae. Many genera are endemic to more narrow areas in North America. Dyspnoi Fam. Acropsopilionidae Acropsopilio Silvestri – USA (New York, Connecticut, Indiana, Maine, Michigan, Ohio, Wisconsin), Canada (Ontario, Quebec) (one sp.) Fam. Caddidae Caddo Banks – USA (New York, Connecticut, DC, Michigan, New Brunswick, New Jersey, etc.) (two spp.) Fam. Ischyropsalididae Subfam. Ceratolasmatinae Acuclavella Shear – Idaho, Washington (seven spp.) Ceratolasma Goodnight et Goodnight – Oregon, California (one sp.) Fam. Taracidae 7.36 North America, North of Rio Grande (incl. Greenland) 511 Crosbycus Roewer – Missouri, Georgia (one sp.) Hesperonemastoma Gruber – North Carolina, California, Washington (five spp.) Taracus Simon – Oregon, California, Colorado, Washington (seven spp.) Fam. Sabaconidae Sabacon Simon – Oregon, Alaska, Washington, North Carolina, California, Kentucky (eight spp.) Fam. Nemastomatidae Subfam. Ortholasmatinae Dendrolasma Banks – USA (Washington, California) (two spp.) Ortholasma Banks – Mexico, USA (California) (five spp.) Trilasma Goodnight et Goodnight – Mexico, Honduras (nine spp.) Laniatores In Canada only three species of Triaenonychidae are known from one province: British Columbia (Paranonychus brunneus Banks, 1893; Sclerobunus nondimorphicus Briggs, 1971; S. parvus Roewer, 1931). From Alaska is known only Paranonychus brunneus. All the other Laniatores, north of Rio Grande (29 genera and 8 families), live in the “uninterrupted” USA. Most states has one to five spp. of Laniatores (in bold the endemics for USA and/or Canada). Exceptions are California (63 species), Texas (18 spp.), and Oregon (15 spp.). The West Coast is particularly rich in Laniatores, and some are considered relicts (Briggs 1971, 1973, 1974). Fam. Triaenonychidae Subfam. Paranonychinae Metanonychus Briggs – Idaho, California, Oregon (three spp.) Paranonychus Briggs – Canada (British Columbia), USA (Alaska, Oregon, Washington) Subfam. Sclerobuninae Sclerobunus Banks (= Cyptobunus Banks) – Canada (British Columbia), USA (California, Colorado, New Mexico, Arizona, Montana, Nevada, Utah, Washington, Idaho, Oregon) (three spp.) Zuma Goodnight et Goodnight – California (two spp.) Subfam. Triaenonychinae Fumontana Shear – North Carolina, Tennessee (one sp.) Fam. Cladonychiidae Cryptomaster Briggs – Oregon (one sp.) (end.) Erebomaster Cope – DC, Maryland, North Carolina, Virginia, Indiana, Kentucky, West Virginia, Ohio (two spp.) Speleomaster Briggs – Idaho (two spp.) Theromaster Briggs – Alabama, Georgia, North Carolina, Tennessee (two spp.) 512 7 Regional Arachnogeography Fam. Pentanychidae Isolachus Briggs – Oregon (one sp.) Pentanychus Briggs – Oregon, Washington (five spp.) Fam. Cosmetidae Calicynorta Goodnight et Goodnight – California (one sp.) Denticynorta Roewer – Ohio (one sp.) Vonones Simon – Florida, Georgia, Louisiana, Oklahoma, Alabama, Illinois, Indiana, Kansas, Texas, North Carolina, Ohio, Tennessee (two spp.) Fam. Phalangodidae Bishopella Roewer – Alabama, North Carolina, Florida, Georgia, South Carolina (two spp.) Banksula Roewer – California (11 spp.) Calicina Ubick et Briggs – California (25 spp.) Microcina Briggs et Ubick – California (six spp.) Phalangodes Tellkampf – Kentucky, Tennessee (one sp.) Phalangomma Roewer – Virginia (one sp.) (endemic) Sitalcina Banks – California, Oregon (ten spp.) Texella Goodnight et Goodnight – California, New Mexico, West Virginia, Texas (21 spp.) Tolus Goodnight et Goodnight – Tennessee (one sp.) (endemic) Undulus Goodnight et Goodnight – Alabama (one sp.) (endemic) Wespus Goodnight et Goodnight – Arkansas (one sp.) (endemic) Fam. Travuniidae Speleonychia Briggs – Washington (one sp.) (endemic) Fam. Stygnommatidae Stygnomma Roewer – Florida, Ohio (one sp.) Fam. Stygnopsidae Hoplobunus Banks – Texas (two spp.) Family uncertain (Phalangodidae) Crosbyella Roewer – Alabama, Arkansas, Florida, Georgia, South Carolina (five spp.) 7.36 North America, North of Rio Grande (incl. Greenland) 513 Amblypygi This order is recorded only from the Southern USA, with six species of Phrynidae (Mullinex 1975, Quintero 1981, Harvey 2013): Acanthophrynus Kraepelin, 1899 – USA (A. coronatus Butler, 1873, Arizona, California) Paraphrynus Moreno, 1940 – USA (Arizona, Florida) (P. mexicanus Bilimek, 1867; P. raptator Pocock, 1902) Phrynus Lamarck, 1801 – USA (Ph. marginemaculatus C.L. Koch, 1841, Florida; Ph. operculatus Pocock 1902, Texas; Ph. fuscimanus C.L. Koch, 1847, Texas) These species live also south of these states (in Mexico, etc.). They mark also the northern limit of Amblypygi in North America. Thelyphonida (Uropygi) In the area North of Rio Grande is known only one species of Thelyphonida (Uropygi): Mastigoproctus giganteus (Lucas, 1835) in Texas (Thelyphonidae). This is the northernmost species of Uropygi in the New World (Rowland and Reddell 1976). Schizomida North of Rio Grande Schizomida are recorded only from four states of the USA: Arizona, California, Florida, and Texas (not counting the American territories Hawaii and Puerto Rico) (Briggs and Hom, 1966, 1972, 1988, Chamberlin 1939, Cook 1899, Gertsch 1940, Reddell and Cokendolpher 1995, Rowland 1971, 1972, 1975). Most species live in California (eight, all of the genus Hubbardia Cook, 1899). Another three species belong to the genera Bamazomus and Stenochrus. From this 11 spp., 9 are endemic: Stenochrus mulaiki (Texas) and all Hubbardia (Arizona and California). Stenochrus portoricensis is known also from Florida, Puerto Rico, Mexico, and many other countries in Central and South America and West Indies and is introduced also in the Canary Islands and England. Endemic extant genus is Hubbardia Cook (Hubbardiidae) (+ the fossil Calcitro, Calcoschizomus, and Onychothelyphonus). The family Protoschizomidae (gen. sp. indet.) is also recorded from Texas. Araneae In Canada and Alaska have been recorded 1413 spp. of 43 families of spiders (highest number in Linyphiidae – 39.5%). The richest territories are Ontario (746 spp.), British Columbia (700 spp.), and Quebec (677 spp.) (Brignoli 1973, Griswold et al. 2012, Paquin et al. 2001, Ubick et al. (eds.) (2005), Paquin et al. 2010). These authors affirm that the 1413 spp. might represent 80% of the actual total of spiders on this huge territory. According to Bennett (1999), the number of spiders in Canada is ca. 1400 spp. Except for five spp. of Mygalomorphae, all the others belong to Araneomorphae. As in most northern countries, dominant are the Linyphiidae (>500 spp.), followed by Salticidae (110 spp.), Gnaphosidae (100 514 7 Regional Arachnogeography spp.), and Theridiidae (100 spp.). These four families form more than half of all Canadian spiders. In Greenland are found 70 spp. of spiders of nine families: Araneidae, Dictynidae, Gnaphosidae, Hahniidae, Linyphiidae (45 spp.), Philodromidae, Tetragnathidae, Theridiidae, and Thomisidae (Larsen and Scharff 2003, updated 2007). Endemic family in the USA is the recently described Trogloraptoridae (Griswold, Audisio & Ledford, 2012) from caves in Oregon and California, with one genus and species Trogloraptor marchingtoni. Further are compared the families of spiders in the Nearctic (56), the Palearctic (55), and the Holarctic (68), including 44 families common between the two regions. The highest level of difference is the suborder Mesothelae, unknown in the Nearctic. The endemism on family level is insignificant. It is to notice that only Cuba, immensely smaller than the Nearctic, are known almost the same number of families (52). Opilioacarida The only representative of Opilioacarida in the USA (Texas, Arizona) is Neocarus texanus Chamberlin et Mulaik (= Neocarus arizonicus Chamberlin et Mulaik). It marks the northern limit of the order Opilioacarida (Chamberlin and Mulaik 1942, Vásquez and Klompen 2002, 2009). Parasitiformes Ixodida Gregson (1956) published from Canada 29 spp. of Ixodida, including 5 Argasidae and 24 Ixodidae. The list of localities includes some of the northernmost findings of Ixodida in the Western Hemisphere. Distribution of Arachnida on the Islands of the Pacific 7.37 7.37.1 Polynesia and Micronesia Geography, General Zoogeography, and Paleogeography In the western part of the largest ocean “...the Pacific plate preserves the oldest ocean floor on Earth at 167 Ma, of Jurassic age” (Koppers et al. 2003). The age and the origin of the Pacific islands have been explained by Neall and Trewick (2008). According to these authors, the paleocontinent of Zealandia has been displaced eastward from Gondwana between 83 and 54 Ma by mid-ocean spreading of the Tasman Sea. Remains of Zealandia are New Zealand, the Chatham Islands, and New Caledonia. Other islands are likely to have existed in the past (between 38 and 21 Ma) (Karig 1970, 1971, 1974, Menard and Hamilton 1963, Stoddart 1992). 7.37 Polynesia and Micronesia 7.37.2 515 Arachnogeography On the islands of Polynesia and Micronesia are recorded Pseudoscorpiones, Opiliones, Araneae, Schizomida, Amblypygi, very few Scorpions, Araneae, and Acari (Mesostigmata, Ixodida, Prostigmata). Some sources: Berland (1942), Kami and Miller (1998), Mayr (1944), Gressitt (1961) After a long series of publications on the spiders of the islands of the Pacific, Berland (1934b) subdivided the Pacific into six provinces: “La province australo-canaque” – Australia, New Zealand, New Caledonia, Loyalty Islands, the islands Chatham, Norfolk, and Lord Howe “La province papouasienne” – New Guinea, the islands around it, the Solomon Island, and even Vanuatu (the New Hebrides) “La province polynésienne” – Samoa, Fiji, Tonga, Marquesas, Society, Rapa Nui, and even Hawaii (? subprovince) “La province micronésienne” “La province neotropicale” (Galapagos and Juan Fernandez Islands) “La province antarctique” – the islands Campbell, Auckland, Bounty, Snares, Macquarie, attached from one side to Tierra del Fuego, on the other side to Kerguelin Lehtinen (1980) Palpigradi – No records Scorpiones On many islands (Palau, Samoa, etc.) is widespread Liocheles australasiae (Hormuridae). From Palau is recorded also Hormurus waigiensis (Gervais) from the same family (Chapin 1957, Vachon 1972). Schizomida Only Hubbardiidae indet are mentioned from Samoa, Belau, Marshall Islands, and FS of Micronesia. From Guam and North and South Mariana Islands (Asuncion, Saipan, Sarigan), Reddell and Cokendolpher (1995) notice the presence of the genus Orientzomus. Cokendolpher and Reddell (2000) recorded from the Marshall Islands the first two (new) species of Schizomida: Apozomus brignolii and Orientzomus ralik, Cokendolpher and Tsurusaki (1994). Villarreal Manzanilla (2010) announced the presence of genus Zomus on Samoa. The list of pseudoscorpions of Polynesia and Micronesia counts eight families: Chthoniidae, Tridenchthoniidae, Geogarypidae, Olpiidae, Cheiridiidae, Atemnidae, Chernetidae, and Withiidae (Beier 1940, 1957, Chamberlin 1934, 1938, 1939a, 1939b, 1947). Endemic genera for Polynesia and Micronesia are: Fam. Chernetidae Meiochernes Beier, 1957 – Caroline Islands (one sp.) 516 7 Regional Arachnogeography Opiliones – seven genera of four families (all Laniatores) Cyphophthalmi are not known in Polynesia and Micronesia. Dyspnoi are not known in Polynesia and Micronesia. Laniatores – Roewer (1912), Goodnight and Goodnight (1957), Suzuki (1941) Fam. Epedanidae – Dibunus Loman, 1906 Fam. Podoctidae – Metibalonius Roewer, 1915, Lomanius Roewer, 1923 Fam. Samoidea – Parasamoa C.J. Goodnight et M.L. Goodnight, 1957; Feretrius Simon, 1879 (one sp.); Samoa Sørensen in L. Koch, 1886 Fam. Zalmoxidae – Zalmoxis Sørensen, in L. Koch 1886 Amblypygi Fam. Charontidae: On Palau has been found Charon grayi (Gervais, 1842) (Esaki 1936, Takashima 1950). Araneae Berland (1934b) subdivided the Pacific islands into four groups. According to him: All information seems to indicate that the settlement of Pacific was done by migrations coming from the Indo-Malaysian area, migrations which would probably have been multiple and following several distinct currents. One of these currents would have been populated from Australia; this is what I call the “australo-canaque” province; a quite distinct current of migration, but of same origin, would have populated Polynesia, whose current archipelagoes constitute probably only the subdivision of a previously more extended continent, with a branch being detached towards Hawaii; another towards Micronesia. There is no visible relation between America and the Pacific; but on the other hand Galapagos as well as Fernandez received their fauna from the south of America, by the austral lands. In addition the islands of Pacific almost always present a very marked level of endemism, which testifies to an extremely old isolation, and should be fix their separation farther back, at least to about the middle of the Tertiary epoch, and perhaps farther. I will add that the study of several zoological groups, as one can see it in this work, arrive, independently from each other, at conclusions very close to mine, and so appear to me to receive a solid confirmation from it. 7.37 Polynesia and Micronesia 517 Map 7.28 Zonation of Micronesia and Polynesia (After Berland 1934) Berland supported the view that the islands of Polynesia were formed as a single large continent, which became subdivided to form the islands. Recent geological evidence has dismissed this view. The archipelagoes of Hawaii, Marquesas and Societies are all volcanic in origin, and formed as volcanic hot spots. The Marquesas range from Nuku Hiva, the oldest in the north at 3.7myrs, to Fatu Hiva the youngest, in the south, at 1.4 myrs. The Society Islands range from Bora Bora, the oldest in the north at 3.3 myrs, to Tahiti the youngest, in the south, at 1.0 myrs. (Gillespie et al. 2000) 518 7 Regional Arachnogeography Map 7.29 Spreading of Araneae to the islands of the Pacific (Berland, 1934) Some endemic genera are: Fam. Linyphiidae Paro Berland, 1942 – Rapa (one sp.) Uahuka Berland, 1935 – Marquesas Islands (two spp.) Uapou Berland, 1935 – Marquesas Islands (one sp.) Fam. Dictynidae Banaidja Lehtinen, 1967 – Samoa (one sp.) Fam. Pisauridae Nukuhiva Berland, 1935 – Marquesas (one sp.) Fam. Tetrablemmidae Rhinoblemma Lehtinen, 1981 – Caroline Islands (one sp.) 7.38 Hawaii 519 Fam. Anyphaenidae Australaena Berland, 1942 – Polynesia (two spp.) Fam. Salticidae Iona Peckham et Peckham, 1886 – Tonga Rarahu Berland, 1929 – Samoa Other sources: Beatty et al. (2008), Berland (1924, 1935a, 1935b, 1935c, 1939, 1942), Evenhuis (2006), Garb (2003), Garb and Gillespie (2006), Gillespie (2003b, c), Gillespie et al.(2002), Lehtinen (1993), Marples (1955), Proszyński (1996) 7.38 7.38.1 Hawaii Geography, General Zoogeography, and Paleogeography Hawaiian islands are the exposed peaks of a great undersea mountain range known as the Hawaiian-Emperor seamount chain, formed by volcanic activity over a hot spot in the Earth’s mantle. The Hawaiian islands are about 3,000 km from the nearest continent. The archipelago consists of eight greater and many smaller islands. The main islands are Hawaii (10,432 km2, with the highest point Mauna Kea, 4205 m), Maui (1883 km2), Oahu (1545 km2), Kauai (1430 km2), Molokai (673 km2), Lanai (364 km2), Niihau (180 km2), and Kahoolawe (116 km2). The total area of the archipelago is 16,636.5 km2. The islands range from Kauai, the oldest in the north at 5.1myrs, to Hawaii the youngest, in the south, at up to 0.4 myrs old. It is considered that the entire fauna of these islands, originally azoic, is a result of the three W (wind, waves, wings) – the ancestors either arrived by rafting or were windblown or have been brought with birds or bats (Wagner and Funk 1995, Zimmerman 1948). 520 7 Regional Arachnogeography Map 7.30 The Hawaiian Islands 7.38.2 Arachnogeography These islands, completely azoic in the beginning, are now housing several orders of Arachnida: Palpigradi, Scorpiones, Schizomida, Pseudoscorpiones, and Araneae and different mites. Even more, there are endemic genera of pseudoscorpions and spiders and amazing cave fauna (troglobites in recently formed lava tubes). Some other arachnids are clearly adventive from other lands, including Thailand and North America (Berland 1934, Nishida 1994, 1997, 2002, Simon 1900). Palpigradi On Hawaii are known two species of Palpigradi, both from the family Eukoeneniidae – Eukoenenia florenciae (Ruckner, 1903) and E. hanseni (Silvestri, 1913). Both species are known also from the continental USA (Louisiana, Texas) (Harvey 2013e, Rémy 1961a). Scorpiones Isometrus maculatus (De Geer) is distributed in Oahu, Maui, and Hawaii (Yates 1993). Schizomida Known is only Bamazomus siamensis (Hansen) (Oahu, also from Thailand, Hong Kong, and the Ryukyu Islands) (Reddell and Cokendolpher 1995). Pseudoscorpiones In the first checklist of Pseudoscorpiones of Oceania (Chamberlin 1934) are listed six spp. from Hawaii. Important contribution to the record on the Pseudoscorpiones of Hawaiian Islands was made by Muchmore 7.38 Hawaii 521 (1979, 1983, 1989, 1996, 2000). Now (2017) from the islands are known the families Chthoniidae, Lechytiidae, Garypinidae, Chernetidae, Cheliferidae, Cheiridiidae, Geogarypidae, Olpiidae, and Withiidae (Beier 1932, 1940, Muchmore 2000, Harvey 1990, Harvey 2013f, Simon 1900, With 1905), according to Nishida (2002), 17 spp. Endemic genera for Hawaiian Islands are: Fam. Chthoniidae: Vulcanochthonius Muchmore, 2000 (three spp.) Fam. Chernetidae: Eumecochernes Beier, 1932 (three spp.) Of special interest are the troglobitic species in the lava tubes (Muchmore, 1979, 1983, 1989). Three species have been described: Tyrannochthonius howarthi, T. stonei, and T. pupukeanus. T. howarthi has been transferred to the new genus Vulcanochthonius Muchmore, 2000, including V. pohakuloae Muchmore, 2000. In his description of Tyrannochthonius howarthi as the first cave pseudoscorpion in Hawaii, Muchmore (1982) expressed some doubt concerning the real place of this species in the system. According to him, “...it appears quite possible that it actually belongs to a presently unrecognized group (genus or subgenus) with a wide distribution in the Pacific.” Opiliones In the checklist of terrestrial arthropods of Nishida (2002) are not mentioned any Opiliones inhabiting Hawaii. Araneae After Simon’s “Fauna Hawaiiensis” (1900), in the catalogue of Suman (1964) have found place 149 spp. of 21 fam. and 66 genera. This number includes 82 endemic species (55%) of 10 fam. and 27 genera. By 2005 on the Hawaiian Islands were known 132+ native spp. of spiders of 36 native genera and 16 native families. Particularly interesting are the blind spiders, found by Howarth (1980) in the relatively recent lava caves (Adelocosa anops Gertsch). At least 128 spp. are endemic species. “Eighty percent of the known Hawaiian native spider species belong to ten genera in seven families Cyclosa (Araneidae), Orsonwelles (Linyphiidae), Pagiopalus and Pedinopistha (Philodromidae), Havaika (Salticidae), Tetragnatha (Tetragnathidae), Argyrodes and Theridion (Theridiidae), and Mecaphasa and Misumenops (Thomisidae)” (Hawai’i Comprehensive Wildlife Conservation Strategy, 2005). Gertsch (1973) identified the collection of 20 spp. of spiders, found in the lava tubes. Most were introduced, but six were eyeless or with eyes reduced to vestiges (troglobites). According to Gertsch (loc. cit.), “Most important are the two new wolf spiders (Lycosidae), one with vestigial eyes and the other a completely eyeless species of this family of big-eyed hunting spiders.” Other sources are: Garb (1999), Garb and Gillespie (2009), Gillespie (1991), Gillespie et al. (1998), Gillespie et al. (2000), Gillespie et al. (2003), Simon (1899 – 1900), Suman (1964, 1965, 1967, 1970). Some endemisms: Fam. Linyphiidae Orsonwelles Hormiga, 2002 Priperia Simon, 1904 (one sp.) 522 7 Regional Arachnogeography Fam. Lycosidae Adelocosa Gertsch, 1973 – Kaua’i, caves (blind) Syroloma Simon, 1900 Fam. Tetragnathidae Doryonychus Simon, 1900 (one sp.) Fam. Philodromidae Subfam. Pedinopisthinae Pagiopalus Simon, 1900 Pedinopistha Karsch, 1880 (five spp.) Fam. Thomisidae Mecaphesa Simon, 1900 Fam. Salticidae Havaika Prószyński, 2002 – 23 spp. on Hawaii, three spp. on Marquesas Islands Also more than 30 local genera Acari According to Garrett and Haramoto (1967), until this time on Hawaiian Islands have been recorded 210 species of mites and ticks of 67 families and 138 genera. According to Nishida (2002), on the islands are recorded 928 spp. of Acari, including 296 endemic (165 families, 505 genera). Acariformes Garrett and Haramoto (1967) indicate on the archipelago 21 families of Prostigmata and 18 families of Oribatida. Parasitiformes (Ixodida and Mesostigmata) Ixodida – until 1967 in Hawaii have been registered four spp. of Argasidae and eight spp. of Ixodidae (mostly quarantine finds). Ixodes laysanensis Wilson, 1964, is described on birds. Mesostigmata – Garrett and Haramoto (1967) indicate 15 families on the islands. 7.39 7.39.1 Antarctic and Subantarctic Arachnofauna Geography, General Zoogeography, and Paleogeography Subantarctic are termed the islands around Antarctica situated north of and adjacent to the Antarctic Convergence (between the 48th and 61st parallel of south latitude). The Falkland Islands, Prince Edward Islands, Crozet Islands, Amsterdam Island, St Paul Island, Tierra del Fuego, and Macquarie Island lie north of the Antarctic 7.39 Antarctic and Subantarctic Arachnofauna 523 Convergence. Other bigger Subantarctic islands of island groups are Antipodes Islands, Auckland Islands, Bounty Islands, Macquarie Island, Prince Edward Islands, The Snares, and Tierra del Fuego. Some of the Antarctic islands also lie north of 60oS: Bouvet Island, Heard Island, McDonald Islands, Kerguelen Islands, South Georgia Islands, South Sandwich Islands, and others (Gressitt 1964, 1965, 1970, Gressitt and Weber 1959, Morrone 2000b). According to Morrone (2000), biogeographically the Subantarctic Subregion belongs to the Andean region of the Austral Kingdom and comprises the Austral Andes from 37oS to Cabo de Hornos, the archipelago of southern Chile and Argentina, and the Malvinas, South Georgia, and Juan Fernandez Islands (six provinces: Maule; Valdivian; Magallanic forest; Magallanic moorland; Malvinas islands, including South Georgia; and Juan Fernandez Islands). For each province are listed, among the other animals, the typical elements of Arachnida. 7.39.2 Arachnogeography On the Antarctic continent live only some groups of mites. From the other groups on the Subantarctic islands are known some pseudoscorpions, opilions, and spiders (Pugh 1993, 1997, 2003). If the most southern South America is included in the term “Subantarctic,” we find many more groups of Arachnida, but the orders listed above still prevail. The ice-covered Antarctica with harsh climate and almost no vegetation and the islands inside the Antarctic Convergence (N) have very limited arachnofauna, composed mostly of Araneae and Acari. This huge area has not been always ice-covered, and there are hypotheses for transantarctic ways of dispersion for many groups of Arachnida, inhabiting the southern continents (Patagonia, Southern Africa, Australia, New Zealand, New Caledonia). Some sources: Dalenius and Wilson (1958), Fleming (1987), Forster (1964, 1970, 1971), Gressitt (1965, 1971), Hill (2009), Marshall and Pugh (1996), Morrone (2000), Pugh (1994, 1997, 2004), Wallwork (1963, 1965, 1966, 1967, 1969, 1973, 1979) Presently “0.34% of its area is currently ever free of snow or ice in the form of terrestrial ecosystems including exposed nunataks, cliffs and seasonally snow and ice-free areas” (Convey 2010, after British Antarctic Survey, 2004). Udvardy (1987) proposed the creation of “biogeographical realm Antarctica” including four provinces: Maudlandia, Marielandia, Insulantarctica, and Neozealandia. After Hill (2009), the Antarctic land bridge, associated with a tropical to temperate climate (Francis et al., 2008), was available to support the dispersal of plants and animals for about 75–95 million years after the separation of Africa. The study of the few arthropods in Antarctica and Subantarctic is important in many ways – to understand the ways of dispersal and the abilities of different groups to exist in harsh and extreme conditions of cold and devoid of vegetation environ- 524 7 Regional Arachnogeography ment. The only groups represented there are Acari (Acaridida, Oribatida, Trombidiformes) and Araneae. Some parasitic ticks and mites are brought to these lands by birds (Ixodida). Pseudoscorpiones From the Subantarctic islands outside the ones belonging to New Zealand and Australia (see New Zealand), Falkland Islands, and Tierra del Fuego (see Patagonia) are recorded the following pseudoscorpions (Chamberlin 1933, Vitali-di Castri 1968): Ile de la Possession, île Crozet – Austrochthonius insularis Vitali-di Castri St Paul – Diplotemnus insularis Chamberlin Opiliones. Forster (1955a) Opiliones reported from Subantarctic and extreme southern cold temperate islands (after Cokendolpher and Lanfranco 1985): Fam. Triaenonychidae Neonuncia campbelli Forster – Campbell Islands (52o33′), cold temperate (Forster 1954, 1964) Neonuncia eastoni Forster – Auckland Islands (50o 32′), cold temperate (Forster 1954, 1964) Neonuncia enderbyi (Hogg) – Auckland and Campbell Islands (50°32”, 50° 33”), cold temperate (Forster 1954, 1964) Nuncia unifalculata (Enderlein) – Crozet Islands (46o30”), Subantarctic (TambsLyche 1954, Hickman 1939) Fam. Gonyleptidae Lycomedicus planiceps (Guerin) – Isla Hoste (55o 10”), cold temperate (Soares and Soares 1954) Haversia defensa (Butler) – Islas Malvinas (52o), cold temperate (Soares and Soares 1949) Haversia [=Hoggellula] vallentini (Hogg) – Islas Malvinas (52o), cold temperate (Soares and Soares 1949) Fam. Megalopsalididae Pantopsalis distincta Forster – Auckland Islands (50° 32′), cold temperate (Forster 1964) Pantopsalis johnsi Forster – Auckland Islands (50o 32′), cold temperate (Forster 1964) Pantopsalis mila Forster – Auckland Islands (50o32′), cold temperate (Forster 1964) Pantopsalis rennelli Forster – Campbell Island (52o33′), cold temperate (Forster 1964, Gressitt et al. 1964) Pantopsalis snaresensis Forster – Snares Island (48o), cold temperate (Forster 1964) 7.39 Antarctic and Subantarctic Arachnofauna 525 Fam. Neopilionidae Thrasychirus dentichelis Simon – Isla de los Estados, Isla Hoste (54o50′, 55o30′), cold temperate (Ringuelet 1959) Cape Horn Archipelago 55o37′–49′ cold temperate Thrasychirus gulosus Simon – Isla de los Estados, Isla Hoste (54o50′, 55o30′), cold temperate (Ringuelet 1959) Thrasychirus modestus Simon – Isla Hoste, Isla Navarino (55o 10′–30), cold temperate (Cekalovic 1976), Isla Deceit (55o49′) cold temperate Araneae In his paper on the spiders of islands Chatham, Bounty, Snares, Auckland, Campbell, Macquarie, and Kerguelen, Berland (1930a) pays special attention to the genus Myro P.-C. (Desidae), known from Kerguelen, Macquarie, South Africa, Tasmania, and the extreme south of South America. His conclusion is that “il jalonne donc admirablement un continent qui auraît réuni l’Amérique à l’Australie par les subantarctiques et la Tasmanie, à Kerguelen et même au sud de l’Afrique.” Now the distribution of this genus is restricted to Crozet, Kerguelen, Macquarie, New Zealand, Marion, and Tasmania. There are other genera of intertidal spiders living in South America (Porteria in Chile and even Desis galapagoensis Hirst on Galapagos Islands), but they make no connection with Australasia or Kerguelen. Berland also affirm that the spiders found on the small islands satelites “n’ont à peu près aucune affinité avec la Nouvelle-Zélande.” Actually, now we know that very few of the numerous genera of Desidae of New Zealand are represented also on the Subantarctic islands (Gasparia Marples, Gohia Dalmas, Myro P.-Cambridge, Laestrygones Urquhart). Forster (1955, 1962, 1970, 1971) completed vastly the knowledge on the spider fauna of Subantarctic islands. Pugh (2004) analyzed the biogeography of spiders on the islands of the Southern Ocean. Araneae In his special paper on the possibility of mites to colonize Antarctica and the islands of the Southern Ocean via air currents, Pugh (2003) indicates that among the spiders “most long-range aerial colonists are minute adult Araneidae, Lycosidae, Theridiidae, and especially Linyphiidae, representing 81% of verified specimens collected over the Southern Ocean and 46% of Southern Ocean species.” The juveniles have a low colonization success. In a biogeographic analysis of the spiders on the islands of the Southern Ocean, Pugh (2004) indicates that only 115 verified species from 26 families are reported from these islands. On the continent the spiders are anthropogenic immigrants. A young spider in good condition was trapped at Marble Point on the Victoria Land coast opposite Ross Island (Gressitt 1964). Endemic genera: Fam. Linyphiidae: Ringina Tambs-Lyche, 1954 – Crozet Fam. Theridiidae: Icona Forster, 1955 – Auckland, Campbell Islands (two spp.) 526 7 Regional Arachnogeography Acari According to the prominent specialist of Antarctic ecology and biogeography Prof. J.L. Gressitt, “The mites and ticks may well prove to be, in the number of species, the richest terrestrial animal group in Antarctica. The mites seem to be particularly tolerant of low temperatures. They have been found equally as far south as Collembola (83°55’S)” (Gressitt 1964). What concerns the dispersal of Acari (and other Arachnida) in Antarctica, Gressitt (1964) says: “If the quaternary climates on the continent was too severe at its coldest for the survival of remnants of the preceding temperate fauna, than it is likely that all or much of the present fauna has resulted from immigration in fairly recent time. This suggests transport in air currents, or by the agency of birds.” He points that a mite has been trapped 3000 meters above sea level at a point halfway between New Zealand and Antarctica. A comprehensive checklist of the Antarctic Acari is due to Pugh (1993). It contains 528 spp. After Pugh (1994), about 70 spp. are nonindigenous. About mites Pugh (2003) concludes that some are probably phoretic (Ascidae, Parasitidae, Uropodidae and Veigaiidae) and that “…all Bdellidae, Eupodidae, Acaridae, Tyroglyphidae, and Oribatida (= Cryptostigmata) relate to local edaphic/ hemiedaphic mites that were locally uplifted by storm and/or imported as shipborne (anthropogenic) contaminants.” Concerning the Oribatida (Cryptostigmata), Wallwork (1969) has adopted the scheme proposed by Holdgate (1964): 1. Subantarctic zone Kerguélin I., Prince Edward I., Crozet I., Macquarie I., South Georgia 2. Maritime Antarctic zone Bouvet Island, South Orkney Islands, South Sandwich Islands, South Shetland Islands, Antarctic Peninsula 3. Continental zone Eastern Antarctic Continent From his analysis Wallwork (1969) has drawn the following (tentative) conclusions: 1. The Continental zone has an impoverished, highly endemic fauna. 2. Endemism in the Maritime zone is low compared with that of the Subantarctic zone, the fauna of the former being clearly derived from that of the latter. 3. The Subantarctic zone supports a much richer fauna than the other two zones, and the relatively high endemicity (probably estimates) of its fauna suggests that this zone may be recognized as a distinct faunal region, as far as the Cryptostigmata are concerned. Stary and Block (1998) consider 105 spp. of 20 fam. of Oribatida to inhabit the Antarctic and Subantarctic territories. 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Aust Nat Antarct Res Exped Reports (B) 1 (Zool.):12–20 Chapter 8 Some Peculiarities of the Distribution of Arachnida Abstract Special attention is payed to the comparison between the arachnofaunas of close areas: Mediterranean and Central European; Mexico and the USA; Central America and the Caribbean Islands; South Africa, Madagascar, Seychelles, and Mascarene; and special areas like Patagonia and New Guinea, Australia, New Zealand, New Caledonia, and Lord Howe Island. 8.1 Mediterranean and Central European Arachnofauna Both the classic Mediterranean and European-Siberian (Eurosiberian) areas are in the Palearctic and usually are considered subregions. Malicky et al. (1983) presented arguments for the existence and the horology of special Middle European (ExtramediterranEuropean) elements in the fauna of Europe. In Middle Europe are missing many orders of Arachnida (Amblypygi, Uropygi, Schizomida, Ricinulei, Opilioacarida, Holothyrida); groups like Palpigradi, Cyphophthalmi, Laniatores, and Scorpiones are represented by symbolic number of species. Well represented are Araneae, Opiliones (Eupnoi, Dyspnoi), and Pseudoscorpiones. Cave Arachnida are much less represented than in the Mediterranean, and troglobites North of the Alps are almost missing. According to Lopatin (1980), the European-Siberian Region within the Palearctic Subkingdom is subdivided into European-Obian subregion (from Great Britain to Kamchatka and Chukotka) and Angaran subregion (Siberia East of Yenisei – the Johansen’s Line – as far as Sakhalin). Both subregions are subdivided into landscape zones: tundra, taiga, steppe, and high mountains. The Asiatic part of EuropeanSiberian area is with much harsher climate than in Middle Europe. Spiders are prevailing; the part of Linyphiidae is bigger in the Nordic areas. 8.2 Arachnofauna of the Ancient Mediterranean According to Lopatin (1980), Palearctic is a subkingdom, including three regions: European-Siberian, Region of the Ancient Mediterranean, and East Asiatic Region. The Region of the Ancient Mediterranean consists of two subregions – Mediterranean © Springer International Publishing AG, part of Springer Nature 2018 P. Beron, Zoogeography of Arachnida, Monographiae Biologicae 94, https://doi.org/10.1007/978-3-319-74418-6_8 625 626 8 Some Peculiarities of the Distribution of Arachnida and Saharo-Gobian. These are territories around the Mediterranean Sea and a large band from North Africa to Central Asia and Gobi. According to the map (Lopatin 1980, p. 125), the southern border of this region follows the northern limit of the Afrotropical Region (Paleotropical Kingdom) and also runs on the northern slope of the Himalaya. The northern limit is following the Pyrenees, Alps, Danube, and the mountains of Central Asia. The Mediterranean Subregion is divided into provinces: Macaronesian, Tyrrhenian or West Mediterranean, and East Mediterranean. The vast territory of the Saharo-Gobian Subregion includes three superprovinces: Saharo-Sindian, Irano-Turanian, and Central Asiatic. In the respective parts of this book are enumerated in details the Arachnida of each of these territories. Some orders are found only in the Mediterranean Subregion, much warmer than the other parts of the Palearctic, or are barely represented. Such groups are the Amblypygi (Charinus, Musicodamon), the majority of Solifugae and Scorpiones, many families of Opiliones and Araneae, most of the Palpigradi, and most of the relicts (scorpions, opilions, pseudoscorpions). Mediterranean or submediterranean elements are found even in Romania and southern Slovakia, but actually the full Mediterranean fauna stops about the mountains on the border between Bulgaria and Greece. Submediterranean elements (Galeodes, scorpions, and other groups) enter the present-day Bulgaria only along the valleys of Struma, Mesta, and Maritza. Otherwise most of the Bulgarian territory is with Central European elements. 8.3 The Boundary Neotropica-Nearctic In North America, north of Mexico, are found 13 orders of Arachnida, but the “southern” orders (Amblypygi, Ricinulei, Schizomida, Uropygi, Opilioacarida, and largely extending to Solifugae and Scorpiones) are confined mostly to Texas, Arizona, California, Florida, Louisiana, and New Mexico, some of them being on the edge of their areals (Dice 1943, Harvey, on line, Rowland and Reddell 1976; Sørensen 1898). 8.3.1 Palpigradi Mexico – Eukoenenia hanseni (Silvestri), E. corozalensis Montaño et Francke, and E. tetraplumata Montaño (Eukoeneniidae). USA – In Texas and Louisiana are recorded Eukoenenia florenciae (Rucker, 1903), E. hanseni (Silvestri, 1913), Prokoenenia californica (Silvestri, 1913) (California), and P. wheeleri (Rucker, 1901) (Texas). In North America, the order is known from the area up to 44oN. 8.3 The Boundary Neotropica-Nearctic 8.3.2 627 Ricinulei Mexico – Nine species of the genus Pseudocellus Platnick. USA – The only recent species of Ricinulei from the USA (Pseudocellus dorotheae (Gertsch et Mulaik)) has been described from Edinburg, Texas, and Hidalgo County, on the border of Mexico. Solifugae According to the checklist of Brookhart and Brookhart (2006), in continental North America have been recorded 196 species of Solifugae, from Eremobatidae (174 sp.) and Ammotrechidae (22 sp.). Scorpiones Mexico – 272 spp. of 37 genera and 8 families. The biggest genus is Vaejovis C.L. Koch (61 sp.). Remarkable is the presence of cave species of the family Typhlochactidae. This family and many genera (Typhlochactas, Stygochactas, Sotanochactas, Troglocormus, Alacran, Hoffmannichadrurus, Chaneke, Koloti, Balsateres, Konetontli, Kuarapu, and others) are endemic for Mexico. USA – In North America, north of Mexico (30 states of the USA), are distributed 92 spp. of scorpions of 13 genera and 5 families. Paruroctonus boreus (Vaejovidae) is the only species that occurs in Canada (southern Saskatchewan, Alberta, and British Columbia). Altogether in the USA are registered 111 spp. of 20 genera and 6 families. The states richest in scorpions are California (51 sp.), Arizona (41 spp.), New Mexico (20 spp.), Texas (20 spp.), Nevada (16 spp.), and Utah (12 spp.). 8.3.3 Pseudoscorpiones Mexico – Recorded are 161 spp. of Pseudoscorpiones of 63 genera and 18 families. Six genera are endemic to Mexico. USA – Now (2017) we may assume that in North America, north of Rio Grande are recorded about 400 species of pseudoscorpions, belonging to 105 genera and 19 families. 8.3.4 Opiliones Mexico – 283 spp. (1 sp. of Cyphophthalmi, 143 Laniatores, and 139 “Palpatores”). Cyphophthalmi – Shearogovea mexasca – is a Gondwanan Neotropical element; the laniatorid genera all have closest relations with the Neotropical fauna, including the 16 endemic genera. About half of the “Palpatores” have close relations with the Nearctic and half with the Neotropical fauna. Acropsopilio is considered to be a relictual genus (Kury and Cokendolpher 2000). USA – The catalogue of Cokendolpher and Lee (1993) facilitates the overview of Cyphophthalmi and “Palpatores” of Greenland, Canada, the USA, and Mexico, 628 8 Some Peculiarities of the Distribution of Arachnida together with a bibliography on the Opiliones of this area. Kury (2003) analyzed the American Laniatores. In the USA are represented all subgenera of Opiliones: Cyphophthalmi (Sironidae). 8.3.5 Amblypygi In Mexico and the southern states of the USA (Arizona, California, Florida, Texas) are found the same genera of Amblypygi (all Phrynidae): Acanthophrynus Kraepelin, Paraphrynus Moreno, and Phrynus Lamarck. They mark also the northern limit of Amblypygi in North America. The family Phrynidae is clearly Neotropical, despite the finding of one species in Indonesia (Flores) (according to Armas 2006, “pudiera ser errónea”). 8.3.6 Uropygi (Thelyphonida) Mexico – Recorded is only one member of Mastigoproctus Pocock, 1894, M. giganteus (Lucas, 1835) (Uroproctinae). The genus is known from South and Central America, the West Indies, and the Southern USA (Texas). USA – One species of Uropygi: Mastigoproctus giganteus (Lucas, 1835) in Texas (Thelyphonidae), the northernmost species of the order in the New World 8.3.7 Schizomida Mexico – Thirty-six species from the genera Mayazomus, Pacal, Stenochrus, Protoschizomus, Agastoschizomus, Sotanostenochrus (all genera, except Stenochrus, are endemic to Mexico.) USA (Arizona, California, Florida, and Texas) – Most species live in California (eight, all of the genus Hubbardia Cook, 1899). Also the genera Bamazomus and Stenochrus. All Hubbardia (Arizona and California) are endemic in the USA. The family Protoschizomidae (gen. sp. indet.) is also recorded from Texas. 8.3.8 Araneae Mexico – The Mexican spider fauna consists of 2295 spp. in 423 genera and 64 families of Araneomorphae (Orthognatha), no endemic families. USA – Endemic family in the USA is the recently described Trogloraptoridae (Griswold, Audisio, and Ledford 2012) from caves in Oregon and California, with one genus and species Trogloraptor marchingtoni. 8.4 Mexico: The Great American Interchange of Fauna 8.3.9 629 Opilioacarida Mexico – At least five species, all of genus Neocarus Chamberlin et Mulaik USA (Texas, Arizona) – One species (Neocarus texanus Chamberlin et Mulaik), marking the northern limit of Opilioacarida 8.4 8.4.1 Mexico: The Great American Interchange of Fauna Geography, General Zoogeography, and Paleogeography South of Rio Grande is Mexico – 1,972,550 km2 of lands with exceptional biodiversity. High mountains form the Cordillera Volcanica Transversal. We had the chance to visit the highest summits and to collect some arachnids and other high mountain animals: Îrisaba (5700 m.), Popocatepetl (5492 m.), and Nevado de Toluca (4680 m.). Between Mexico and Colombia is stretching one of the richest in biodiversity territories on Earth – Central America. Geographically part of North America, this territory has nevertheless many distinctive patterns and is often considered a subcontinent (like India). There are two main definitions of the notion of Central America. The first comprises the seven states between Mexico and Colombia: Belize, Guatemala, El Salvador, Honduras, Nicaragua, Costa Rica, and Panama (total surface of 524 000 km2). According to the other concept, Central America starts with the Tehuantepec land bridge in Mexico – the 200 km. shortest distance between the Atlantic and the Pacific Oceans. This concept includes in Central America the very rich in biodiversity Yucatan Peninsula – I find this concept more natural, as Yucatan is narrowly attached to the land south of it. Along both seashores of Mexico, the Neotropical elements penetrate north. In the central parts extends dry plateau with coniferous forests and mostly of Nearctic caractere (Freeland and Dietz 1971; Martin and Harrell 1957; Barrera 1962; Escalante et al. 2007; Espinosa et al. 2000; Ferrusquía-Villafranca 1990; Halffter 1978, 1987, 2003; Marshall and Liebherr 2000; Morrone et al. 2002). The Mexican entomologist Prof. Gonzalo Halffter introduced (Halffter 1977) a particular term within Mexican Transition Zone, called in Spanish “Mesoamericano de Montaña” (the mountain rain forests, populated with animals, having evolved in the “Central American Nucleus”). Halffter (1976) describes in details the distribution of insects in Mexican Transition Zone. Halffter’s Mexican Transition Zone is a complex area in which the Neotropical and Nearctic regions overlap. The generalized tracks obtained correspond broadly to Halffter’s Nearctic and Mesoamerican patterns, respectively; however, in contrast to Halffter’s Mesoamerican pattern, the southern generalized track extends further north through the lowlands of the Pacific and Mexican Gulf coasts (Morrone and Márquez 2001). The Mesoamerican distributional pattern includes Neotropical taxa that evolved in situ in humid montane habitats (Morrone 2005). 630 8 Some Peculiarities of the Distribution of Arachnida According to Morrone (2005), Mexican Transition Zone between Nearctic and Neotropic includes five provinces: Western Sierra Madre, Eastern Sierra Madre, Trans-Mexican Volcanic Belt, the basins Balsas, and Sierra Madre del Sur. Morrone (2006) also includes in the biogeographical subdivision of “Latin America” the notion of “Mexican Transition Zone.” The development of the Mexican Transition Zone is summarized by Morrone (2015) into five stages: (a) Jurassic-Cretaceous, four Paleoamerican cenocrons extend in Mexico; (b) Late Cretaceous-Paleocene, dispersal from South America of the Plateau cenocron; (c) Oligocene-Miocene, dispersal from Central America of the Mountain Mesoamerican cenocron; (d) Miocene-Pliocene, dispersal from North America of the Nearctic cenocron; and (e) Pleistocene, dispersal from South America of the typical Neotropical cenocron. Arachnogeography The arachnofauna of Mexico is very rich and interesting as a transition zone between two kingdoms – Holarctic (the Nearctic Region) and Neotropica (Vázquez 1995, 1996). In this country are lacking only the suborders Palaeoamblypygi and Mesothelae. All orders of Arachnida are represented in Mexico, some of them with many taxa. Especially interesting are the many cave scorpions of the family Typhlochactidae and the 272 species of Opiliones in this transitional country. Map 8.1 Mexico and the Southern USA 8.4 Mexico: The Great American Interchange of Fauna Ricinulei: Amblypygi: Cyphophthalmi: Opilioacaridae: Holothyrida: 631 Pseudocellus – ◇ Phrynus – ● Paraphrynus – ▶ Acanthophrynus – ▼ Heterophrynus – ◑ Charinus – ◻ Shearogovea – ♣ Metasiro – ♠ Neocarus – ☒ Caribeacarus – △ Caribothyrus – ◓ Palpigradi From Mexico are known Eukoenenia hanseni Silvestri, 1913; E. corozalensis Montaño et Francke, 2006; and E. tetraplumata Montaño, 2007 (Eukoeneniidae) (Condé 1986; Harvey 2013e). Ricinulei From Mexico are known nine species of the genus Pseudocellus Platnick, 1980 (Bolivar y Pieltain 1941, 1946; Brignoli 1974; Chamberlin and Ivie 1938; Coronado-Gutiérrez 1970; Gertsch 1971; Goodnight and Goodnight 1952; Marquez and Conconi 1974; Valdez-Mondragón and Francke 2011). Solifugae In Mexico are known 77 spp. of Solifugae from 12 genera and 2 families: Eremobatidae – Eremorhax Roewer, Eremobates Banks, Eremocosta Roewer, Eremothera Muma, Eremochelis Roewer, Chanbria Muma, Hemerotrecha Banks Ammotrechidae – Ammotrechella Roewer, Ammotrechula Roewer, Ammotrecha Banks, Branchia Muma, Ammotrechesta Roewer Most of these genera are shared with the adjacent states of the USA (all Eremobatidae) or with Florida, Central America, South America, or the Antilleans (Ammotrechidae) (Muma 1976; Vazquez Rojas 1981, 1995; Brookhart and Muma 1987; Brookhart and Brookhart 2006). Scorpiones One of the richest countries with scorpions, According to Díaz Nájera (1975), in Mexico live 134 spp. of Scorpiones, belonging to 15 genera and 4 families – Buthidae, Chactidae, Scorpionidae (incl. Diplocentridae), and Vaejovidae. Since this publication many changes occurred, especially with the cave scorpions (Francke 1977, 1981, 1982; Lourenço 1994; Lourenço and Sissom 2000; Mitchell 632 8 Some Peculiarities of the Distribution of Arachnida 1968, 1971; Mitchell and Peck 1977; Prendini et al. 2010; Soleglad and Fet 2005, 2006, 2008; Williams 1970). Now (2017) the scorpions recorded from Mexico belong to 272 spp. of 37 genera and 8 families. The biggest genus is Vaejovis C.L. Koch (61 spp.). Remarkable is the presence of cave species of the genera Typhlochactas Mitchell, Stygochactas Vinoli et Prendini, Sotanochactas Francke, Troglocormus Francke, and Alacran Francke (Typhlochactidae). According to Lourenço (1994), in Mexico live 11 out of the 13 troglobite scorpions in the world. Since then, some more troglobitic genera and species have been published (Francke 2009). The family Typhlochactidae and many genera (Typhlochactas, Stygochactas, Sotanochactas, Troglocormus, Alacran, Hoffmannichadrurus, Chaneke, Koloti, Balsateres, Konetontli, Kuarapu, and others) are endemic to Mexico (Table 8.1). Pseudoscorpiones From Mexico have been recorded 161 spp. of Pseudoscorpiones of 63 genera and 18 families (Beier 1933; Ceballos 2004; Chamberlin 1947; Hentschel and Muchmore 1989; Hoff 1945; Muchmore 1973b, 1975, 1977, 1982, 1986). Genera endemic for Mexico are: Mexichthonius Muchmore, 1975 – Campeche (one sp.) Apohya Muchmore, 1973b (one sp.) Vachonium J.C. Chamberlin, 1947 – Mexico, Belize (eight spp.) Mexichelifer Muchmore, 1973b (one sp.) Ancalochernes Beier, 1932 (one sp.) Epichernes Muchmore, 1982 (one sp.) Opiliones The outline of Kury and Cokendolpher (2000) presents the overall picture of the opilionid fauna of Mexico and a review of the biogeographic affinities of the genera (we are relying on them in our analysis). In this vast country, situated in the “zona de transition” between the Nearctic and the Palearctic, have been recorded 283 spp. (1 sp. of Cyphophthalmi, 143 Laniatores, and 139 “Palpatores”), of which 56 remained in 2000 unnamed. Further data could be found in the papers of Cokendolpher and Lee (1993); Goodnight and Goodnight (1942, 1945, 1951, 1953, 1971, 1977); Giribet (2011), Kury (2003, 2013), Morales Soto (1980), Roewer (1912, 1947), Shear (1977, 1980, 2006, 2010); Šilhavý (1974, 1977), and others. Cyphophthalmi The only Cyphophthalmid recorded from Mexico is Neogovea mexasca Shear, 1977 (endemic, troglobitic), with Neotropical affinities, now transferred to a new genus Shearogovea Giribet, 2011, with uncertain position. 8.4 Mexico: The Great American Interchange of Fauna Table 8.1 Scorpions in Mexico and the USA Country Меxico Number of species 272 Taxa Fam. Buthidae + Centruroides Marx 40 Chaneke Francke et al. 2 Tityus C.L. Koch − Fam. Caraboctonidae + Hadrurus Thorell 6 Hoffmannihadrurus Fet et Soleglad 2 Fam. Chactidae − Anuroctonus Pocock 2 Nullibroteas Williams 1 Uroctonus Thorell − Fam. Euscorpiidae + Megacormus Karsch 4 Plesiochactas Pocock 2 Troglocormus Francke 2 Fam. Scorpionidae + Bioculus Stahnke 5 Diplocentrus Peters 48 Koloti Santiban-Lopez et al. 2 Fam. Superstitioniidae + Superstitionia Stahnke 1 Fam. Typhlochactidae + Alacran Francke 3 Sotanochactas Francke 1 Stygochactas Vignoli et Prendini 1 Typhlochactas Mitchell 6 Fam. Vaejovidae + Balsateres G.-Sant. et Prendini 1 Chihuahuanus G.-Sant. et Prendini 8 Franckeus Soleglad et Fen 6 Gerschius Graham et Soleglad 2 Kochius Soleglad et Fen 9 Konetontli G.-Sant. et Prendini 5 Kovarikia Soleglad, Fen et Graham − Kuarapu Francke et P.S. 1 Maaykuyak G.-Sant. et Prendini 2 Mesomexovis G.-Sant. et Prendini 6 Paravaejovis Williams 12 Paruroctonus Werner 15 Pseudouroctonus Stahnke 9 Serradigitus Stahnke 14 633 USA 111 + − − 1 + 4 − + − − 3 − − − 1 + − 5 − + 1 − − − − − + − 4 − 1 2 − 3 − 1 1 4 22 9 6 (continued) 634 8 Table 8.1 (continued) Some Peculiarities of the Distribution of Arachnida Country Smeringurus Haradon Stahnkeus Soleglad et Fen Syntropis Kraepelin Thorellius Soleglad et Fen Uroctonites Williams et Savary Vaejovis C.L. Koch Vejovoidus Stahnke Viscaino G.-Sant. et Prendini Wernerius Soleglad et Fen Меxico 2 4 3 3 − 39 1 1 − USA − 4 − − 4 20 − − 3 The 143 spp. of Mexican “Palpatores” are distributed as follows: Eupnoi Fam. Sclerosomatidae Subfam. Gagrellinae Geaya Roewer, 1910 – Ten endemic spp., the genus with Neotropical affinities Holcobunus Roewer, 1910 – One endemic species, genus with Nearctic affinities (Eastern USA) Krusa G. et G., 1947 – Eleven endemic spp. (partly undescribed), Neotropical affinities Parageaya Mello-Leitão – Three endemic spp. (genus known from Uruguay and Argentina) Prionostemma Pocock, 1903 – Fifteen spp. (endemic and known from Central America) Romerella G. et G., 1943 – Two endemic spp. (Neotropical affinities) Trachyrhinus Weed, 1892 – Two spp. with Nearctic affinities (known also from the USA) Subfam. Leiobuninae Cosmobunus G.et G., 1946 – One sp. (endemic, Neotropical affinities, misidentified, acc. to Kury and Cokendolpher 2000) Hadrobunus Banks, 1900 – Two endemic spp. (Nearctic affinities, Eastern USA) Leiobunum C.L. Koch, 1839 – Thirty-one described and 7 undescribed spp., including 29 endemics (Holarctic, mostly temperate) Leuronychus Banks, 1900 – Two spp. (one endemic) (Nearctic affinities) Nelima Roewer, 1910 – Three endemic spp. (Holarctic, mostly temperate) Paranelima di Caporiacco, 1938 – Five endemic spp. (Neotropical affinities, Central America) 8.4 Mexico: The Great American Interchange of Fauna 635 Undescribed subfamily Dalquestia Cokendolpher, 1984 – Four spp. (three endemic) (Nearctic affinities) Diguetinus Roewer, 1912 – Two endemic spp., endemic genus Eurybunus Banks, 1893 – One described and five undescribed spp. (Nearctic affinities) Globipes Banks, 1893 – Two endemic spp. (Nearctic affinities) Metopilio Roewer, 1911 – Fourteen described and six undescribed spp. (Mixed affinities: Southwestern USA, Central America) Fam. Protolophidae Protolophus Banks, 1893 – Two endemic spp. (genus with Nearctic affinities, Western USA) Dyspnoi Fam. Acropsopilionidae Acropsopilio Silvestri, 1904 – A. chomulae Goodnight et Goodnight, 1948 (endemic) Fam. Nemastomatidae Subfam. Ortholasmatinae Martensolasma Shear, 2006 (endemic genus) Ortholasma Banks, 1894 – Several spp. from Mexico, partly undescribed (USA, Mexico, Honduras) Laniatores The 143 spp. of Mexican Laniatores are distributed as follows: Fam. Cosmetidae Acromares G. et G., 1942 – One endemic sp. (Neotropical affinities) Boneta G. et G., 1944 – One endemic sp. (Neotropical affinities) Colima G. et G., 1945 – One sp. (endemic genus) Cynorta C.L. Koch, 1839 – Nineteen end. species (Neotropical affinities) Cynortoides G. et G., 1945 – One end. sp. (Neotropical affinities) Cynortula Roewer, 1912 – Two end. spp. (Neotropical affinities) Erginiperna Roewer, 1947 – One end. sp. (endemic genus) Erginulus Roewer, 1912 – Twelve spp., including five end. (Neotropical affinities) Eucynorta Roewer, 1912 – Two end. spp. (Neotropical affinities) Eucynortoides Roewer, 1912 – One end. sp. (Neotropical affinities) Eucynortula Roewer, 1912 – One end. sp. (Neotropical affinities) Gueroma G. et G., 1942 – One end. sp. (Neotropical affinities) 636 8 Some Peculiarities of the Distribution of Arachnida Heterovonones Roewer, 1912 – One end. sp. (Neotropical affinities, Cuba) Holovonones Roewer, 1912 – One sp. (Neotropical affinities) Kevonones Chamberlin, 1925 – One end. sp. (Neotropical affinities) Metaconomma P.-C., 1905 – One end. sp. (endemic genus) Metacynorta P.-C., 1904 – Two spp. (one end.) (Neotropical affinities) Metavonones P.-C., 1904 – One end. sp. (Neotropical affinities) Meterginus P.-C., 1905 – Three spp.(one end.) (Neotropical affinities) Michella G. et G., 1942 – One end. sp. (Neotropical affinities) Paecilaema C.L. Koch, 1839 – Three spp., two end. (Neotropical affinities) ?Palpinus – One sp. recorded, unrecognizable Paravonones P.-C., 1904 – Three end. spp. (Neotropical affinities) Poala G. et G., 1942 – Two end. spp. (Neotropical affinities) Prosontes G. et G., 1945 – One end. sp. (endemic genus) Vononana Roewer, 1927 – One end. sp. (Neotropical affinities) Vonones Simon, 1879 – Seven spp. (six end.) (mixed affinities, Caribbean, Ecuador, USA) Fam. Phalangodidae – Three spp. Guerrobunus Goodnight et Goodnight, 1945 – Three spp. (? misplaced) Fam. Samoidae – Three spp. Akdalima Šilhavý, 1977 – One end. sp. (Neotropical affinities, Haiti) Arganotus Šilhavý, 1977 – One end. sp. (Neotropical affinities, Guatemala, Haiti) Pellobunus Banks, 1905 – One end. sp. (Neotropical affinities) Fam. Stygnommatidae – Eight spp. Stygnomma Roewer, 1912 – Eight spp. (seven end., one in Belize) (Neotropical affinities) Fam. Stygnopsidae – Forty-nine spp. Hoplobunus Banks, 1900 – Eleven described spp., 9 undescribed spp. (19 end.) (Nearctic affinities, Texas) Karos Goodnight et Goodnight, 1944 – Eleven described spp., seven undescribed spp. (endemic genus) Mexotroglinus Šilhavý, 1977 – One sp. (endemic genus) Paramitraceras O.P.-Cambridge, 1905 – Three spp., one endemic sp. (Neotropical affinities) Sbordonia Šilhavý, 1977 – Two spp. (endemic genus) Stygnopsis Sørensen, 1902 – Two end. spp. (endemic genus) Tampiconus Roewer, 1949 – One sp. (endemic genus) Troglostygnopsis Šilhavý, 1974 – Two spp. (endemic genus) Fam. Zalmoxidae – Four spp. Ethobunus Chamberlin, 1925 – Two spp. (one endemic) (Neotropical affinities) 8.4 Mexico: The Great American Interchange of Fauna 637 Metapachylus O. Pickard-Cambridge, 1905 – One sp. (endemic genus) Pachylicus Roewer, 1923 – One sp. (Neotropical affinities) The transitional position of Mexico reflects in its fauna of Opiliones. The analysis was made by Kury and Cokendolpher (2000), and we shall borrow from it. For Cyphophthalmi the endemic Shearogovea mexasca (Shear 1977) is clearly a Gondwanan Neotropical element, member of the Afro-Neotropical family Neogoveidae. “The occurrence of the endemic genus Guerrobunus in Mexico represents the southernmost limit of occurrence of true Phalangodidae in the New World. This family is typically distributed in the Holarctic Region and has 79 species recorded in the USA. This is the single example of penetration of typical Nearctic laniatorid fauna into Mexico. The other two genera of Laniatores which reach the Southern USA are Vonones and Hoplobunus, both typical members of the Neotropical opiliofauna, their maximum northern extension being here. Of the other 40 laniatorid genera, all have closest relations with the Neotropical fauna, including the 16 endemic genera.” About half of the “Palpatores” have close relations with the Nearctic and half with the Neotropical fauna. Acropsopilio is considered to be a relictual “Palpatores” genus. Since this analysis, were published the catalogue of Kury (2003) and the papers of Shear (2006) and other authors, and the number of identified opilionid species in Mexico was augmented. Actually, in seven of the northern states of Mexico are known only two to five species of Opiliones, and these states are surely understudied. A different situation is observed with Nuevo Leon (17 spp.), Tamaulipas (34 spp.), Guerrero (30 spp.), San Luis Potosi (26 spp.), and Veracruz (50 spp.), the highest number being in Chiapas (53 spp.). Amblypygi In Mexico are represented 3 genera with 19 spp. of 1 family (Armas 1994, 1995, 2006; Armas and Gadar 2004; Bilimek 1867; Butler 1873; Gervais 1842; Mullinex 1975, 1979; Pocock 1894, 1902; Quintero 1981; Rowland 1973; Wood 1893). Fam. Phrynidae Subfam. Phryninae Acanthophrynus Kraepelin, 1899 – Mexico, USA (Arizona, California) (one sp.) Paraphrynus Moreno, 1940 – (16 spp., 11 Mexican, all endemics) Five known from the USA, Central America, and West Indies Phrynus Lamarck, 1801 – Twenty-seven American spp., in Mexico are known (seven spp., four endemic); the others known also from South America, USA (Texas, Florida), Central America, and West Indies. The family Phrynidae is clearly Neotropical, despite the finding of one species in Indonesia (Flores). Thelyphonida (Uropygi) From Mexico has been recorded only one member of Mastigoproctus Pocock, 1894 – M. giganteus (Lucas, 1835) (Thelyphonidae, 638 8 Some Peculiarities of the Distribution of Arachnida Uroproctinae). The genus is known from South and Central America, the West Indies, and the Southern USA (Texas) (Harvey 2003). The localities in Mexico and Texas mark the northern limit of Uropygi in the New World. Schizomida Mexico is very rich in Schizomida, especially in caves (Cokendolpher and Reddell 1988; Rowland 1971a, b, 1973a, 1973b; Rowland and Reddell 1977; Reddell and Cokendolpher 1991, 1995). So far 41 species have been recorded, from the genera Mayazomus, Pacal, Stenochrus, Protoschizomus, Agastoschizomus, and Sotanostenochrus (all genera, except Stenochrus, are endemic to Mexico). Practically endemic is the family Protoschizomidae (known also in southern Texas). This fauna is different from the fauna of the neighboring regions (Central America and the Southern USA), largely because of the cave endemism. Actually, Mayazomus with its seven species from Chiapas and Tabasco, is also more Central American than connected to the rest of Mexico, so are two of the three species of Pacal Reddell et Cokendolpher. As a whole, Mexico has the third most numerous list of Schizomida in the world (after Australia and Cuba). Araneae According to Hoffmann (1976), the rich spider fauna of Mexico consists of three suborders, Orthognata, Labidognatha, and Cribellatae, and 50 families. Now (2017) this number is greatly increased. According to Francke (2011) and Aguayo-Morales et al. (2012), the Mexican spider fauna consists of 2295 spp. in 423 genera and 64 families. They all belong to contemporary Araneomorphae (Orthognatha). Opilioacarida The modern research on Mexican Opilioacarida by Vásquez and Klompen (2002, 2004, 2009) revealed the existence in this country of at least five species, all of the genus Neocarus Chamberlin et Mulaik. They have been collected in Baja California Sur (N. bajacalifornicus), Jalisco (N. bajacalifornicus chamelaensis), in Veracruz (N. veracruzensis), and in Campeche and Quintana Roo (N. calakmulensis, N. nohbecanus, N. siankaanensis). The other members of Neocarus are known from the neighboring USA (Texas, Arizona), Nicaragua, Cuba, and South America. Acariformes Trombidiformes – Prostigmata Fam. Erythraeidae Endemic genus: Tepotztlana Hoffman et Mendez, 1973 Parasitiformes Ixodida Fam. Argasidae – Nothoaspis reddelli Keirans et Clifford, 1975 (endemic genus and species, Yucatan). 8.5 Arachnida of the Antilleans and Mesoamerica 8.5 639 Arachnida of the Antilleans and Mesoamerica 8.5.1 Central America 8.5.1.1 Geography, General Zoogeography, and Paleogeography Central America is extending from the Isthmus of Tehuantepec in southern Mexico southeastward to the Isthmus of Panama where it connects to the Colombian Pacific Lowlands in northwestern South America. Alternatively, the Trans-Mexican Volcanic Belt delimits the region on the north. Central America has an area of some 592,000 square kilometers, shared between seven states, Belize, Guatemala, Honduras, Costa Rica, Nicaragua, El Salvador, and Panama, but the Mexican Yucatan is also part of it. Most of Central America rests atop the Caribbean Plate (Halffter 1976, 1987; Morrone 2001; Khudoley and Meyerhoff 1971; Ryan). Central America has been assigned to different zoogeographical units (Kraus 1960, 1962) but is usually considered part of the Neotropical Region. Using the comparative chorological method (De Lattin 1957), Müller (1972) outlined in Central America (Yucatan included) seven centers of dispersal: Central American Rainforest, Central American Montane Forest, Yucatan, Central American Pacific, Coco, Costa Rica, and Talamanca-Paramo. According to Morrone (2005), Central America is included in the Caribbean Subregion of the Neotropical Region (Mesoamerican Dominion with five provinces). Central America is part of the Mesoamerican Biodiversity hot spot, with 7% of the world’s biodiversity. As a bridge between North and South America, Central America has many species from the Nearctic and the Neotropic ecozones. Halffter (1978) says that his “Patron Mesoamericano de Montaña” (Mountain Mesoamerican Pattern) is composed of elements having evolved in the Central American Nucleus (determined by Halffter 1978, as “La mesa Central y las dos cordilleras de Chiapas, asi como las tierras altas y sus declives de Guatemala, Honduras, El Salvador y norte de Nicaragua”). The Entomologist Krzhanovskiy (2002) outlined the fauna of Central American Subregion and included in it the “Galapagos Province.” In Central America (from the Isthmus of Tehuantepec to Panama), we can distinguish clearly two mountain massifs: north of the Central American Nucleus, including 80% of the high grounds (above 600 m.) of Central America and in the south la Sierra de Talamanca in Costa Rica and the west of Panama (Halffter 1978). 640 8 Some Peculiarities of the Distribution of Arachnida Arachnogeography Central America, bridge between North and South America and facing the Caribbean, is zoogeographically important and is with rich and varied nature (Armas 2004). In the seven countries between Mexico and Colombia are represented 12 orders: Ricinulei (18 spp. of 2 genera), Solifugae (12 spp. of 5 genera, all Ammotrechidae), Scorpiones (6 fam., 11 genera – none is endemic), Pseudoscorpiones (13 fam., 33 gen., 1 endemic), Opiliones (1 genus of Eupnoi, 56 genera of 8 families of Laniatores, mostly Cosmetidae), Amblypygi (three genera of the families Phrynidae and Charinidae, four endemic spp.), Uropygi (three spp. of two genera, one is endemic), Araneae (many), Opilioacarida (two genera), plus the superorders Acariformes and Parasitiformes (Banks 1909a; Kury 2003; Roewer 1954). Not recorded in Central America are Palpigradi and Holothyrida and the suborders Palaeoamblypygi, Cyphophthalmi, Dyspnoi, and Mesothelae. If we consider the entire area of Central America (incl. Yucatan) and the Caribbean, we may say that this is the most varied area in Arachnida in the world – all orders and almost all suborders are present (except for Palaeoamblypygi and Mesothelae). Palpigradi – Not recorded from Central America Ricinulei In the 7 countries of Central America (without South Mexico) are registered 18 species of Ricinulei (Cooke and Shadab 1973; Fage 1921, 1938; Merret 1960; Platnick and Pass 1982; Platnick and Shadab 1981; Chamberlin and Ivie 1938). Cryptocellus Westwood, 1874 – Panama, Costa Rica, Honduras, Nicaragua (12 spp.) Pseudocellus Platnick, 1980 – Guatemala, Honduras, El Salvador, Panama (six spp.) These are the two genera of Ricinulei (Ricinoididae), known in the New World. They are represented also in Mexico and the Caribbean (Cuba). In Yucatan, which is geographically also part of Central America, are known five more Ricinuleid species. Solifugae From the 7 countries of Central America (without South Mexico) are known 12 spp. of Solifugae of 5 genera, all belonging to Ammotrechidae family (Armas 1993, 1996, 2000, 2004, with supplement; Muma 1970, 1976, 1986; Pocock 1902; Roewer 1934). The highest diversity has been recorded in Nicaragua (six spp.) and Guatemala (five spp.). According to Armas (1996), this is only half of the presumed number of species in Central America (23). Ammotrechinae Ammotrecha Banks, 1900 Ammotrecha limbata (Lucas, 1835) – Guatemala A. nigrescens Roewer, 1934 – Costa Rica, Guatemala A. picta Pocock, 1902 – Guatemala A. stollii (Pocock, 1895) – Costa Rica, Guatemala, USA Ammotrechella Roewer, 1934 8.5 Arachnida of the Antilleans and Mesoamerica 641 Ammotrechella pseustes (Chamberlin, 1925) – Panama, California, Puerto Rico tabogana Chamberlin, 1919 – Panama Ammotrechesta Roewer, 1934 Ammotrechesta garcetei Armas, 1993 – Nicaragua maesi Armas, 1993 – Nicaragua schlueteri Roewer, 1934 – Honduras brunnea Roewer, 1934 – Costa Rica tuzi Armas, 2000 – Mexico Innesa Roewer, 1934 Innesa vittata (Pocock, 1902) – Guatemala Scorpiones The list of scorpions in Central America (Mexico excluded) of Armas and Maes (1998) contains data on 38 species of 11 genera and 5 families. From them 24 species are Central American endemics. Meanwhile, some changes occurred in the names of families, the place of the genera, etc. Now there are 11 genera but of 6 families (Francke 1978; Francke and Stockwell 1987; Stockwell 1988; Lourenço 1996c; Lourenço and Méndez 1984; Teruel and Stockwell 2002; Viquez 1999) (Table 8.2). Pseudoscorpiones In the 7 countries of Central America are recorded pseudoscorpions of 33 genera and 13 families (Chthoniidae, Tridenchthoniidae, Bochicidae, Neobisiidae, Syarinidae, Ideoroncidae, Sternophoridae, Garypinidae, Atemnidae, Olpiidae, Cheliferidae, Chernetidae, Withiidae) (Beier 1931, 1932, 1953, 1955, 1976; Heurtault 1998; Hoff 1944; Mahnert 1987; Muchmore 1973). In the six countries of Central America (no data for Honduras), the number of the pseudoscorpion species is as follows (Harvey 2003): Belize, 10; Guatemala, 17; Costa Rica, 23; El Salvador, 10; Nicaragua, 1; and Panama, 15. Endemic genus for Central America: Fam. Chernetidae Coprochernes Beier – Costa Rica Opiliones Many authors, Cokendolpher and Cokendolpher (1984), Goodnight and Goodnight (1983), Kury (2003), Cruz-López et al. (2016), Pickard-Cambridge (1904–1905), Roewer (1949a), Shear (2010), Šilhavý (1979), Townsend et al. (2010), and others, have explored the rich opilionid fauna of Central America. Cyphophthami – Unknown Eupnoi Fam. Sclerosomatidae Geaya Roewer, 1910 – Belize, Costa Rica Prionostemma Pocock, 1903 – Guatemala, El Salvador, Panama, Costa Rica Dyspnoi Fam. Nemastomatidae 642 Table 8.2 Scorpiones in Central America (Without Mexico) Guat. 16 + − 6 − − − − − + − + 1 + 6 − + 1 Honduras 9 + − 5 − − − − − + − − − + 3 1 − − Salvador 2 + − 1 − − − − − − − − − + − 1 − − Nic. Costa Rica 4 15 + − 1 3 4 − 1 − 6 − + − − 1 − − − − − + − 1 + + − − 2 1 − − − − Panama 17 + 1 4 1 10 + 1 2 − − + 1 − − − + 1 Central America 8 + 1 10 1 8 + 1 2 + + 2 + 8 + 1 Congo DR Congo Gabon Centr. AR Angola 18 14 9 4 24 + + + + + 10 Some Peculiarities of the Distribution of Arachnida Belize 4 + − 1 − − − − − − − − − + 4 − − − 8 Country Number of species Fam. Buthidae Ananteris Thorell Centruroides Marx Isometrus Ehrenberg Tityus C.L. Koch Fam. Chactidae Neochactas Sol. et Fet Chactas Gervais Fam. Euscorpiidae Plesiochactas Pocock Fam. Hormuridae Opisthacanthus Peters Fam. Scorpionidae Diplocentrus Peters Didymocentrus Kraep. Fam. Vaejovidae Vaejovis C.L. Koch 8.5 Arachnida of the Antilleans and Mesoamerica 643 Ortholasmatinae Trilasma Goodnight et Goodnight – Honduras (one sp.) Laniatores The catalogue of Kury (2003) enumerates 198 spp. of Laniatores from the 7 countries of Central America of 56 genera and 8 families as follows: Fam. Cosmetidae Acromares Goodnight et Goodnight – Belize (one sp.) Bokwina G.et G. – Belize (one sp.) Boneta G. et G. – Guatemala (one sp.) Cosmetus Perty – Panama (one sp.) Cynorta C.L. Koch – Belize, Costa Rica, Panama, Honduras, Guatemala, El Salvador (20 spp.) Cynortellana Roewer – Costa Rica (one sp.) Cynortoperna Roewer – Costa Rica (one sp.) Cynortula Roewer – Costa Rica, El Salvador, Nicaragua, Guatemala (nine spp.) Erginoides Pickard – Cambridge-Panama, Costa Rica (two spp.) Erginulus Roewer – Guatemala, Honduras, Belize, Costa Rica, El Salvador (21 spp.) Eucynorta Roewer – Costa Rica, Panama, El Salvador, Guatemala (24 spp.) Eucynortella Roewer – Guatemala, Panama (five spp.) Eucynortoides Roewer – Costa Rica (one sp.) Eucynortula Roewer – Costa Rica, Nicaragua, Guatemala, Belize, Panama (seven spp.) Eugnidia Roewer – Costa Rica (one sp.) Eupoecilaema Roewer – Costa Rica, Panama (two spp.) Flirtea C.L. Koch – Honduras, Panama, Costa Rica (two spp.) Holovonones Roewer – Belize, Costa Rica, Guatemala (one sp.) Kevonones Chamberlin – Costa Rica (one sp.) Metacynorta Pickard – Cambridge-Guatemala (one sp.) Metarhaucus Pickard – Cambridge-Costa Rica (one sp.) Metavonones Pickard – Cambridge-Costa Rica (two spp.) Meterginus Pickard – Cambridge-Guatemala, Costa Rica, El Salvador (six spp.) Paecilaema C.L. Koch – Costa Rica, Panama, Guatemala, Honduras, Belize (12 spp.) Paecilaemana Roewer – Panama, Costa Rica (two spp.) Paracynorta G. et G. – Panama (one sp.) Paravonones Pickard – Cambridge-El Salvador (one sp.) Poecilaemula Roewer – Costa Rica (one sp.) Reimoserius Roewer – Costa Rica (one sp.) Tajumulcia G. et G. – Guatemala (one sp.) Vonones Roewer – Belize (one sp.) Vononesta Roewer – Guatemala (one sp.) Vononula Roewer – Guatemala (one sp.) 644 8 Some Peculiarities of the Distribution of Arachnida According to Townsend et al. (2010), the family Cosmetidae in Central America registered 133 spp. of 33 genera. Fam. Cranaidae Comboyus Roewer – Panama (one sp.) Phareicranaus Roewer – Costa Rica, Panama (two spp.) Fam. Gonyleptidae Glysterus Roewer – Costa Rica (six spp.) Hernandaria Soerensen – Costa Rica (two spp.) Hernandarioides Picard – Cambridge-Panama (one sp.) Nesopachylus Chamberlin – Panama (two spp.) Fam. Manaosbiidae Barrona Goodnight C. J. et M. L. Goodnight – Panama (one sp.) Bugabitia Roewer – Panama (one sp.) Poassa Roewer – Costa Rica (one sp.) Zygopachylus Chamberlin – Panama (one sp.) Fam. Samoidae Arganotus Šilhavý – Guatemala (one sp.) Neocynortina Goodnight C. J. et M. L. Goodnight – Costa Rica (one sp.) Pellobunus Banks – Panama (two spp.) Fam. Stygnommatidae Stygnomma Roewer – Belize, Costa Rica (six spp.) Fam. Pyramidopidae Jarmilana Cruz-López et al. – Belize (one sp., troglobite) Fam. Stygnopsidae Paramitraceras P. – Cambridge-El Salvador, Guatemala, Costa Rica (two spp.) Fam. Zalmoxidae Ethobunus Chamberlin – Panama, Guatemala, El Salvador, Costa Rica (15 spp.) Pachylicus Roewer – Panama, Costa Rica (eight spp.) Panopiliops Roewer – Costa Rica (two spp.) Phalangoduna Roewer – Costa Rica, Panama (one sp.) Stygnoleptes Banks – Costa Rica, Panama, El Salvador (three spp.) 8.5 Arachnida of the Antilleans and Mesoamerica 645 Family uncertain Costabrimma Goodnight et Goodnight – Costa Rica (three spp.) Isaeolus Roewer – El Salvador (one sp.) Amblypygi The amblypygids are represented in Central America by the family Phrynidae and the genera Phrynus Lamarck (four spp. in Nicaragua, Guatemala, Honduras, Belize, Costa Rica, Panama) and Paraphrynus Moreno (four spp. in Belize, Guatemala, Costa Rica, El Salvador, Nicaragua, Panama, Honduras). Both genera live also in South America, Mexico, and the Antilleans. The genus Charinus (fam. Charinidae) is found in Panama (Armas 2004; Armas de and Gonzalez 2001; Armas de and Maes 2000; Armas de and Víquez 2001; Avila Calvo and Armas 1997; Gervais 1842; Mullinex 1975, 1979; Pocock 1893, 1894, 1902; Roewer 1954; Quintero 1981; Víquez et al. 2012). There are four species endemic to Central America (between Mexico and Colombia): Phrynus Lamarck – Ph. maesi Armas, 1996 (Nicaragua), Ph. parvulus Pocock, 1902 (Guatemala, Belize) Paraphrynus Moreno – P. emaciatus Mullinex, 1975 (Guatemala), P. leptus Mullinex, 1975 (Guatemala) Thelyphonida (Uropygi) In Central America are found five species of Uropygi and three endemic genera Valeriophonus (Viquez et de Armas, 2005, Mayacentrum Viquez et Armas, 2006, and Mimoscorpius Butler, 1872). They live in Costa Rica (Valeriophonus nara Valerio), El Salvador (Mayacentrum tantalus Roewer), Honduras (M. pijol Viquez et de Armas), Belize (M. guatemalae Viquez et de Armas), and Guatemala (Mimoscorpius pugnator Butler) (Roewer 1954; Valerio 1981; Viquez and de Armas 2005, 2006; Armas and Viquez 2007). Schizomida In Central America (between Mexico and Colombia) are recorded nine species, but there is also unidentified material from many places (Armas et al. 2010; Armas and Viquez 2010; Brignoli 1973; Rowland and Reddell 1977; Reddell and Cokendolpher 1995). Besides the largely distributed Stenochrus portoricensis (Guatemala, Honduras, Nicaragua, Belize), from Central America are known the genera Heteronochrus (Guatemala), Hansenochrus (Costa Rica, Panama), Rowlandius, Piaroa and Surazomus (Costa Rica), and Belicenochrus from Belize. Surazomus Reddell et Cokendolpher is known also from many South American countries. Rowlandius Reddell et Cokendolpher is widespread in the Caribbean Islands. Hansenochrus Reddell et Cokendolpher is found also in South America and the Caribbean. 646 8 Some Peculiarities of the Distribution of Arachnida Map 8.2 Schizomida in Central America and the Caribbean Belicenochrus – ◊ Stenochrus – ■ Hansenochrus – ▲ Rowlandius – ▼ Surazomus – ●` Stewardpeckius – ♣ Antillostenochrus – Piaroa – Luisarmasius – Cokendolpherius – Cubacanthozomus – Heterocubazomus – Caribezomus – ► Heteronochrus – ◄ Cubazomus – X Guanazomus – □ Reddellzomus – ♦ Troglocubanus – ○ Araneae In Central America (without Mexico) are known at leat 59 families of spiders: Microstigmatidae, Dipluridae, Nemesiidae, Theraphosidae, Paratropidae, Barychelidae, Cyrtaucheniidae, Idiopidae, Ctenizidae, and Actinopodidae from infraorder Mygalomorphae and Filistatidae, Drymusidae, Scytodidae, Sicariidae, 8.5 Arachnida of the Antilleans and Mesoamerica 647 Leptonetidae, Ochyroceratidae, Telemidae, Pholcidae, Caponiidae, Tetrablemmidae, Oonopidae, Segestriidae, Hersiliidae, Oecobiidae, Mimetidae, Deinopidae, Uloboridae, Anapidae, Araneidae, Linyphiidae, Synphytognatidae, Tetragnathidae, Theridiidae, Theridiosomatidae, Ctenidae, Lycosidae, Oxyopidae, Pisauridae, Senoculidae, Trechaleidae, Udubidae, Agelenidae, Amaurobiidae, Anyphaenidae, Dictynidae, Hahniidae, Sparassidae, Selenopidae, Zodariidae, Zoropsidae, Clubionidae, Miturgidae, Titanoecidae, Gnaphosidae, Prodidomidae, Philodromidae, Thomisidae, Salticidae, and Corinnidae from infraorder Araneomorphae. Most of the endemic genera have been described from Panama by Chikering (1946, Panama). No endemic families were noted. Some endemic spider genera in Central America and the Caribbean: Fam. Microstigmatidae Micromygale Platnick et Forster, 1982 – Panama Fam. Theraphosidae Aenigmarachne Schmidt, 2005 – Costa Rica Antillena Bertani, Huff et Fukushima, 2017 – Dominican Republic Barropelma Chamberlin, 1940 – Panama (one sp.) Caribena Fukushima et Bertani, 2017 – Puerto Rico, Cuba, US Virgin Islands, Martinique Crassicrus Reichling et West, 1996 – Belize Longilyra Gabriel, 2014 – El Salvador Mygalarachne Ausserer, 1871 – Honduras Reichlingeria Rudloff, 2001 – Belize Sphaerobothria Karsch, 1879 – Costa Rica, Panama Stichoplastoris Rudloff, 1997 – El Salvador, Costa Rica, Panama Fam. Ochyroceratidae Fageicera Dumitrescu et Georgescu, 1992 – Cuba Fam. Barychelidae Troglothele Fage, 1929 – Cuba Fam. Agelenidae Neowadotes Alayón, 1995 – Hispaniola Fam. Cyatholipidae Pokennips Griswold, 2001 – Jamaica Fam. Tetragnathidae Ancinosphenus Simon, 1895 – West Indies Antillognatha Bryant, 1945 – Hispaniola Hispanognatha Bryant, 1945 – Hispaniola 648 8 Some Peculiarities of the Distribution of Arachnida Fam. Linyphiidae Lomaita Bryant, 1948 – Hispaniola Primerigonina Wunderlich, 1995 – Panama Sthelota Simon, 1894 – Panama, Guatemala Fam. Filistatidae Antilloides Breskovit et al., 2016 – Cuba, Dominican Republic, Virgin Islands, Puerto Rico Fam. Paratropididae Anisaspis Simon, 1891 – Saint Vincent Fam. Amaurobiidae Tugana Chamberlin, 1948 – Cuba, Hispaniola Fam. Anyphaenidae Thaloe Brescovit, 1993 – Cuba, Hispaniola Fam. Theridiidae Jamaitidion Wunderlich, 1995 – Jamaica Fam. Liocranidae Laudetia Gertsch, 1941 – Dominican Republic, Puerto Rico Mesobria Simon, 1897 – Saint Vincent (one sp.) Fam. Ctenidae Ciba Bloom et al., 2014 – Cuba, Dominican Republic Ohvida Polotow et Brescovit, 2009 – Cuba, Puerto Rico, Bahamas Trujillina Bryant, 1948 – Hispaniola, Puerto Rico Fam. Pholcidae Bryantina Brignoli, 1985 – Cuba Ciboneya Perez, 2001 – Cuba Platnicknia Özdikmen et Demir, 2009 – Cuba Tainonia Huber, 2000 – Hispaniola Fam. Tetrablemmidae Micromatta Lehtinen, 1981 – Belize Fam. Miturgidae Hoedillus Simon, 1898 – Guatemala, Nicaragua 8.5 Arachnida of the Antilleans and Mesoamerica 649 Fam. Prodidomidae Caudalia Alayón, 1980 – Cuba Cubanopillus Alayón et Platnick, 1993 – Cuba Fam. Sparassidae Decaphora Franganillo, 1931 – Cuba Defectrix Petrunkevitch, 1925 – Panama Fam. Thomisidae Rejanellus Lise, 2005 – Cuba, Puerto Rico, Hispaniola Fam. Salticidae Albionella Chikering, 1946; Banksetosa Chickering, 1946; Carabella Chickering, 1946 Gorgasella Chickering, 1946; Micalula Strand, 1932; Monaga Chickering,1946 Orvilleus Chickering, 1946; Toloella Chickering, 1946; Udalmella Galiano, 1994 Uluella Chickering, 1946 – All from Panama Allodecta Bruyant, 1950 – Jamaica Antillattus Bryant, 1943 – Hispaniola, Cuba Bythocrotus Simon, 1903 – Hispaniola Caribattus Bryant, 1950 – Jamaica Cerionesta Simon, 1901 – Saint Vincent Commoris Simon, 1902 – Guadeloupe, Dominica Corticettus Zhang et Maddison, 2012 – Puerto Rico, Hispaniola Paraplexippus Franganillo, 1930 – Cuba Parasaitis Bruyant, 1950 – Jamaica Parathiodina Bruyant, 1943 – Hispaniola Popcornella Zhang et Maddison, 2012 – Hispaniola, Puerto Rico Truncattus Zhang et Maddison, 2012 – Hispaniola Opilioacarida From Central America (between Mexico and Colombia) are known two spp. of Opilioacarida, Caribeacarus panamensis Vásquez et Klompen (Panama) and Neocarus nicaraguensis (Vásquez et Klompen) (Nicaragua), but we can add three species of Neocarus Chamberlin et Mulaik, 1942, from Yucatan Peninsula in Mexico. Both Neocarus and Caribeacarus live also on the Antilleans. The species are (so far) endemic (Vásquez and Klompen 2002, 2009). Parasitiformes Ref.: Beron (2014), Chant and Baker (1965), Hoffmann (1962). 650 8 Some Peculiarities of the Distribution of Arachnida The purely Neotropical family of Spelaeorhynchidae (parasites on bats of the families Phyllostomatidae and Mormoopidae) is found, together with these bats, in Cuba, Jamaica, Puerto Rico, and Dominican Republic but also in Central America. Holothyrida – Unknown 8.6 8.6.1 Antillean (Caribbean) Islands Geography, General Zoogeography, and Paleogeography “The West Indies and Central America, the question of the relationships between Atlantic and Pacific Ocean, between North and South America, as well as the question of possible land-bridges between the islands on one hand and the continent on the other, have been dealt with in an almost interminable procession of biogeographical and geological publications.” Rutten (1935) Antillean islands form the greater part of the West Indies in the Caribbean. The Antilleans are divided into two major groups: the “Greater Antilles” to the north and west, including the larger islands of Cuba, Jamaica, Hispaniola (Haiti and the Dominican Republic), and Puerto Rico, and the smaller “Lesser Antilles” on the southeast – comprising the northerly Leeward Islands, the southeasterly Windward Islands, and the Leeward Antilles just north of Venezuela. The Bahamas, though part of the West Indies, are generally not included among the Antillean islands. Geologically, the Greater Antilles are made up of continental rock, as distinct from the Lesser Antilles, which are mostly young volcanic or coral islands. Cuba – area of 109,886 km2, highest point Pico Turquino (1974 m.) (Isla de la Juventud, 2199 km2). Remain only ca. 15% of the original forest cover. Hispaniola – Area 76 480 km2, highest point Pico Duarte (3087 m.) Jamaica – Area 10 990 km2, highest point Blue Mountain Peak (2256 m.) Puerto Rico – Area 9104 km2, highest point Cerro de Punta (1338 m.) The Lesser Antilles, also known as the Caribbees, are part of the Antilles, which together with the Bahamas, the Cayman Islands, the Turks and Caicos Islands, and Greater Antilleans form the West Indies. The islands are a long partly volcanic island arc, most of which wrap around the Eastern end of the Caribbean Sea on the Western boundary with the Atlantic Ocean. The two main groups of the Lesser Antilleans are the Windward Islands in the south and the Leeward Islands in the north. The Leeward Antilleans in the west form the third group; they consist of the Dutch islands just off the coast of Venezuela, plus a group of Venezuelan islands. The main Lesser Antilles are (from north to south to west): Leeward Islands: Virgin Islands, Anguilla (91 km2, Saint Martin (87 km2, 414 m – Pic Paradis), Saint-Barthélemy, Saba, Saint Eustatius, Saint Kitts, Nevis, Antigua, Redonda, Montserrat, Guadeloupe, Dominica 8.6 Antillean (Caribbean) Islands 651 Windward Islands: Martinique, Saint Lucia, Saint Vincent, Grenadines Barbados (a continental island found 160 km east of the Windward chain) – 430 km2, 336 m (Mount Hillaby) Grenada – 344 km2, 840 m (Mount Saint Catherine) Tobago – 300 km2 Trinidad – 4768 km2 Trinidad and Tobago are situated on the shelf of South America, 11 km far from the mainland. Dutch Antilles – Islands north of the Venezuelan coast (from west to east) Aruba – 193 km2, 188 m (Jamanota) Curaçao – 444 km2, 375 m (Christoffelberg) Bonaire – 294 km2, 240 m (Brandaris) Many papers deal with the paleogeography of the Caribbean. Among them are Blair Hedges (2001, 2006), Bonatti and Gardner (1973), Burke et al. (1984), Curtis et al. (2001), Freeland and Dietz (1971), Hedges (1982, 2001), Hedge et al. (1994), Iturralde-Vinent and MacPhee (1999), Jeannel (1939), Khudoley and Meyerhoff (1971), Koopman (1959), Meschede and Frish (1998, 2001), Meyerhoff and Meyerhoff (1972), Morrone (2001), Page and Lydeard (1994), Schuchert (1935), Weyl (1964), Woodring (1954), Wyatt Durham (1985), and others. “A major deformation began in the Miocene, with folding of the Lower Tertiary sediments and uplifting of the cores of the present-day islands. During the Late Miocene and Pliocene, extensive peneplains were formed, which, as a result of the youngest crustal movements in the Antilles, now stand at elevations as great as 2000 m” (Weyl 1966). “Sykes et al. have demonstrated that the Caribbean plate has moved eastnortheast about 1400 km since late Eocene time (38 Ma). This movement changes or affects the interpretation of many biogeographic problems of that region” (Wyatt Durham 1985). “In the Middle Eocene ends entirely the movement of the plate and Caribbean Sea and its surroundings acquired approximately their actual configuration” (Iturralde-Vinent 1975). The Antilles and the West Indies in broader sense (including the Bahamas Bank) have been interesting to biogeographers even since Wallace (1881). The Greater Antilles (Cuba, Hispaniola, Jamaica, and Puerto Rico) are large enough to allow radiative speciation within the islands and between them and even to disperse some lines to the mainland (some examples among the vertebrates have been quoted by Blair Hedges (2006)). This author made an analysis of the paleogeography of the Antilles and the identity of the 1319 native terrestrial vertebrates on the islands. The problem of how and when these animals arrived to the islands has been discussed (Darlington 1938; Weyl 1973; Morrone 2005, and many others). Among the non-flying terrestrial vertebrates, the endemicity is high: nearly all 173 spp. of amphibians, 96% of the 499 spp. of reptiles, 96% of the freshwater fishes, but only 35% of birds and 50% of bats. It would be interesting to compare these figures with the various orders of Arachnida (although with the vertebrates, we 652 8 Some Peculiarities of the Distribution of Arachnida have to deal with classes). Thanks to many local and foreign researchers, the general picture of the Arachnida on the Antilles is more or less complete and a sound basis for discussions. Actually, the main argument is centered on the importance (and feasibility) of the three ways of “arrival” of the animals: dispersal, vicariance, and land bridges. The age of the various groups is also important. There are few fossils of Arachnida on the islands, mostly from Dominican amber (Miocene 15–20 million years to Upper Eocene ca. 30–45 million years), so hypotheses should be based on speculations. Morrone (2001) includes the Antilleans as 7 of the 29 provinces of the Caribbean Subregion of Neotropical Region (together with Central America, most of Mexico, Ecuador, Galapagos, and other areas). According to Morrone (2005), the Antillean Dominion is divided into eight provinces: the Yucatan Peninsula province (the states of Yucatan, Campeche, and Quintana Roo), Bahamas province, Cuba province, the Cayman Islands province, Jamaica province, Hispaniola province, Puerto Rico province, and Lesser Antilles province. These provinces reflect the distribution of some orders of insects in the area. Morrone (2017) provided a detailed regionalization of the Neotropics, dividing the area in three subregions: Antillean, Brazilian, and Chacoan. Based on insects (mostly Coleoptera), Krzhanovskiy (2002) considered the Antillean Subregion part of the Neotropical Region. For him the explanation is simple: volcanic archipelago raised in the late Tertiary and populated by casual migrants from Central and South America. Some speciation took place after that. Arachnogeography The arachnofauna of the islands is varied and with many endemics, well studied by Cuban, Romanian, and other specialists (Armas 1982, 2004; Banks 1909b; PerezGelabert 2008; Teruel and de Armas 2005; Armas and Avila 2015). Represented are all orders. Remarkable are the findings of an endemic genus and species of Holothyrida, many endemic genera of Schizomida, and very rich fauna of scorpions (130 spp., only in Cuba there are 55 spp.). According to these authors, there is a zoogeographical barrier between Trinidad and Tobago on one side and Grenada on the other, “the bond line.” Lourenço (loc. cit.) indicates that the number of scorpion species doesn’t seem to be correlated negatively with the distance from South America but rather with the surface of the islands – five on Martinique, three on Saint Lucie, and one on each of the smaller islands. Lourenço (1999b) made an important analysis of the origins and the affinities of the scorpions of the Greater Antilleans. He indicates the Northern and Central American origins of some elements (Centruroides) and the “undoubtedly” Southern American origin of others (Ischnuridae = Hemiscorpiidae and other Buthidae). The explanation is the formation of a “cluster of islands,” known as Proto-Antilleans, which drifted to South America, allowing the passage of some elements from this continent. We may indicate that, studying the cave Isopoda of Cuba, Vandel (1973) stressed on their exclusively South American origin. 8.6 Antillean (Caribbean) Islands 653 Some attempts have been made to explain the connections of the present Greater Antilleans with North or Central America by the lowering by 100–150 m during the Pleistocene glaciations (Donn et al. 1962 and others) and the supposed “stepping stones” as a consequence. However, we find hard to explain why all this rich arachnofauna used these “stones” and many other taxa did not. In the analysis of Weyl (1964), no connection between the islands and any continental ground is shown after the probable “bridge” between Jamaica-Hispaniola and Central America (the so-called Sapperland) in the Paleocene (65–55 Ma ago). On the islands are lacking the suborders Cyphophthalmi, Dyspnoi, Palaeoamblypygi, and Mesothelae. Palpigradi The only troglobite Palpigradi in the New World (Eukoenenia orghidani Condé et Juberthie) was described from Cuba. Some non-described Palpigradi have been recorded from Guadeloupe and Haiti, and the subspecies Eukoenenia berlesei virginea Condé (1984) was described from the Virgin Islands (Condé 1984, 1986; Condé and Juberthie 1982; Remy 1948). Solifugae From the Antilleans are known three genera of Solifugae, two of them endemic, all belonging to Ammotrechidae (Armas 1994, 2004; Armas and Teruel 2005; Muma 1970, 1976, 1986; Muma and Nezario 1971), as follows: Ammotrechella apejii Muma, 1981 – Jamaica (endemic) bahamica Muma, 1986 – Bahamas (endemic) bonariensis (Werner, 1925) – Bonaire (endemic) [Ammotrechona] cubae (Lucas, 1835) – Cuba (endemic) geniculata (C.L. Koch, 1842) – Bahamas, Curacao, Saint Vincent, Guadeloupe, also USA, Mexico, and Panama jutisi Armas et Teruel, 2005 – Cuba (endemic) hispaniolana Armas et Alegre, 2001 – Dominican Republic (endemic) maguirei Muma, 1986 – Caicos (endemic) pallida Muma et Nezario, 1971 – Puerto Rico (endemic) pseustes (Chamberlin, 1925) – Puerto Rico, Panama Ammotrechinus gryllipes (Gervais, 1842) – Jamaica, Haiti (endemic genus and species) Antillotrecha iviei Armas, 2002 – Leeward Islands (endemic genus for the Caribbean) fraterna Armas, 1994 – Dominican Republic (endemic sp. for Dominican Rep.) disjunctodens Armas et Teruel, 2005 – Cuba (endemic sp.) Ammotrechella apejii Muma, 1981 – Jamaica (endemic) bahamica Muma, 1986 – Bahamas (endemic) bonariensis (Werner, 1925) – Bonaire (endemic) [Ammotrechona] cubae (Lucas, 1835) – Cuba (endemic) geniculata (C.L. Koch, 1842) – Bahamas, Curacao, Saint Vincent, Guadeloupe, also USA, Mexico, and Panama jutisi Armas et Teruel, 2005 – Cuba (endemic) 654 8 Some Peculiarities of the Distribution of Arachnida hispaniolana Armas et Alegre, 2001 – Dominican Republic (endemic) maguirei Muma, 1986 – Caicos (endemic) pallida Muma et Nezario, 1971 – Puerto Rico (endemic) pseustes (Chamberlin, 1925) – Puerto Rico, Panama Ammotrechinus gryllipes (Gervais, 1842) – Jamaica, Haiti (endemic genus and species) Antillotrecha iviei Armas, 2002 – Leeward Islands (endemic genus for the Caribbean) fraterna Armas, 1994 – Dominican Republic (endemic sp. for Dominican Rep.) disjunctodens Armas et Teruel, 2005 – Cuba (endemic sp.) Prevails the genus Ammotrechella, found also in Panama, Mexico, and the USA (Texas and Florida). In Cuba there are eight spp. of Solifugae and in Jamaica – two in Dominican Republic – three spp. (Armas 2004, with catalogue and analysis of the distribution of Solifugae in the Caribbean and Central America). Since only few taxa have been added (Armas and Teruel 2005 and others). Ricinulei The only Ricinuleids known in the Greater Antilleans were found in Cuba (one cave inhabiting Pseudocellus, described almost simultaneously by Cooke (1972) and by Dumitresco and Juvara-Bals (1973), and Pseudocellus silvai (Armas, 1977)). Both species seem endemic for Cuba. Judson and Hardy (2001) described a protonymph of Cryptocellus sp. from Tobago Island. Scorpiones The information on the scorpions in the Antillean fauna was generalized by Armas (2001, 2009b) and Teruel and Kovařik (2012), who described many of the existing taxa in the islands. Zoogeographical analysis of the origin and the affinities of the Antillean scorpions is by Lourenço (1999a). On the Antilleans have been recorded 4 families (Buthidae, Chactidae, Hormuridae, and Scorpionidae), 17 genera, and 130 species. Four genera with 21 species are endemic in the Antilleans (Armas 1973, 1974, 1982, 1983, 1999, 2001, 2009b; Armas and Maes 1998; Armas and Marcano 1987; Francke 1978; Francke and Sissom 1980; Kjellesvig-Waering 1966; Lourenço 1984, 1986, 1987; Lourenço et al. 1991; Lourenço and Huber 1999; Lourenço and Vachon 1996; Kovařik and Teruel 2014; Moreno 1940; Teruel 2005, and many others). The scorpions of the Antilleans belong to 17 genera of 4 families, but Chactidae (with 3 genera) is known only on the islands of Trinidad and Tobago near the coast of Venezuela (also the genus Ananteris). From the other 13 genera by far, the richest is the fauna of Cuba (55 spp.), followed of Hispaniola (Dominican Republic and Haiti), with a total of 41 spp. The list of the scorpions in the Lesser Antilleans (16 spp.) is provided by Lescure et al. (1991), the one for Pinos (Isla de la Juventud, three spp. Centruroides, Rhopalurus, no endemic) by Armas (1983). Family Buthidae Alayotityus Armas, 1973 (a Cuban endemism, seven species) 8.6 Antillean (Caribbean) Islands 655 Tityopsis Armas, 1974 (a Cuban endemism, two species) Family Scorpionidae (incl. Diplocentridae) Cazierius Francke, 1978 (a Greater Antillean endemism, ten species) Oiclus Simon, 1880 (a Lesser Antillean endemism, two species from the Leeward Islands) The other 11 genera are distributed as follows: Ananteris Borelli, 1910 (one species from Trinidad and Tobago) Centruroides Marx, 1890 (28 species on the Antilles) Isometrus Ehrenberg, 1828 (one introduced species) Microtityus Kjellesvig-Waering, 1966 (15 species, incl. 6 recent spp. from Hispaniola) Rhopalurus Thorell, 1876 (eight species from Cuba and Hispaniola) Tityus C. L. Koch, 1836 (20 species on the Antilles, incl. 13 recent spp. from Hispaniola) Broteochactas Pocock, 1893 (two species from Trinidad and Tobago) Chactas Gervais, 1844 (one species from Trinidad and Tobago) Opisthacanthus Peters, 1861 (one species from Hispaniola) Didymocentrus Kraepelin, 1905 (nine species from Lesser Antilles and central Cuba) Heteronebo Pocock, 1899 (14 species from Greater Antilles) (Table 8.3). Pseudoscorpiones The fauna of Pseudoscorpiones in the Caribbean belongs to 16 families as follows (indicated are only the species endemic for West Indies) (Barba Diaz and Pérez 2001; Barba Diaz and Barroso 2013; Beier 1976; Cosgrove et al. 2016; Dumitresco and Orghidan 1977, 1981; Harvey 2013b; Heurtault and Rebière 1983; Hoff 1945, 1946, 1959, 1963, 1964, 1976; Muchmore 1967, 1979, 1982, 1984, 1992, 1998; Tullgren 1907; Vitali-di-Castri 1984). According to the table in Cosgrove (2016), the genera are 74 and the species 147. Fam. Chthoniidae Caribchthonius butleri Muchmore, 1976 – US Virgin Islands (endemic) Pseudochthonius thibaudi Vitali-di-Castri, 1984 – Guadeloupe (endemic) Tyrannochthonius guadeloupensis Vitali-di-Castri, 1984 – Guadeloupe (endemic) T. imitatus Hoff, 1959 – Jamaica, Dominican Republic T. proximus Hoff, 1959 – Jamaica, Dominican Republic Fam. Lechythiidae Lechytia delamarei Vitali-di-Castri, 1984 – Guadeloupe (endemic) L. trinitatis Beier, 1970 – Dominican Republic, Trinidad 656 8 Some Peculiarities of the Distribution of Arachnida Table 8.3 Scorpions in the Antilleans Country Cuba Hispaniola (40 + 6) Number of species 55 41 Fam. Buthidae + + Alayotityus Armas 8 − Ananteris Thorell − − Centruroides Marx 14 5 Isometrus Ehrenberg Microtityus 9 12 K. – W. Rhopalurus 7 3 Thorell Tityopsis Pocock 2 − Tityus C.L. Koch 1 13 Fam. Chactidae − − Neochactas Sol. et − − Fet Chactas Gervais − − Broteochactas − − Pocock Fam. Hormuridae − + Opisthacanthus − 1 Peters Fam. + + Scorpionidae Casierus Francke 7 2 Didymocentrus 4 − Kraepelin Cryptoiclus Ter. et 1 − Kov. Heteronebo 2 5 Pocock Oiclus Simon − − Puerto Jamaica Rico Lesser Bahamas Antilles Trinidad and Tobago 6 + − − 4 17 + − − 5 2 − − 2 16 + − − 8 11 + − 1 1 − 2 − − 2 − 1 − − − − 1 − − − 7 − − − − − − − 4 − − − 4 + 1 − − − − − − − − 1 1 − − − − − − − − − − + + − + + − − 1 − − − 1 3 − − − − − − − 1 1 − 5 − − − − 3 − Fam. Tridenchthoniidae Tridenchthonius cubanus (Chamberlin, 1929) – Cuba Fam. Syarinidae Ideobisium balzanii With, 1905 – Guadeloupe, Dominica, Saint Vincent puertoricense Muchmore, 1982 – Dominican Republic, Puerto Rico puertoricense cavicola Muchmore, 1982 – Puerto Rico yunquense Muchmore, 1982 – Puerto Rico Ideoblothrus carinatus (Hoff, 1964) – Jamaica 8.6 Antillean (Caribbean) Islands 657 insularum (Hoff, 1945) – Jamaica, Puerto Rico pygmaeus (Hoff, 1964) – Jamaica, Martinique truncatus (Hoff, 1964) – Jamaica Fam. Bochicidae Antillobisium mitchelli Dumitresco et Orghidan, 1977 – Cuba (endemic genus) vachoni Dumitresco et Orghidan, 1977 – Cuba Troglobochica Muchmore, 1984 – Jamaica (endemic genus) T. jamaicensis Muchmore, 1984 – Jamaica T. pecki Muchmore, 1984 – Jamaica Mexobisium armasi Muchmore, 1980 – Cuba M. cubanum Muchmore, 1973 – Cuba M. sierramaestrae Muchmore, 1980 – Cuba Nannobisium mollis (Hoff, 1964) – Jamaica Fam. Ideoroncidae Typhloroncus coralensis Muchmore, 1979 – US Virgin Islands (endemic) (the other four Typhloroncus are from Mexico) Fam. Sternophoridae Fam. Olpiidae Aphelolpium thibaudi Heurtault et Rebière, 1893 – Guadeloupe, Martinique Novohorus subfuscus Hoff, 1945 – Jamaica, Puerto Rico Olpiolum monae (Hoff, 1964) – Mona, Jamaica, Dominican Republic, Florida O. amplum Hoff, 1964 – Jamaica O. aureum (Hoff, 1945) – Mona, Puerto Rico O. confundens (Hoff, 1945) – Puerto Rico O. puertoricensis (Hoff, 1945) – Puerto Rico Fam. Garypidae Planctolpium arboreum Hoff, 1964 – Jamaica, Dominican Republic, Mexico Fam. Garypinidae Fam. Cheiridiidae Cheiridium insulare Heurtault et Rebière, 1893 – Guadeloupe (endemic) Fam. Pseudochiridiidae Pseudochiridium insulae Hoff, 1964 – Cuba, Dominican Republic Fam. Atemnidae Paratemnoides elongatus (Banks, 1895) – Cuba, Dominican Republic, US Virgin Islands, Central America, Mexico, USA P. nidificator (Balzan, 1888) – Haiti, Saint Vincent, Central and South America 658 8 Some Peculiarities of the Distribution of Arachnida Fam. Cheliferidae Cubachelifer stator Hoff, 1946 – Dominican Republic, Cuba Parachelifer dominicanus Beier, 1976 – Dominican Republic (endemic) P. parvus Muchmore, 1981 – US Virgin Islands (endemic) Tyrannochelifer cubanus Hoff, 1964 – Cuba (endemic) T. floridanus (Banks, 1891) – Dominican Republic, Florida T. macropalpus (Tullgren, 1907) – Haiti (endemic) Fam. Chernetidae Americhernes puertoricensis Muchmore, 1976 – Puerto Rico Antillochernes cruzensis Muchmore, 1984 – US Virgin Islands (endemic) Bituberochernes jonensis Muchmore, 1979 – US Virgin Islands (endemic) Byrso.chernes caribicus Beier, 1976 – Dominican Republic (endemic) Caribochernes pumilus Beier, 1976 – Dominican Republic (endemic) Chelanops ? [Dinocheirus] altimanus (Ellingsen, 1910) – St. Thomas (endemic) Chernes hispaniolicus Beier, 1976 – Dominican Republic (end.) Dinochernes chalumeaui Heurtault et Rebière, 1893 – Guadeloupe (endemic) Hesperochernes vespertilionis Beier, 1976 – Dominican Republic (endemic) Gomphochernes communis (Balzan, 1888) – South America, Trinidad, Dominican Republic Lustrochernes mauriesi Heurtault et Rebière, 1893 – Guadeloupe (endemic) Parachernes (Scapanochernes) compressus (Tullgren, 1907) – Dominican Republic, Haiti, Florida P. dominicanus Beier, 1976 – Dominican Republic, Haiti (endemic) Parazaona klapperichi Beier, 1976 – Dominican Republic (endemic) Fam. Withiidae Cacodemonius segmentidentatus [in Beier, 1976 – serratidentatus] (Balzan, 1891): Dominican Republic, South America Dolichowithius canestrinii (Balzan, 1887) – Virgin Islands, St. Thomas, South America D. simplex Beier, 1932 – Dominican Republic, Puerto Rico Endemic genera for the Antilleans are: Antillobisium Dumitresco et Orghidan, 1977 – Cuba (two spp.) Caribochernes Beier, 1976 – Dominican Republic (one sp.) Cubachelifer Hoff, 1946 – Cuba, Dominican Republic (one sp.) Troglobochica Muchmore, 1984 – Jamaica (two spp.) The pseudoscorpions are distributed between the main countries as follows: Cuba, 27; Jamaica, 31; Puerto Rico, 18; Haiti, 3; Dominican Republic, 26; Barbados, 1; US Virgin Island, 12; Dominica, 2; Martinique, 7; Guadeloupe, 8. 8.6 Antillean (Caribbean) Islands 659 Antilleans The fauna of Pseudoscorpiones in the Antilleans consists of 39 genera and 16 families. Genera and families in both areas: ---------------------------------------------------------------------------------------------------Central America Antilleans Chthoniidae Caribchthonius====================== Caribchthonius Muchmore Mundochthonius Chamberlin Pseudochthonius==========================Pseudochthonius Balzan Tyrannochthonius =====================Tyrannochthonius Chamberlin Lechythiidae – Lechytia Balzan Tridenchthoniidae – Tridenchthonius Balzan Bochicidae – Antillobisium Dumitresco et Orghidan (end. genus, Cuba) Mexobisium========================== Mexobisium Muchmore – Troglobochica Muchmore (end. genus, Jamaica) Neobisiidae Microbisium Chamberlin – Syarinidae – Ideobisium Balzan Ideoblothrus Balzan – Ideoroncidae – Typhloroncus Muchmore Albiorix Chamberlin – Pseudalbiorix Barba et Pérez – Sternophoridae Garyops ===================================== Garyops Banks – Idiogaryops Hoff Cheiridiidae – Cheiridium Menge – Cryptocheiridium Chamberlin Pseudochiridiidae – Pseudochiridium With Garypinidae Serianus ==================================Serianus Chamberlin Atemnidae – Atemnus Canestrini –– Oratemnus Beier Paratemnoides ======================== Paratemnoides Harvey Olpiidae 660 8 Some Peculiarities of the Distribution of Arachnida – Aphelolpium Hoff Olpiolum ====================================== Olpiolum Beier Garypidae – Planctolpium Hoff Cheliferidae – Cubachelifer Hoff (Dominican Rep., Cuba – end. genus) Parachelifer ==============================Parachelifer Chamberlin – Tyrannochelifer Chamberlin Chernetidae – Antillochernes Muchmore – Bituberochernes Muchmore – Byrsochernes Beier – Caribochernes Beier (Dominican Rep., endemic genus) – Chelanops Gervais – Chernes Menge – Dinochernes Beier – Hesperochernes Chamberlin Lustrochernes ==== =========== ================ Lustrochernes Beier Parachernes (Parachernes) Chamberlin Parachernes (Scapanochernes)========Parachernes (Scapanochernes Beier) – Parazaona Beier Withiidae Cacodemonius Chamberlin===================Cacodemonius Chamberlin Dolichowithius (D.) Chamberlin============Dolichowithius (D.) Chamberlin ========================================================= Opiliones Cyphophthalmi – Not recorded on the Antilleans Eupnoi (Avram 1970, 1973a, b, 1977a, b, 1981; Banks 1909; Cokendolpher and Camilo – Rivera, 1989, Goodnight and Goodnight 1942; Rambla 1969; Roewer 1910, 1947; Šilhavý 1971, 1973, 1976, 1979; Staręga 1970) Fam. Sclerosomatidae – Geaya Roewer, 1910 (Haiti) Dyspnoi – Not recorded Laniatores According to Kury (2003), in the Caribbean area are known 117 species of Opiliones Laniatores, belonging to 65 genera and 11 families: Cosmetidae, Agoristenidae, Stygnidae, Stygnommatidae, Kimulidae (Minuidae), Biantidae, Samoidae, Podoctidae, Manaosbiidae, Phalangodidae, and Zalmoxidae. The subfamily Stenostygninae of Biantidae consists almost entirely of taxa from the Caribbean (eight of the nine genera). By far the best studied island is Cuba, not only because of its size but mainly as a result of the studies of the Cubano-Romanian Expeditions (S. Avram and V. 8.6 Antillean (Caribbean) Islands 661 Šilhavý described from Cuba and other islands 58 new species and many new genera). Here are listed the endemic taxa in the Antilleans: Fam. Cosmetidae Arucillus hispaniolicus Šilhavý, 1971 – Dominican Republic Cynortula garna Goodnight et Goodnight, 1942 – Bahamas (Andros) C. sinensis Goodnight et Goodnight, 1942 – Bahamas (New Providence) C. juncta (Gervais, 1844) – Cuba C. fraterna Banks, 1909 – Cuba C. quinquesignata Franganillo Balboa, 1926 – Cuba Cynorta sextuberculata Franganillo Balboa, 1926 – Cuba C. hassleri Goodnight et Goodnight, 1942 – Haiti C. lithoclasica Avram 1981 – Cuba C. poaensis Avram 1981 – Cuba C. quibijana Avram 1981 – Cuba Cynortoides caraibicus (Sørensen, 1932) – US Virgin Islands, St. Thomas C. cubanus cubanus (Banks, 1909) – Cuba C. cubanus signatus Roewer, 1912 – Cuba C. lateralis Roewer, 1947 – Jamaica C. roeweri (Henriksen, 1932) – Puerto Rico C. quadrispinosus Goodnight et Goodnight, 1942 – Jamaica C. roeweri (Henriksen, 1932) – Cuba, Puerto Rico C. caraibicus (Sørensen, 1932) – Dominican Republic C. marginatus Goodnight et Goodnight, 1942 – Dominican Republic C. v-album (Simon, 1879) – Dominican Republic, Haiti, Tortuga Island Cynortesta laevis Roewer, 1947 – Windward Islands C. granulata Roewer, 1947 – Windward Islands: Saint Vincent and the Grenadines Eucynortoides antillarum Roewer, 1947 – Windward Islands: Saint Vincent and the Grenadines Cynortellana quadrimaculata (Gervais, 1844) – Cuba C. bisignata (Banks, 1909) – Cuba Heterovonones insularis Roewer, 1947 – Cuba Erginulus castaneus (Banks, 1906) – Bahamas (Andros, New Providence) E. quadricostatus (Franganillo Balboa, 1926) – Cuba Metacynortoides bilineatus Goodnight et Goodnight, 1942 – Dominican Republic M. obscurus dorsalis Roewer, 1916 – US Virgin Islands: St. Croix, St. Thomas M. obscurus obscurus (Banks, 1901) – Haiti, Jamaica, Puerto Rico, US Virgin Islands, St. John M. romanus Goodnight et Goodnight, 1942 – Dominican Republic M. transversalis Goodnight et Goodnight, 1942 – Dominican Republic M. scabrosus (Banks, 1909) – Cuba Paecilaema luquillense H. Soares, 1990 – Puerto Rico 662 8 Some Peculiarities of the Distribution of Arachnida P. conspicillatus Simon, 1879 – Windward Islands, Martinique Platycynorta secunda Roewer, 1947 – Cuba Prasiana fallax (Sørensen, 1932) – West Indies Proerginus lineatus Roewer, 1917 – Dominican Republic Trinimontius darlingtoni Šilhavý, 1970 – Cuba Vonones sayi (Simon, 1879) – Cuba V. granulatus Roewer, 1947 – Leeward Islands, Antigua and Barbuda V. planus Goodnight et Goodnight, 1942 – Leeward Islands, Dominica Fam. Samoidae Akdalima jamaicana Šilhavý, 1979 – Jamaica Arganotus robustus Šilhavý, 1979 – Haiti Hummelinckiolus parvus Šilhavý, 1979 – Leeward Islands: Guadeloupe, Montserrat, St. Kitts and Nevis Maracaynatum cubanum Šilhavý, 1979 – Cuba M. stridulans Šilhavý 1979 – Cuba Orsa daphne Šilhavý, 1979 – Haiti Pellobunus haitiensis (Šilhavý, 1979) – Haiti Reventula amabilis Šilhavý, 1979 – Jamaica Fam. Stygnidae Heterostygninae Stygnoplus antiguanus (Roewer, 1943) – Leeward Islands, Antigua and Barbuda S. flavitarsis (Simon, 1879) – Leeward Islands, Guadeloupe S. tuberculatus (Goodnight et Goodnight, 1942) – Leeward Islands, Dominica Fam. Stygnommatidae Stygnomma spiniferum bolivari (Goodnight et Goodnight, 1945) – Cuba S. spiniferum spiniferum (Packard 1888) – Florida Keys (USA): Tortugas, Jamaica S. spirulina (Goodnight et Goodnight, 1942) – Puerto Rico S. fiskei Rambla, 1969 – Jamaica Fam. Agoristenidae Agoristeninae Agoristenus cubanus Šilhavý, 1973 – Cuba haitiensis Šilhavý, 1973 – Dominican Republic Ahotta hispaniolica Šilhavý, 1973 – Haiti Calmotrinus turquinensis Šilhavý, 1973 – Cuba Dumitrescuella ornata Avram, 1977 – Cuba Haitimera paeninsularis Šilhavý, 1973 – Haiti Lichirtes hexapodoides Šilhavý, 1973 – Cuba Meriosfera gertschi Šilhavý, 1973 – Haiti M. lineata Šilhavý, 1973 – Haiti Orghidaniella granpiedrae Avram, 1977 – Cuba Piratrinus calcaratus Šilhavý, 1973 – Cuba 8.6 Antillean (Caribbean) Islands 663 Torreana poeyi Avram, 1977 – Cuba T. spinata Avram, 1977 – Cuba Vampyrostenus kratochvili Šilhavý, 1976 – Puerto Rico Yunquenus portoricanus Šilhavý, 1973 – Puerto Rico Fam. Phalangodidae Phalangodes flavipes (Banks, 1908) – Cuba Fam. Biantidae Stenostygninae Caribbiantes cubanus Šilhavý, 1973 – Cuba Galibrotus carlotanus Šilhavý, 1973 – Cuba G. matiasis Avram, 1977 – Cuba G. riedeli Šilhavý, 1973 – Cuba Bidoma indivisa Šilhavý, 1973 – Haiti Decuella cubaorientalis Avram, 1977 – Cuba Negreaella fundorai Avram, 1977 – Cuba N. palenquensis Avram, 1977 – Cuba N. rioindiocubanicola Avram, 1977 – Cuba N. vinai Avram, 1977 – Cuba N. yumuriensis Avram, 1977 – Cuba Manahunca bielawskii Šilhavý, 1973 – Cuba M. cuevajibarae Avram, 1977 – Cuba M. silhavyi Avram, 1977 – Cuba Martibianta virginsulana Šilhavý, 1973 – US Virgin Islands, St. John Vestitecola haitensis Šilhavý, 1973 – Haiti Fam. Podoctidae Ibaloniinae [Santobius cubanus = Ibantila cubana (Šilhavý, 1969) – Cuba (probably introduced from Melanesia, see Kury and Machado (2009))] Fam. Kimulidae (Minuidae) Kimula levii Šilhavý, 1969 – Cuba K. banksi Šilhavý, 1969 – Cuba K. goodnightiorum Šilhavý, 1969 – Cuba K. cokendolpheri Pérez et Armas, 2000 – Dominican Republic K. elongata Goodnight et Goodnight, 1942 – Puerto Rico K. tuberculata Goodnight et Goodnight, 1943 – Cuba K. turquinensis Šilhavý, 1969 – Cuba Metakimula botosaneanui Avram, 1973 – Cuba Minuides milleri Šilhavý 1978 – Cuba Fam. Manaosbiidae Cranellus balthazar Roewer, 1932 – Windward Islands 664 8 Some Peculiarities of the Distribution of Arachnida Sanvincentia tarsalis Roewer, 1943 – Windward Islands: Saint Vincent and the Grenadines Fam. Zalmoxidae Cersa kratochvili Šilhavý 1979 – Cuba Ethobunus cubensis (Šilhavý 1979) – Cuba E. goodnighti (Rambla, 1969) – Jamaica E. pecki (Rambla, 1969) – Jamaica E. zebroides (Šilhavý 1979) – Cuba Pachylicus castaneus (Šilhavý 1979) – Cuba Family uncertain Anamota custodiens Šilhavý 1979 – Cuba Caribula longimana Šilhavý 1979 – Cuba Jimeneziella negreai Avram, 1970 – Cuba (end. genus) J. decui Avram, 1970 – Cuba Metapellobunus unicolor (Roewer, 1912) – US Virgin Islands, St. Thomas Mirda insulanus (Banks, 1901) – Haiti, Puerto Rico Neoscotolemon pictipes (Banks, 1908) – Cuba N. lutzi Goodnight et Goodnight, 1942 – Leeward Islands, Dominica Paraconomma ovala Goodnight et Goodnight, 1942 – Puerto Rico P. spinooculorum Goodnight et Goodnight, 1942 – Puerto Rico Pseudomitraceras minutus Goodnight et Goodnight, 1942 – Puerto Rico Turquinia montana Šilhavý 1979 – Cuba Valifema blanda Šilhavý 1979 – Cuba Endemic genera of Opiliones in the Caribbean are: Fam. Minuidae Jimeneziella Avram, 1970 – Cuba (two spp.) Kimula Goodnight et Goodnight, 1942 – Cuba, Dominican Republic, Puerto Rico (eight spp.) Fam. Biantidae (Stenostygninae) Bidoma Šilhavý, 1973 – Haiti (one sp.) Caribbiantes Šilhavý, 1973 – Cuba (one sp.) Decuella Avram, 1977 – Cuba (one sp.) Galibrotus Šilhavý, 1973 – Cuba (three spp.) Manahunca Šilhavý, 1973 – Cuba (three spp.) Martibianta Šilhavý, 1973 – US Virgin Islands (one sp.) Negreaella Avram, 1977 – Cuba (five spp.) Vestitecola Šilhavý, 1973 – Haiti (one sp.) Fam. Samoidae Neoorsa Ozdikmen, 2006 (Orsa Šilhavý, 1979) – Haiti (one sp.) Reventula Šilhavý, 1979 – Jamaica (one sp.) 8.6 Antillean (Caribbean) Islands 665 Vlachiolus Šilhavý, 1979 – Cuba (one sp.) Amblypygi On the Antillean Archipelago have been recorded 29 species of Amblypygi from 4 genera (Charinus, Heterophrynus, Paraphrynus, Phrynus) and 2 families (Charinidae and Phrynidae). The endemism is only on species level (25 species), most species being confined to only 1 island (Armas 2004, 2006, 2007, 2009a, 2010, 2013; Armas and Avila Calvo 2000; Armas and Pérez Gonzalez 1994, 2001, 2002; Armas and Teruel 1997; Avila Calvo and Armas 1997; Franganillo 1926; Mullinex 1975; Pocock 1893; Quintero 1981, 1983, 1986; Teruel 2011; Teruel and Questel 2011; Teruel et al. 2009). Four species (three Phrynus and one Heterophrynus) are known also from South America (or Florida). This is in contrast with the order of scorpions, where 4 of the 15 genera are endemic to the Antilleans. As Armas (2009b) points it, Cuba has the most diverse amblypygid fauna (2 fam., 3 genera, 16 species, incl. endemics). Follows Hispaniola with two families, two genera, and seven species (incl. endemics). Now (2017) the distribution of the Amblypygi in the Antillean is: Fam. Charinidae – Charinus Simon (12 spp., Cuba, Jamaica, Dominican Rep., Puerto Rico, Lesser Antilleans) Fam. Phrynidae – Heterophrynus Pocock (one sp., Trinidad and Tobago), Paraphrynus Moreno (three spp., Bahamas and Cuba), Phrynus Lamarck (14 spp.) Thelyphonida (Uropygi) According to the list of Rowland and Cooke (1973), on the Antillean islands live only two spp. of Uropygi: Mastigoproctus baracoensis Franganillo, 1931 (Cuba), and the second described species in Uropygi M. proscorpio (Latreille, 1806) (Dominican Rep., Haiti, and Martinique). Another Mastigoproctus (M. pelegrini Armas, 2000) was described later from Cuba. Armas (2002) described a new Uropygid from the Dominican Republic, first as Telyphonellus wetherbeei and then raised in a new genus Ravilops Viquez et Armas, 2005. So far the two Cuban Mastigoproctus and Ravilops wetherbeei are considered endemic species; Ravilops is also an endemic genus for the Dominican Republic (Armas 2000, 2004; Franganillo 1931; Viquez and de Armas 2005). Schizomida Identified Schizomida are known (Ávila Calvo and Armas 1997; Armas 1977, 1989, 2004, 2011; Armas and Abud Antun 1990, 2002; Armas and Teruel 2002; Camilo and Cokendolpher 1988; Dumitresco 1973, 1977; Hilton 1933; Quintero 1983; Reddell and Cokendolpher 1995; Rowland and Reddell 1977; Teruel 2003, 2004, 2007) from: Cuba – Antillostenochrus alejandroi, A. alticola, A. cokendolpheri, A. gibarensis, A. holguin, A. planicauda, Cubazomus armasi, C. montanus, C. orghidani, C. rowlandi, Guanazomus armatus, Reddellzomus cubensis, Rowlandius abeli, R. alay- 666 8 Some Peculiarities of the Distribution of Arachnida oni, R. baracoae, R. biconouros, R. cubanacan, R. cupeyalensis, R. decui, R. digitiger, R. falcifemus, R. florentiae, R. gladiger, R. gracilis, R. guantanamero, R. labarcae, R. littoralis, R. marianae, R. melici, R. mixtus, R. monticola, R. negreai, R. ramosi, R. recuerdo, R. reyesi, R. serrano, R. siboney, R. teruel, R. toldo, R. vinai, Stenochrus alejandroi, S. portoricensis, Troglocubazomus orghidani, T. rowlandi Pinos (Isla de la Juventud) – Luisarmasius insulaepinorum Hispaniola (Dominican Rep. and Haiti) – Antillostenochrus brevipatellatus, A. subcerdoso, Cokendolpherius ramosi, Rowlandius anasilviae, R. casabito, R. ducoudrayi, R. engombe, R. isabel, R. jarmillae, R. lantiguai, R. longipalpus, R. naranjo, R. virginiae, Stenochrus subcerdoso, S. portoricensis Dominica – Stenochrus portoricensis Puerto Rico (incl. Isla Desecheo, Mona Island) – Antillostenochrus cerdoso, Luisarmasius yunquensis, Rowlandius desecheo, R. monensis Jamaica – Caribezomus laurae, Rowlandius cousinensis, R. peckorum, R. primibiconourus, R. viridis, Stenochrus portoricensis, Stewardpeckius troglobius Navassa – Rowlandius steineri Virgin Islands – Stenochrus portoricensis Martinique – Hansenochrus dispar, Rowlandius insignis There is also unidentified material from Barbados and Cayman Islands. Except Stenochrus portoricensis, all species are endemic to the respective islands. Endemic to the Caribbean are also the genera Cokendolpherius (Cuba), Luisarmasius (Cuba, Pinos, Puerto Rico), Reddellzomus (Cuba), Stewardpeckius (Jamaica), Cubazomus (Cuba), Troglocubazomus, and Heterocubazomus (Cuba). Araneae Simon (1888b); Alayón (1994, 1995, 2005), Bruyant (1940, 1947, 1948), Dumitrescu (1973), Dumitrescu, and Georgescu (1992) According to Alayón (2000), the spiders recorded from Cuba are 567 spp. of 52 families, including 247 endemic to the Antilleans (43.56%) with 7 endemic genera (Troglothele Fage, Bryantina Brignoli, Fageicera Dumitresco et Georgesco, Caudalia Alayón, Cubanopillus Alayón et Platnick, Decaphora Franganillo, Paraplexippus Franganillo). From the Antilleans are known some endemic genera of spiders, but there are no endemic families: Fam. Theraphosidae Antillena Bertani, Huff et Fukushima, 2017 – Dominican Republic Caribena Fukushima et Bertani, 2017 – Puerto Rico, Cuba, US Virgin Islands, Martinique Cubanana Ortiz, 2008 – Cuba Nesipelma Schmidt et Kovarik, 1996 – Nevis Fam. Ochyroceratidae Fageicera Dumitrescu et Georgescu, 1992 – Cuba 8.6 Antillean (Caribbean) Islands 667 Fam. Barychelidae Troglothele Fage, 1929 – Cuba Fam. Agelenidae Neowadotes Alayón, 1995 – Hispaniola Fam. Cyatholipidae Pokennips Griswold, 2001 – Jamaica Fam. Tetragnathidae Ancinosphenus Simon, 1895 – West Indies Antillognatha Bryant, 1945 – Hispaniola Hispanognatha Bryant, 1945 – Hispaniola Fam. Linyphiidae Lomaita Bryant, 1948 – Hispaniola Fam. Filistatidae Antilloides Breskovit et al., 2016 – Cuba, Dominican Republic, US Virgin Islands, Puerto Rico Fam. Paratropididae Anisaspis Simon, 1891 – Saint Vincent Fam. Amaurobiidae Tugana Chamberlin, 1948 – Cuba, Hispaniola Fam. Anyphaenidae Thaloe Brescovit, 1993 – Cuba, Hispaniola Fam. Theridiidae Jamaitidion Wunderlich, 1995 – Jamaica Fam. Liocranidae Laudetia Gertsch, 1941 – Dominican Republic, Puerto Rico Mesobria Simon, 1897 – Saint Vincent Fam. Ctenidae Ciba Bloom et al., 2014 – Cuba, Dominican Republic Ohvida Polotow et Brescovit, 2009 – Cuba, Puerto Rico, Bahamas Trujillina Bryant, 1948 – Hispaniola, Puerto Rico Fam. Pholcidae Bryantina Brignoli, 1985 – Cuba 668 8 Some Peculiarities of the Distribution of Arachnida Ciboneya Perez, 2001 – Cuba Platnicknia Özdikmen et Demir, 2009 – Cuba Tainonia Huber, 2000 – Hispaniola Fam. Prodidomidae Caudalia Alayón, 1980 – Cuba Cubanopillus Alayón et Platnick, 1993 – Cuba Fam. Sparassidae Decaphora Franganillo, 1931 – Cuba Fam. Thomisidae Rejanellus Lise, 2005 – Cuba, Puerto Rico, Hispaniola Fam. Salticidae Allodecta Bruyant, 1950 – Jamaica Antillattus Bryant, 1943 – Hispaniola, Cuba Bythocrotus Simon, 1903 – Hispaniola Caribattus Bryant, 1950 – Jamaica Cerionesta Simon, 1901 – Saint Vincent Commoris Simon, 1902 – Guadeloupe, Dominica Corticettus Zhang et Maddison, 1902 – Porto Rico, Hispaniola Paraplexippus Franganillo, 1930 – Cuba Parasaitis Bruyant, 1950 – Jamaica Parathiodina Bruyant, 1943 – Hispaniola Popcornella Zhang et Maddison, 2012 – Hispaniola, Puerto Rico Truncattus Zhang et Maddison, 2012 – Hispaniola Opilioacarida From Cuba and the Dominican Republic has been described the new genus Caribeacarus Vásquez et Klompen, 2009, from which two species have been described from Cuba, one also from the Dominican Republic, and one from Panama. Another species, described from Cuba (and the Antillean Islands), was Neoacarus orghidani (Juvara-Balş et Baltac 1977). Two of the three Cuban species and Caribeacarus panamensis live in caves (Beron 2014; Juvara-Balş and Baltac 1977; Vásquez and Klompen 2009). Parasitiformes (Mesostigmata and Ixodida): Ref.: Beron (2014), Cruz (2001) From Capromys pilorides (Rodentia, Capromyidae) has been described the endemic Ixodes capromydis Černy, 1966 (endemic subgenus Alloixodes). The purely Neotropical family of Spelaeorhynchidae (parasites on bats of the families Phyllostomidae and Mormoopidae) is found, together with these bats, in Cuba, Jamaica, Puerto Rico, and Dominican Republic but also in Central America. Holothyrida The only holothyrid recorded from the Antilleans is the endemic genus Carabothyrus Kontchán et Mahunka, 2004, with one species C. barbatus Kontchán et Mahunka, 2004, described from the Dominican Republic. 8.6 Antillean (Caribbean) Islands 8.6.2 669 Comparison Between the Faunas of Central America and the Antilleans Palpigradi – Known only from the Antilleans Ricinulei Central America In the 7 countries of Central America (without South Mexico) are registered 18 species of Ricinulei: Cryptocellus Westwood, 1874 – 12 spp. Pseudocellus Platnick, 1980 – Six spp. Antilleans Two endemic spp. of Cryptocellus from Cuba Solifugae – Endemic genera in bold Central America – From the seven countries of Central America (without South Mexico) are known ten spp. of Solifugae of five genera, all belonging to Ammotrechidae family (Ammotrechinae): Ammotrecha Banks, 1900 (four spp.); Ammotrechella Roewer, 1934 (two spp.); Ammotrechesta Roewer, 1934 (five spp.); Innesa Roewer, 1934 (one sp., endemic genus); and Ammotrechula Roewer. Antilleans – From the Antilleans are known four genera of Solifugae (all from Ammotrechidae), three of them endemic, all belonging to Ammotrechidae: Ammotrechinus Roewer, 1934 (one sp.); Ammotrechella Roewer, 1934 (nine spp.); Ammotrechona Roewer, 1934 (one sp.); and Antillotrecha Roewer, 1934 (three spp.). All Solifugae in both areas belong to the same family. There is only one genus that is common (Ammotrechella); the species are endemic. Amblypygi Central America – The amblypygids are represented in Central America by nine spp. Fam. Charinidae – Charinus Simon Fam. Phrynidae – Phrynus Lamarck, Paraphrynus Moreno Antilleans – Twenty-nine species of Amblypygi from four genera (Charinus, Heterophrynus, Paraphrynus, Phrynus) and the same two families (Charinidae and Phrynidae). The endemism is only on species level (25 species). Except Heterophrynus, known from the islands Trinidad and Tobago by the Venezuelan coast, the generic composition of Mesoamerica and the Antilleans is the same. 670 8 Some Peculiarities of the Distribution of Arachnida Thelyphonida (Uropygi) Central America – Five species (the endemics for Costa Rica Valeriophonus Viques et Armas, Mayacentrum tantalus Viquez et Armas for El Salvador, M. pijol V. et A. for Honduras, M. guatemalae V. et A. for Belize, and Mimoscorpius pugnator Butler for Guatemala). Antilleans – Four species (three Mastigoproctus Pocock, Ravilops Viques et Armas). Three species, including the genus Ravilops, are endemic for the Antilleans. Schizomida Central America In Central America (between Mexico and Colombia) are recorded seven species, but there is also unidentified material from many places (Reddell and Cokendolpher 1995). Besides the largely distributed Stenochrus portoricensis (Guatemala, Honduras, Nicaragua, Belize), from Central America, are known the genera Hansenochrus (Costa Rica, Panama), Rowlandius, and Surazomus (Costa Rica). Surazomus Reddell et Cokendolpher is known also from many South American countries. Rowlandius Reddell et Cokendolpher is widespread in the Caribbean Islands. Hansenochrus Reddell et Cokendolpher is found also in South America and the Caribbean. Antilleans From the Antilleans are known schizomids of nine genera (six endemic), all belonging to one family – Hubbardiidae. Central America Antilleans – Cubazomus Reddell et Cokendolpher – Heterocubazomus Teruel Rowlandius===========================Rowlandius Red. et Cok. Stenochrus ======================== =========Stenochrus Chamberlin Piaroa Manz. et al. – – Luisarmasius Red. et Cok. – Stewardpeckius Red. et Cok. – Reddellzomus Armas – Cokendolpherius Armas Surazomus – Hansenochrus========================Hansenochrus Red. et Cok. Mayazomus Red. et Cock. (Heteronychus Armas et Viques) Guatemala) – Belicenozomus Armas et Viques – ========================================================== 8.6 Antillean (Caribbean) Islands 671 Ref.: Ávila Calvo and Armas (1997), Armas (1977, 1989, 2002a, b, 2004, 2010), Armas and Abud Antun (1990, 2002), Armas and Teruel (2002), Camilo and Cokendolpher (1988), Dumitresco (1973, 1977), Hilton (1933), Quintero (1983a), Reddell and Cokendolpher (1995), Rowland and Reddell (1977) Identified Schizomida are known from: Cuba – Cubazomus armasi, C. orghidani, C. rowlandi, Rowlandius alayoni, R. baracoae, R. biconouros, R. cubanacan, R. decui, R. digitiger, R. gladiger, R. labarcae, R. negreai, R. recuerdo, Stenochrus alejandroi, S. portoricensis Pinos (Isla de la Juventud) – Luisarmasius insulaepinorum Hispaniola (Dominican Rep. and Haiti) – Rowlandius anasilviae, R. brevipatellatus, R. casabito, R. ducoudrayi, R. engombe, R. isabel, R. jarmillae, R. lantiguai, R. longipalpus, R. naranjo, R. virginae, Stenochrus subcerdoso, S. portoricensis Dominica – Stenochrus portoricensis Puerto Rico (incl. Isla Desecheo, Mona Island) – Luisarmasius yunquensis, Rowlandius desecheo, R. monensis Jamaica – Rowlandius cousinensis, R. peckorum, R. primibiconourus, R. viridis, Stenochrus portoricensis, Stewardpeckius troglobius Virgin Islands – Stenochrus portoricensis Martinique – Hansenochrus dispar, H. insignis There is also unidentified material from Barbados and Cayman Islands. Except Stenochrus portoricensis, all species are endemic to the respective islands. Endemic to the Caribbean are also the genera Cokendolpherius (Cuba), Luisarmasius (Cuba, Pinos, Puerto Rico), Reddellzomus (Cuba), Stewardpeckius (Jamaica), and Cubazomus (Cuba). Pseudoscorpiones Central America In the 7 countries of Central America are recorded pseudoscorpions of 18 genera and 12 families (Chthoniidae, Bochicidae, Neobisiidae, Syarinidae, Ideoroncidae, Sternophoridae, Garypinidae, Atemnidae, Olpiidae, Cheliferidae, Chernetidae, Withiidae). Antilleans The fauna of Pseudoscorpiones in the Antilleans consists in 39 genera and 16 families. 672 8 Some Peculiarities of the Distribution of Arachnida Genera and families in both areas Central America Antilleans Chthoniidae Caribchthonius Muchmore =============== Caribchthonius Muchmore Pseudochthonius Balzan====================== Pseudochthonius Balzan Tyrannochthonius Chamberlin===========-==Tyrannochthonius Chamberlin Mexobisium Muchmore =================== Mexobisium Muchmore Syarinidae Ideobisium Balzan ======================== Ideobisium Balzan Ideoblothrus Balzan ============================Ideoblothrus Balzan Sternophoridae Garyops Banks===================================Garyops Banks Serianus Chamberlin============================Serianus Chamberlin Atemnidae Paratemnoides Harvey====================Paratemnoides Harvey Olpiidae Olpiolum Beier================================== Olpiolum Beier Cheliferidae Parachelifer Chamberlin=====================Parachelifer Chamberlin Chernetidae Lustrochernes Beier ===========================Lustrochernes Beier Parachernes (Scapanochernes Beier)===Parachernes (Scapanochernes Beier) Withiidae Cacodemonius Chamberlin==================Cacodemonius Chamberlin Dolichowithius (D.) Chamberlin===========Dolichowithius (D.) Chamberlin Both areas have 8 families and 15 genera of pseudoscorpions in common. No endemic families, no end. genera in Central America, and four end. genera in the Antilleans. Scorpiones Central America Antilleans (without Trinidad and Tobago) Buthidae – = Alayotityus Armas Centruroides ==================================Centruroides Marx – Microtityus K.-W. – Rhopalurus Thorell – Tityopsis Pocock Tityus ======================================== Tityus C.L. Koch Chactidae – Chactas Gervais – Hormuridae Opisthacanthus ==============================Opisthacanthus Peters Scorpionidae 8.6 Antillean (Caribbean) Islands 673 – Cazierus Francke – Cryptoiclus Ter. et Kov. Didymocentrus ==============================Didymocentrus Kraep. – Heteronebo Pocock Diplocentrus Peters – Euscorpiidae Plesiochactas Pocock – Vaejovidae Vaejovis C.L. Koch – ========================================================= 8.6.3 Conclusion In the seven countries of Central America (without counting Yucatan) are known eight spp. of Ricinulei of both American genera. From the Antilleans (Cuba) have been recorded only two (endemic) species. The genus Cryptocellus Westwood is common in both areas. From the seven countries of Central America (without South Mexico) are known Solifugae of five genera, all belonging to Ammotrechidae family (Ammotrechinae). Two genera seem endemic. From the Antilleans are recorded four genera (all from the same family Ammotrechidae), three of them endemic. There is only one genus in common among both areas (Ammotrechella); the species are endemic. The Amblypygi of the Antilleans are much numerous and varied (29 spp. of 4 genera), than in the 7 countries of Central America (9 spp. of 2 genera, the families are the same). Again differences in the research? Both areas have different fauna of Thelyphonida (Uropygi). Central America has three endemic genera, the Caribbean one. Mastigoproctus is known from Cuba, Martinique, and Haiti but is so far absent from Central America. The Schizomida of both areas belong to one family, but the genera are very different (seven genera, one endemic in Central America, eight genera in the Antilleans, six endemic). Both areas have three genera in common. Again, differences in the research? 8.6.4 Opiliones Cyphophthami – Unknown in both areas Eupnoi: Sclerosomatidae (in both areas, the same genera Geaia and Prionostemma); Dyspnoi – in Honduras the genus Trilasma Goodnight et Goodnight (1942) (Nemastomatidae, Ortholasmatinae); in the Antilleans, Dyspnoi are not recorded. 674 8 Some Peculiarities of the Distribution of Arachnida Laniatores The catalogue of Kury (2003) enumerates from the seven countries of Central America 56 genera and 9 families of Laniatores (Cosmetidae, Cranaidae, Gonyleptidae, Stygnommatidae, Samoidae, Pyramidopidae, Manaosbiidae, Stygnopsidae, Zalmoxidae). From the Antilleans in the same catalogue are listed 117 species of Opiliones Laniatores, belonging to 65 genera and 11 families: Cosmetidae, Agoristenidae, Stygnidae, Stygnommatidae, Kimulidae (Minuidae), Biantidae, Samoidae, Podoctidae, Manaosbiidae, Phalangodidae, and Zalmoxidae. From this number, there are five families (underlined) in common. The endemic genera of opilions in the islands are 13 (Biantidae, Kimulidae). There are no endemic families. The fauna of pseudoscorpions in the Antilles seems much richer (39 genera, 16 families) than the fauna of the 7 countries of Central America (33 genera, 13 families). Both areas have 15 genera in common. This comparison shows that from 8 genera of scorpions in Central America and 11 genera in the Antilleans, there are 4 genera in common. Three families in Central America (Chactidae, Euscorpiidae, and Vaejovidae) are not recorded in the Antilleans. Fact is that the scorpions in the islands have more genera and species than the scorpions of Central America region with land connection with such rich areas like Mexico and northern South America. One explanation is the very active speciation, especially in Cuba (55 spp.) and the Dominican Republic (40 spp.). The other explanation is the presence in Cuba of such specialists as Armas and Teruel. Opilioacarida – Two genera from the same family (Neocarus Chamberlin et Mulaik and Caribeacarus Vásquez et Klompen) live in both areas. The species are (so far) endemic. Holothyrida are unknown in Central America. From the Dominican Republic has been described the endemic species Caribothyrus barbatus Kontchán et Mahunka, 2004. In general, there is considerable difference (artificial) between the archaeofauna of both areas but on low (genus, species) level. The Antilleans seem richer (or better explored). 8.7 8.7.1 Patagonia: South Chile, Argentina, Uruguay, and Tierra del Fuego Geography, General Zoogeography, and Paleogeography Argentina’s continental area is between the Andes mountain range in the west and the Atlantic Ocean in the east. Also claims the Falkland Island (Malvinas) and some parts of Antarctida. The territory is ca. 2 767 000 km2 (the second biggest in Latin America), stretching from the tropical forest of Iguazú Falls to Tierra del Fuego and the Subantarctic climate. The southern part of Argentina and Chile is called Patagonia (cool semidesert region of some 500 000 km2, integrating the 8.7 Patagonia: South Chile, Argentina, Uruguay, and Tierra del Fuego 675 southernmost section of the Andes mountains to the southwest toward the Pacific Ocean, and from the east of the cordillera to the valleys it follows south through the Colorado River toward Carmen de Patagones in the Atlantic Ocean). The Chilean part includes the southern provinces and regions of Valdivia, Los Lagos Region, and Greater Island of Chiloé and is composed of two subregions: the northern Aisen and, to its south, Magallanes. The altitudinal difference is the biggest in the Western Hemisphere – from Aconcagua (6962 m.) to the sea level. Some sources: Cabrera and Willink (1973), Harrington (1962), Kuschel (1960), Markgraf (1985), Mattoni (2007), Riccardi (1987), Roig-Juñent et al.(2006), Spalletti and Franzese (1996), and Uliana and Biddle (1988). Among the 40 centers of dispersal in the Neotropical Region, outlined by Müller (1972), only 2 (Patagonian and Nothofagus centers) are situated in South Chile and Patagonia. According to Crisci et al. (1991), “The northern South American biota is most closely related to that of North America, and southern South America constitutes a monophyletic group together with Australia, Tasmania, New Guinea, New Caledonia, and New Zealand, reflecting the existence of an ancient austral biota. Four conflicting hypotheses of area relationships concerning southern South America and the other austral areas are proposed, suggesting that southern South America may be a composite area in itself.” Among the biogeographical areas of Latin America of Morrone (2005), based on the entomofauna, is also the Andean Region, which “…is included in the Austral Kingdom, which corresponds to western Gondwanaland and comprises five other regions: Antarctic (Antarctica), Cape or Afrotemperate (South Africa), Neoguinean (New Guinea and New Caledonia), Temperate Australian (southeastern Australia), and Neozealandic (New Zealand).” These ideas are developed also in the papers of Brundin (1966), Rapoport (1968), and others. The Andean Subregion of Morrone includes Central Chilean Subregion (between latitudes 30 and 34oSouth), Subantarctic Subregion (from lat. 36 south to Cape Horn), and Patagonian Subregion with two provinces (Subandean Patagonia and Central Patagonia). 8.7.2 Arachnogeographical Character of South Chile and Argentina We consider here the area south of 30oS, as it is important to check from arachnological point of view the sharp delimitation (for some groups of vertebrates and insects) across Argentina and Chile of two Kingdoms – the Neogea (region Neotropica) and the Notogea. Is it such sharp difference what concerns the Arachnida and where is the limit (and/or transition zone)? In this area are registered families and even genera common with faraway lands, remnants from the Gondwana (South Africa, Australia, New Zealand) (Cabrera and Willink 1980; Cokendolpher and Lanfranco 1985; Mello-Leitão 1939; Ringuelet 676 8 Some Peculiarities of the Distribution of Arachnida 1957, 1959, 1961; Simon 1886). A catalogue of the Arachnida of the southernmost Chile (Magallanes) and the “Antarctica Chilena” was due to Cecalovic (1976). Amblypygi, Uropygi, Schizomida, and Ricinulei – No records in Argentina, Chile, Uruguay, and Paraguay Palpigradi From Chile has been described Prokoenenia chilensis Hansen (Prokoeneniidae) (Cekalovic 1984; Hansen 1901). Solifugae Muma (1971) lists for Chile 13 spp. of Solifugae in 9 genera and 2 families – Ammotrechidae and Amacataidae, described by him. Later have been described Chileotrecha Maury (1987) (endemic genus for Chile). Maury (papers from 1970 to 1998) provided information on many Solifugae inhabiting Argentina of the genera Pseudocleobis including P. alticola Pocock (1900) up to 4700 m, Gaucha Mello-Leitao, Valdesia Maury, and the families Ammotrechidae, Mummuciniidae, and Daesiidae (Kraus 1966; Maury 1976, 1980, 1981, 1982, 1983, 1987). Scorpiones There is vast information on the scorpions, mostly of Argentina (see table). In Argentina are known 66 spp. of only 2 families – Bothriuridae (53 spp.) and Buthidae (13 spp.) (Ojanguren-Affilastro, 2005 and suppl.). In Chile are recorded 53 spp. of 3 families – Bothriuridae (50 spp.), Buthidae (two spp.), and Caraboctonidae (one sp.). In Patagonia is represented only the family Buthriuridae with the genera Bothriurus Peters, Brachistosternus Pocock, and Urophonius Pocock. They live even to the southern end of Tierra del Fuego (Provincia Fitogeografica Subantarctica). It is clear that Bothriuridae (mainly the genera Bothriurus Peters and Brachistosternus Pocock) is strongly prevailing in both countries (Maury (1968, 1975a, 1979), MelloLeitao (1934, 1943, 1945), Ringuelet (1953), Cekalovich (1983)). The genera Centromachetes Lönnberg, Phoniocercus Pocock, Rumikiro Ojanguren-Affilastro et al., and Tehuankea Cekalovic are endemic to Chile. Uruguay has only five native genera (four Bothriuridae and one Buthidae). Pseudoscorpiones After the very careful collection of G. Topál in the most southern Andes (42oS), Beier (1964a) concluded that the pseudoscorpion fauna of Patagonia is rather poor (20 species of 11 genera and 5 families – Chthoniidae, Gymnobisiidae, Geogarypidae, Garypinidae, Chernetidae) (Beier 1962, 1967; Mahnert et al. 2011; Vitali-di Castri 1970; Cekalovic 1976). Endemic genus is Diplothrixochernes Beier (1962); the species are endemic to Patagonia in Argentina and South Chile. Beier (1964b) is emphasizing the importance of the Andes as a barrier along the meridian between the faunas of South Chile and South Argentina. From Chile (incl. Juan Fernandez Isl.) are known 83 spp. of pseudoscorpions (Cekalovic, 1984 and suppl.), from the whole of Argentina 70 spp. and from Uruguay 5 spp. In Argentina and Chile have been recorded pseudoscorpions of 55 genera and 16 families: Chthoniidae, Tridenchthoniidae, Pseudotyrranochthoniidae, Lechytiidae, 8.7 Patagonia: South Chile, Argentina, Uruguay, and Tierra del Fuego 677 Gymnobisiidae, Garypidae, Geogarypidae, Garypinidae, Ideoroncidae, Atemnidae, Olpiidae, Menthidae, Cheliferidae, Cheiridiidae, Chernetidae, and Withiidae. From Argentina are known 38 genera, from Chile also 38 genera, with 21 genera in common. Most intriguing is the distribution of family Gymnobisiidae. From the 11 species, 1 (Beierobisium oppositum Vitali-di Castri) is an endemic genus and species for the Falkland Islands, 2 (the genus Gymnobisium Beier) are endemic for South Africa, and 8 have been described from Chile, Argentina, and Bolivia. Opiliones From Argentina, Uruguay, and Chile of 30oS are known Opiliones of the families Caddidae, Sclerosomatidae, Neopilionidae, Acropsopilionidae, Triaenonychidae, Zalmozidae, Cosmetidae, and Gonyleptidae (Acosta 2002; Acosta and Maury 1998; Canals 1936; nine papers from 1932 to 1943; Capocasale 1968; Shear 1993b; Cokendolpher and Lanfranco 1985; Hunt and Cokendolpher 1991; Maury 1986, 1988, 1991, 1992, 1993; Maury et al. 1996; Ringuelet 1955; 26 papers from 1953 to 1963; Roewer 1961, 1962; Cecalovic 1968, 1976, 1985). From 120 species and subspecies, 79 belong to the family Gonyleptidae (mostly to Pachylinae). Zoogeographical analysis has been made several times, mostly by Mello-Leitão (1939) and Ringuelet (1957, 1959). Eupnoi Recorded are the families Caddidae, Neopilionidae, and Sclerosomatidae. Endemic genera: Fam. Neopilionidae Ballarrinae – Chile, Australia, South Africa (ten spp.) Americovibone Hunt et Cokendolpher, 1991 – Chile (one sp.) Enantiobuninae – Brazil, Chile, Argentina (four spp.) Thrasychirus Simon, 1884 – Chile, Argentina (three spp.) Dyspnoi – Fam. Acropsopilionidae, Acropsopilio Forster Laniatores Recorded are opilions of the families Gonyleptidae, Triaenonychidae, Zalmoxidae, and Cosmetidae. Endemic genera – Fam. Triaenonychidae: Picuchenops Maury, 1988, Argentina (one sp.) The zoogeography of Opiliones in Argentina has been treated several times (Mello-Leitao 1939; Ringuelet 1957, 1959; Acosta and Maury 1998; Acosta 2002). Mello-Leitão (1939) lists 79 spp. of Opiliones in Argentina and makes a subdivision of Argentina after the Arachnida into 5 zoogeographical provinces, including “Provincia patagónica” (the area south of Rio Negro). The most important monograph on Argentinian Opiliones is due to Ringuelet (1959). The highest number of species exists in the three wetter areas: the Andino-Patagonian forests, the selva and the yungas (Acosta and Maury 1998). Ringuelet (1959) proposed a subdivision of Argentina based on Opiliones: several zones in the northeast and a narrow stripe (mostly in Chile), called “zona austral,” including the “subzone malvinense” on the 678 8 Some Peculiarities of the Distribution of Arachnida Falkland Islands. Between these areas is spread a huge zone “Andina-SubandinaPatagonica,” considered negative for opilions. Later some endemic (relic?) species have been detected in this area. The opiliofaunistical conclusions of this author (Ringuelet 1957, 1959) have been altered by Acosta and Maury (1998) and Acosta (2002). After the last author, in the “área Andino-patagónica” live 22 species/subspecies of Opiliones (2 of Caddidae, 3 of Neopilionidae, 7 of Triaenonychidae, 10 of Gonyleptidae). From the cave system, Cuchillo Curá Maury (1988) described the only Argentinian troglobitic opilion Picunchenops spelaeus, belonging to a group far from the other Triaenonychidae. According to Acosta (2002), by this time the Argentinian fauna of Opiliones includes120 “entidades” (species and subspecies), relatively few for the territory of almost 2 800 000 km2. This is due to the dry conditions of most of the country. Family Gonyleptidae is predominant with 79 “entidades” (69 of them belong to Pachylinae, 57% of all Argentinian Opiliones). Araneae In the austral part of South America are registered at least 62 families of spiders, some of which are zoogeographically very interesting (Forster and Platnick 1984; Platnick and Forster 1989; Merian 1913; Richardson 2010; Zapfe 1961). The families are Mecicobothriidae, Microstigmatidae, Hexathelidae, Dipluridae, Nemesiidae, Cyrtaucheniidae, Idiopidae, Migidae, and Actinopodidae from infraorder Mygalomorpha and Austrochilidae, Filistatidae, Drymusidae, Scytodidae, Sicariidae, Diguetidae, Pholcidae, Caponiidae, Dysderidae, Oonopidae, Orsolabidae, Segestriidae, Hersiliidae, Oecobiidae, Mecysmaucheniidae, Palpimanidae, Malkaridae (incl. Pararchaeidae), Mimetidae, Deinopidae, Uloboridae, Anapidae (incl. Micropholcommatidae), Araneidae (incl. Nephilidae), Linyphiidae, Synotaxidae, Tetragnathidae, Theridiidae, Lycosidae, Oxyopidae, Pisauridae, Senoculidae, Trechaleidae, Agelenidae, Amphinectidae, Amaurobiidae, Anyphaenidae, Desidae, Dictynidae, Hahniidae, Sparassidae, Selenopidae, Zodariidae, Zoropsidae, Clubionidae, Miturgidae (incl. Zoridae), Titanoecidae, Gallienellidae, Gnaphosidae, Prodidomidae, Trochanteriidae, Philodromidae, Thomisidae, Salticidae, and Corinnidae from suborder Arachnomorphae. It is worth to analyze the distribution of some families. Fam. Mecysmaucheniidae – The genera Mecysmauchenius Simon and Mecysmauchenioides Forster et Platnick, 1984, are outlined by Grismado and Lopardo (2003). The 17 species of this genus live in Patagonia and the islands Malvinas (Falkland Island) and Juan Fernandez. Other genera of this family in Chile and/or Argentina are Chilarchaea Forster et Platnick, 1984; Mesarchaea Forster et Platnick, 1984; and Semysauenius Forster et Platnick, 1984. The last two genera of the family are known from the other end of the world – New Zealand: Aotearoa Forster et Platnick, 1984, and Zearchaea Wilton, 1946. This is one of the amazing cases of faraway disjunctions. Fam. Malkaridae – This small family (46 spp.) included the genera Perissopmerus Butler, 1932, from the Australian continent; Malkara Davies, 1980, from Queensland; Carathea Moran, 1986, from Tasmania; and, surprisingly, Chilenodes 8.8 Falkland Islands (Malvinas) 679 Platnick et Forster, 1987, from Chile and Argentina. Later to it was added the family Pararchaeidae with six genera from Australia, Tasmania, and New Zealand. Fam. Amphinectidae – From the 35 known genera of this family, 18 live in New Zealand and 16 in Australia (incl. Tasmania), but 2 genera are known from Chile (Calacadia Exline, 1960) and Western coast of South America (Metaltella MelloLeitão, 1932). Opilioacarida Neocarus platensis (Silvestri, 1905) from “La Plata” (ArgentinaUruguay) is the southernmost member of Opilioacarida in the New World. Acariformes Oribatida – Hammer (1962a, 1962b) Parasitiformes – Ref.: Beron (2014) From Panama, Brazil, and Uruguay is known the family Dasyponyssidae – endemic to the Neotropics, as is composed by parasites on Armadillos (Cingulata, Dasypodidae). 8.8 8.8.1 Falkland Islands (Malvinas) Geography, General Zoogeography, and Paleogeography The Falkland Islands (Islas Malvinas) are an archipelago in the Southern Atlantic Ocean, on 51o45’S and about 480 km. from Argentina. The archipelago consists of two main islands of East and West Falkland and ca. 200 small islands with total surface area of 12 173 km2, on the Patagonian Shelf. The climate is cold marine, with occasional snow all year, except in January and February, and with strong winds throughout the year. The highest point is Mount Usborne (705 m). Falkland Islands were originally part of South Africa. They have achieved their present position during the fragmentation of Gondwana (Marshall 1994). Due to the strong winds, they are treeless, with only dwarf shrubs, but have 163 spp. of native vascular plants. The islands are biogeographically part of the mild Antarctic zone, with strong connections to the flora and fauna of Patagonia (McDowall 2005). According to Morrone (2000), biogeographically the Falkland Islands (Malvinas) are a province within the Subantarctic subregion of the Andean Region. Arachnogeography Only one pseudoscorpion, two opilions, and some spiders and mites are known from the Falkland Islands. They are among the southernmost arachnids (Ringuelet 1955). Pseudoscorpiones Beierobisium oppositum Vitali-di-Castri, 1970, is the only pseudoscorpion known from these islands (endemic genus and species of Gymnobisiidae). 680 8 Some Peculiarities of the Distribution of Arachnida Opiliones Information about the few opilions on the islands is due to Soares and Soares (1949), Hogg (1913), and Butler (1875). Laniatores Fam. Gonyleptidae: Haversia defensa (Butler), Haversia (Hoggellula) vallentini (Hogg) Araneae The spiders belong to the families Orsolobidae, Linyphiidae, Zodariidae, and others (Forster and Platnick (1985), Hogg (1913), Lavery and Snazell (2013), Platnick (1977), Schiapelli and Gershman de Pikelin (1974), Grismado and Platnick (2008), Usher (1983)). Endemic genera: Fam. Orsolobidae Falklandia Forster et Platnick, 1985 Fam. Linyphiidae Falklandoglenes Usher, 1983 Linga Lavery et Snazell, 2013 8.9 8.9.1 The Boundary Between Palearctic and Paleotropical The Boundary Between Western Palearctic and Afrotropical Arachnofaunas It is a common knowledge that a biogeographical boundary is not a line but rather a more or less wide transitional zone. We have also to take into account the historical changes during the last several thousand years. Part of these changes is due to climatic factors, others to human activities (also causing changes in the climate), like deforestation, diverting of rivers, etc. The traditional zoogeographic regions are delimitated mainly on the distribution of terrestrial vertebrates. The difference between the regions and kingdoms is usually on much higher taxonomical level when we consider vertebrates, than with the Arachnida. Differences between Afrotropical and Neotropical regions, or between Holarctic and Paleotropical, are between orders or suborders and very much between families. Historically it was not as it is today. In green Sahara and Mediterranean Africa lived elephants, crocodiles, and giraffes; in Atlas Mountains, there were until recently ostriches, lions, and bears. They were exterminated by man. We do not speak about geological periods when these animals (plus Tubulidentata, Marsupialia, 8.10 Arachnida in the Afrotropical and Palearctic Regions 681 tapirs, and many others) lived in Europe. The present-day biogeographical subdivision should consider the picture of the last several thousand years. We don’t know what was the distribution of many arachnids over the green Sahara. It was not influenced by the rare human population during the millenia, as it certainly was in Europe and North America with the disappearing of forests and their replacement with anthropogenic landscape. Important were also new crops like maiz and potatoes, chemization, and other kinds of human intruding into the ecosystems. If we follow the classical subdivision (Africa south of Sahara and the southernmost Arabia form the Afrotropical Region), north of this “line” is the Palearctic (different subregions). This “line” (actually transitional zone) is separating two kingdoms, so differences should be substantial. Let us check how the known distribution of Arachnida fits into the classical scheme (Vachon 1951a, b, 1952; Beron 2017a). 8.10 8.10.1 Arachnida in the Afrotropical and Palearctic Regions Analysis and Comments The Afrotropical Region is bordering only the Western part of the huge Palearctic Region (Saharo-Sindian Province). Many higher taxa are marked “present” for both regions; but actually they may be distributed in tropical Africa and, say, Japan or Korea. Sometime in between there is a gap of thousand kilometers, and the difference between the arachnofauna of tropical Africa and Europe is much bigger. So some comments on the distributions of the various taxa are needed. Palpigradi In tropical Africa are known eight described species of Palpigradi; one genus (Allokoenenia Silv.) is endemic. In the Western Palearctic (Europe and Lebanon) are recorded species, all from the genus Eukoenenia Börner (Eukoeneniidae). Ricinulei In the Old World this order is represented only by 11 species of 1 genus (Ricinoides, endemic for West Africa). Solifugae In the Afrotropical Region are registered eight families (three of which from South Africa). One of the striking facts in zoogeography is the absence of Solifugae in Madagascar. Scorpiones In Africa south of 20oN live scorpions of five families with genera (Bothriuridae, Buthidae, Hormuridae, Scorpionidae, Lisposomatidae). The underlined families are known only from Southern Africa. Schizomida The northern limit of Schizomida in Africa is determined by their thermopreferendum and the sands of Sahara – from Liberia to Kenya and Somalia. 682 8 Some Peculiarities of the Distribution of Arachnida There is one endemic subfamily in Southern Africa and five genera – all endemic for narrow areas of Africa. Thelyphonida (Uropygi) Only one genus (Etienneus) is represented in Africa (the Western part). In the Palearctic the order is also almost not represented (one species of Typopeltis in the far East of Russia). Amblypygi In the Afrotropical Region (incl. South Africa) live members of five genera of Euamblypygi of two families (Phrynichidae, Charinidae). Remarkable is the finding in Guinea-Bissau of the most primitive member of the order – Paracharon caecus Hansen, 1921 – sole living representative of a special family (Paracharontidae) and even suborder (Palaeoamblypygi). Very few amblypygids have been recorded in the Palearctic; none in Europe; Charinus in Turkey, the Dodecanese, Egypt, Israel, and Pakistan; and Musicodamon in Algeria and Morocco. Araneae We counted 81 families of spiders (out of the 112 families of spiders in the world), related to both regions (living in one of them or in both). In both regions occur 47 families, only in the Afrotropical Region 72 and also in the Palearctic Region 72 families. From the 72 families, represented in the Afrotropical Region, 10 are found only in Southern Africa (south of Zambezi-Kunene). Opilioacarida This peculiar order is represented in both territories by one family but with different genera: Opilioacarus within South Europe and Algeria, with two genera, endemic to West Africa (Panhaetes Naudo and Phalangiacarus Coineau and Van der Hammen) and with Salfacarus Van der Hammen, known from Southern Africa and Madagascar. Differences in Subordinal Level The orders Ricinulei and Schizomida and the suborder Palaeoamblypygi are found in Afrotropical Region but are absent in the Palearctic. The suborders Mesothelae and Dyspnoi are represented in the Palearctic, but not in tropical Africa (Table 8.4). This boundary has been analyzed by Beron (2016) and other authors. Martens (1984) concluded from his long experience in Himalaya that We must take into account that the Himalayan fauna is predominantly one which invaded the Himalayas at the time of or after the uplift of the mountain chain during the Tertiary and that Himalayan fauna has only partially undergone an evolution of its own. Both climatic belts and vegetation zones are largely in accordance with the areas of origin outside the Himalayas of the various faunal elements. The Himalayan fauna is mainly an immigration fauna. We distinguish five main centers of origin and thus five categories of Himalayan fauna, three of which fall into the Palearctic (Central Asian, Himalayan West Asian, Himalayan West Chinese) and two in the Oriental realm (Himalayan Indochinese, Peninsular Indian. When trekking along Kali Gandaki River, one have the strange feeling to cross a boundary between two kingdoms – Holarctic and Paleotropic. 8.10 Arachnida in the Afrotropical and Palearctic Regions 683 Table 8.4 The boundary between the Palearctic and the Indomalayan regions, indicated by Arachnida Group Order Palpigradi Fam. Eukoeneniidae Order Ricinulei Fam. Ricinoididae Order Solifugae Fam. Galeodidae Fam. Karschiidae Fam. Daesiidae Fam. Solpugidae Fam. Ceromidae Fam. Melanoblossiidae Fam. Gylippidae Fam. Hexisopodidae Fam. Rhagodidae Order Scorpiones Fam. Bothriuridae Fam. Buthidae Fam. Pseudochactidae Fam. Euscorpiidae Fam. Scorpiopidae Fam. Troglotayosicidae Fam. Iuridae Fam. Diplocentridae Fam. Hemiscorpiidae Fam. Hormuridae Fam. Scorpionidae Fam. Akravidae Fam. Lisposomidae Order Schizomida Fam. Hubbardiidae Order Uropygi Fam. Hypoctonidae Order Amblypygi Suborder Neoamblypygi Fam. Charinidae Fam. Phrynichidae Suborder Palaeoamblypygi Fam. Paracharontidae Order Opiliones Suborder Cyphophthalmi Region Afrotropical Present Present Present Present Present Present Absent Present Present Present Present (Southern Africa) Present (Southern Africa) Present (Southern Africa) Present Present Present (Namibia) Present Absent Absent Absent Absent Absent Absent Absent Present Present Absent Present (South Africa) Present Present Present (relict) Present Present Present Present Present Present Present Present Present Region Palearctic Present Present Absent Absent Present Present Present Present Present Absent Absent Present Absent Present Present Indian Himalaya Present Present Present Present ? Present Present Present Present Present Present Present Absent Absent Absent Present Present Present Present Present Present Absent Absent Present Present (continued) 684 8 Some Peculiarities of the Distribution of Arachnida Table 8.4 (continued) Fam. Sironidae Fam. Ogoveidae Fam. Neogoveidae Fam. Pettalidae Suborder Eupnoi Fam. Caddidae Fam. Phalangiidae Fam. Sclerosomatidae Fam. Neopilionidae Suborder Dyspnoi Fam. Ischyropsalididae Fam. Sabaconidae Fam. Dicranolasmatidae Fam. Trogulidae Fam. Nemastomatidae Fam. Nipponopsalididae Suborder Laniatores Fam. Cladonychiidae Present Present Present Present (South Africa) Present Present (South Africa) Present Present Present (South Africa) Absent Absent Absent Absent Absent Absent Absent Present Absent Fam. Travuniidae Fam. Triaenonychidae Fam. Assamiidae Fam. Biantidae Fam. Epedanidae Fam. Phalangodidae Fam. Podoctidae Fam. Pyramidopidae Fam. Samoidae Fam. Zalmoxidae Order Pseudoscorpiones Suborder Epiocheirata Fam. Chthoniidae Fam. Tridenchthoniidae Fam. Pseudotyrannochthoniidae Fam. Lechytiidae Fam. Feaellidae Suborder Iocheirata Fam. Ideoroncidae Fam. Hyidae Fam. Gymnobisiidae Fam. Neobisiidae Fam. Syarinidae Absent Present (South Africa) Present Present Absent Absent Present Present Present (Mozambique) Seychelles Present Present Present Present, India, Seychelles Absent Present Absent Absent Absent Present Present (Japan) Present Present Absent Present Present Present Present Present Present Present (Japan, Korea) Present Present (Holoscotolemon) Present Present (Japan, Korea) Absent Nepal, India Japan, India, Taiwan Present Present (Japan, India) Absent Absent Absent Present Present Present Japan, Bhutan, Nepal, Present Present Present Present Present Absent Present (South Africa) Present Present Present India Present Present India Present Present Present (continued) 8.10 Arachnida in the Afrotropical and Palearctic Regions 685 Table 8.4 (continued) Fam. Garypidae Fam. Geogarypidae Fam. Larcidae Fam. Cheiridiidae Fam. Pseudochiridiidae Fam. Olpiidae Fam. Garypinidae Fam. Menthidae Fam. Sternophoridae Fam. Withiidae Fam. Cheliferidae Fam. Atemnidae Fam. Chernetidae Order Araneae Suborder Mesothelae Fam. Liphistiidae Suborder Orthothelae Infraorder Mygalomorphae Fam. Microstigmatidae Fam. Hexathelidae (Macrothelinae) Fam. Dipluridae (Euagrinae) Fam. Nemesiidae Fam. Theraphosidae Fam. Atypidae Fam. Antrodiaetidae Fam. Cyrtaucheniidae Fam. Idiopidae Fam. Ctenizidae Fam. Migidae Infraorder Araneomorphae Fam. Archaeidae Fam. Hypochilidae Fam. Austrochilidae Fam. Filistatidae Fam. Drymusidae Fam. Scytodidae Fam. Sicariidae Fam. Leptonetidae Fam. Ochyroceratidae Fam. Telemidae Fam. Pholcidae Fam. Caponiidae Present Present Absent Present Present Present Present (South Africa) Present (Socotra) Present Present Present Present Present Present Absent Absent Present Present Present (South Africa) Present Present Present Present India, Nepal Present Present Present (Israel) India Present Present Present Present Present Present Present Present Present Absent Present Present Present (Southern Africa) Present Present Absent Present Present Present (Southern Africa) Present Present Present (Southern Africa) Absent Present (Southern Africa) Present Present (South Africa) Present Present Absent Present Present Present Present Present Present Present Present Present Present (Japan) Present Present Present Absent Present Absent Present Absent Present Absent Present Present Present ? Present (China) Present Present Absent (continued) 686 8 Some Peculiarities of the Distribution of Arachnida Table 8.4 (continued) Fam. Tetrablemmidae Fam. Dysderidae Fam. Oonopidae Fam. Orsolobidae Fam. Segestriidae Fam. Eresidae Fam. Hersiliidae Fam. Oecobiidae Fam. Palpimanidae Fam. Mimetidae Fam. Deinopidae Fam. Uloboridae Fam. Anapidae Fam. Araneidae Fam. Cyatholipidae Fam. Linyphiidae Fam. Sinopimoidae Fam. Symphytognathidae Fam. Synaphridae Fam. Tetragnathidae Fam. Theridiidae Fam. Theridiosomatidae Fam. Ctenidae Fam. Lycosidae Fam. Oxyopidae Fam. Pisauridae Fam. Psechridae Fam. Trechaleidae Fam. Udubidae Fam. Zoropsidae Fam. Agelenidae Fam. Amaurobiidae Fam. Anyphaenidae Fam. Cybaeidae Fam. Desidae Fam. Dictynidae Fam. Hahniidae Fam. Sparassidae Fam. Selenopidae Fam. Zodariidae Fam. Chummidae Fam. Clubionidae Present Present Present Present (Southern Africa) Present Present Present Present Present Present Present Present Present Present Present Present Absent Present Absent (but present on Madagascar) Present Present Present Present Present Present Present Absent Absent Present Present (South Africa) Present Present Present (South Africa) Absent Present Present Present Present Present Present Present (South Africa, end.) Present India Absent Present Absent Present Present Present Present Present Present Present Present Present Present Present Present China (doubtful status) Present (one sp. in Japan) Present Present Present Present Present Present Present Present Present Present Absent Present Present Present Present Present Present Present Present Present Present Present Absent Present (continued) 8.10 Arachnida in the Afrotropical and Palearctic Regions 687 Table 8.4 (continued) Fam. Miturgidae Fam. Phyxelididae Fam. Titanoecidae Fam. Ammoxenidae Fam. Cithaeronidae Fam. Gallieniellidae Fam. Gnaphosidae Fam. Prodidomidae Fam. Trochanteriidae Fam. Philodromidae Fam. Thomisidae Fam. Salticidae Fam. Corinnidae Fam. Liocranidae Order Opilioacarida Fam. Opilioacaridae Order Holothyrida Order Ixodida Fam. Argasidae Fam. Ixodidae Fam. Nuttalliellidae Order Mesostigmata Order Sarcoptiformes Order Trombidiformes Present Present Absent Present (Southern Africa) Present Present Present Present Present Present Present Present Present Present Present Present Absent (only Seychelles) Present Present Present Present (Southern Africa) Present Present Present Present Present Present Absent Present Absent Present Present Present Present Present Present Present Present Present Present Absent Present Present Present Absent Present Present Present In his analysis of the distribution of Mammals in the Himalayan ranges, Hoffmann (2001) concludes that, because of the strong altitudinal gradient, “Palearctic elements dominate higher, and Indomalayan elements, lower elevations” (being in the Himalaya almost equally represented). Further Hoffmann (2001) analyzes in details the transitional zone in such complicated area as North Burma, Sichuan, and Yunnan (Map 8.2). Concerning the insects, Krijanovskij (2002) accepted the existence of an IndoMalayan Region with three subregions: Malayan (Malacca, the Sunda Islands, and the Philippines), Indian (most of Hindustan and Sri Lanka), and Indo-Chinese (Assam, Manipur, Indo-China without Tenasserim, Malacca, and the high Northern Burma, also the tropical South China and the islands Hainan, Taiwan, and Ryukyu). According to the catalogue of Fet (1988) and other sources, from the present territory of Russia (including Crimea) are known five spp. of scorpions. The neighboring Kirghizstan has also only the same two Mesobuthus (caucasicus and eupeus), but Tajikistan, situated near the scorpion-rich (29 spp.) Afghanistan, has already 6 spp., including Hottentotta, Orthochirus, Psammobuthus, and Pseudochactas. Gromov (2001) outlined the northern limit of the distribution of Scorpiones in Central Asia. From the area in the Palearctic East from Yenisei River are known 688 8 Some Peculiarities of the Distribution of Arachnida scorpions of the families Euscorpiidae, Buthidae, and Hemiscorpiidae (Russia East of Yenisei, North China, Mongolia, Korea, and Palearctic Japan). In Mongolia live only three species of genus Mesobuthus Vachon (M. eupeus, M. caucasicus, M. martensii). In China north of Yangtze are recorded very few scorpions. In Korea are recorded only two scorpions: Mesobuthus martensii and the widespread Liocheles australasiae. Japan has also two widespread species – the same Liocheles australasiae and Isometrus maculatus. The scorpiofauna of Pakistan reflects the transitional fauna of this country: 46 species of the families Buthidae (Androctonus Ehr., Baloorthochirus Kov., Buthacus Birula, Buthoscorpio Werner, Compsobuthus Vachon, Hemibuthus Pocock, Hottentotta Birula, Isometrus Ehr., Lychas C.L. Koch, Mesobuthus Vachon, Odontobuthus Vachon, Orthochirus Karsch, Plesiobuthus Pocock, Sassandiothus Farzanpay, Vachonus Tikader et Bastawade), Euscorpiidae (Euscorpiops Vachon, Scorpiops Vachon), Hemiscorpiidae (Hemiscorpius Peters), and Scorpionidae (Heterometrus Ehr.). 8.11 The Boundary Across China According to Corbet (1978), the Yangtze River was “…just beyond the southern limit of the [Palearctic] region,” and further “…in lowland China the boundary is taken very arbitrarily as Latitude 35oN, corresponding in part with the Hwang Ho (Yellow River).” The broad area between these two rivers has warm temperate climate (Fan 1990) and transitional zone between the two realms. Corbet and Hill (1992) confirm the idea that the arbitrary northern boundary between the Palearctic and the Indomalayan region is about 35oN (Yellow River). Zhang and Zhao (1978) placed the median line “a little south of Yangtze.” Again Corbet and Hill (1992) defined three divisions of the transitional zone between the Hwang Ho and the Yangtze (see Hoffmann 2001 for details). All these subdivisions were based on vertebrates, a group with many anthropogenic changes during the last centuries. It would be interesting to compare this discussion with the results obtained by a detailed analysis on the distribution of all orders of Arachnida (so far much less known in this area). 8.11.1 The Transitional Zone We discussed already the map and the analysis on mammals (Hoffmann 2001). Recently Morrone (2015) published a biogeographic map, showing the “Chinese Transitional Zone.” A higher category has been assigned to this (or a similar) zone in the subdivision of Holt et al. (2013) – an independent “Sino-Japanese Realm.” Most families (93) of Arachnida are in common to both regions. There are no endemic orders or suborders in any of them. What concerns the Arachnida is that 8.11 The Boundary Across China 689 their distribution does not justify the sharp difference between the two kingdoms (Paleotropical and Holarctic) in Eastern Eurasia. The transitional zone (SinoJapanese Realm) of Holt et al. (2013) also does not satisfy the criteria for outlining an area on the same footing as Palearctic and Indomalayan Realms. Cluster analyses, based again on terrestrial vertebrates (He et al. 2017), revealed nine major zoogeographical regions in China: South China, the Yungui Plateau, Taiwan, North China, the Inner Mongolia Plateau, Northwest China, the Longzhong Plateau, the Tibetan Plateau, and East Himalaya. He et al. (2017) suggest a major south-north-oriented Palearctic-Oriental boundary in China. Other sources: Palestrini and Zunino (1986), Palestrini et al. (1986). 8.11.2 Arachnogeography of Himalaya Typical for the Himalaya is the giant mountain system on the limit or transition between two regions and even kingdoms. The prominent arachnologist and explorer of the Himalaya, Jochen Martens, analyzed in several articles the distribution of the Palearctic and Indomalayan elements in the mountain (Martens 1973, 1983, 1984, 1987, 1993). The arachnofauna of Himalaya was analyzed by Beron (2001a, 2001b, 2008), Mani (1959), and others. Map 8.3 The transitional zone between Palearctic and Indomalayan regions (After Hoffmann, 2001, modified) 690 8.11.3 8 Some Peculiarities of the Distribution of Arachnida East Asiatic (Himalayan-Chinese) Region of Lopatin (1980) Although much smaller (the extratropical parts of Southeast Asia, including parts of the Himalaya, Tibet, Northeast China, Primorye, Korea, and the three bigger islands of Japan South of Hokkaido), this area is very interesting and contains some “southern” elements like Uropygi (Typopeltis). Still missing are Amblypygi, Schizomida, Palpigrada, Opilioacarida, and Holothyrida; there are very few scorpions (Mesobuthus, Liocheles). Some of the arachnids of Japan are clearly relicts (opilions like Suzukiellus, Yuria, spiders of suborder Mesothelae); some genera and even species (Caddo agilis), as well as the subfamily Paranonychinae, show affinities with the Nearctic fauna. The climate of this area is very different from the climate of the European-Obian and the Angaran Subregions. The lowlands have mild, even warm climate, the proximity of the ocean is causing high humidity, and the forests are broadleaf, some of them even evergreen. Of course, the high mountains have many belts with different climates. In the area there are many. 8.12 The Cape Arachnofauna 8.12.1 Africa South of Zambezi-Kunene 8.12.1.1 Geography, General Zoogeography, and Paleogeography This arbitrary-outlined area includes South Africa, Lesotho, Swaziland, Botswana, Namibia, Zimbabwe, and Mozambique (south of Zambezi). Southern Africa has a wide diversity of ecoregions including grassland, bushveld, karoo, savannah, and riparian zones. It has complex plateaus that create massive mountain structures along the South African border. The highest mountains are Drakensberg (Mafadi, 3450 m) in South Africa and Lesotho and Brandberg (2606 m) in Namibia. The highest peaks in South Africa not part of the Drakensberg Escarpment are located in the Karoo range in the southwestern part of the country. The two highest peaks in the Karoo are Compassberg and Seweweekspoortpiek, with summit elevations of 2504 m and 2325 m, respectively (Killick 1978; Mahaney (Ed.)1989; Martin 1961; Moreau 1952, 1963; Taylor 1978; White 1978; van Zinderen Bakker 1975). 8.12 The Cape Arachnofauna 691 Map 8.4 Map of Southern Africa The Mediterranean climates are restricted to the extreme southwestern Cape. The Cape flora is so distinct that it is recognized as a floral kingdom of its own (Capensis). This is justified by the richness of species, the high degree of endemism (especially of families), and the disjunct distribution of many of its members (Werger 1978). Arachnogeography Southern Africa is very different botanically from the continent more to the North, so different, that is, considered a kingdom. Much less are the differences in the fauna – usually we speak about Cape Subregion of the Afrotropical Region. This land of deserts, mountains, and an analogue of the Mediterranean climate still contains many endemic groups, mostly on a level genus. There are many riddles in this southern end of Africa – connections with other southern lands and sharp difference with the fauna of Madagascar and the Mascarene. The arachnofauna of southern parts of Africa is rich and relatively well studied by people like Lawrence, Beier, Prendini, Starega, and other researchers. We now 692 8 Some Peculiarities of the Distribution of Arachnida know that in the area south of Kunene-Zambezi are present the orders Palpigradi (1 end. sp.), Solifugae (the richest part of the world that concerns this order), Scorpiones (140 spp., 1 end. subfamily), Opiliones (3 end. genera of Cyphophthalmi, 1 end. subfamily of Eupnoi and several endemic genera, 1 end. subfamily in Laniatores, and 164 spp., all South African genera are endemic), Pseudoscorpiones (122 spp., 73% endemic, 14 end. genera), Amblypygi (rich fauna with 1 endemic genus), and Schizomida (1 endemic subfamily, all species are endemic). Araneae (64 families, 2 of them endemic), Opilioacarida (2 spp., part of a genus known in Madagascar), Ixodida (1 extremely interesting endemic family, known from Tanzania, South Africa, and Namibia). There are endemic genera also in group like Prostigmata and other Acari. Lacking from Southern Africa are the orders Ricinulei and Holothyrida and the suborders Palaeoamblypygi, Mesothelae, and Dyspnoi (Haddad 2004; Lawrence 1955, 1963) (Monod 1957; Newlands 1978; Werger 1978). A total of 821 spp., 296 genera, and 69 families of non-acarine arachnids (Araneae, Solifugi, Scorpiones, Pseudoscorpiones, Opiliones, and Amblypygi) were known from Namibia (Griffin 1998); now (2017) they are certainly more. Palpigradi Eukoenenia lawrencei Remy, 1957, is the only species of this order, known from the described area. Solifugae The area under consideration (South Africa, Lesotho, Namibia, Swaziland, Botswana, Zimbabwe, southern part of Mozambique) is extremely rich in solpugids. Here are represented 6 of the 12 families (incl. 17 end. genera) and many of the 153 known genera of Solifugae, including the endemic family Hexisopodidae and the endemic genera Hexisopus (15 spp.) and Chelypus (9 spp.) (Dippenaar-Schoenman and González Reyes 2006; Hewitt 1919; Lawrence 1955; Purcell 1903; Roewer 1941; Wharton 1981). With 127 spp. of Solifugae, 47 (37%) of which are endemic and 9 genera occurring only in Namibia, this country appears to be the richest in the world in Solifugae (Griffin 1998). An older, but most reliable, source of information on the group is the checklist of Lawrence (1955), listing from this area 181 species of 33 genera and 6 families. Wharton (1981) outlined the solpugids of Namibia. According to Harvey (2013b), in South Africa are recorded 163 spp. of Solifugae of 18 genera and 6 families. The families Gyllipidae and Rhagodidae are shared with Asia, Solpugidae only with the rest of Africa, Ceromidae with southern part of tropical Africa, and only Daesiidae also with South America. Fossil Ceromid is recorded from Brazil. Endemic genera of Solifugae south of Zambezi-Kunene: Fam. Ceromidae – Ceromella Roewer (three spp.), Toreus Purcell (one sp.) Fam. Gyllipidae – Bdellophaga Wharton (one sp.), Trichotoma Lawrence (three spp.) Fam. Hexisopodidae – Hexisopus Karsch (15 spp.), Chelypus Purcell (9 spp.) Fam. Daesiidae – Namibesia Lawrence (one sp.) 8.12 The Cape Arachnofauna 693 Fam. Melanoblossiidae (Melanoblossiinae end. in Southern Africa) – Daesiella Hewitt (one sp.), Lawrencega Roewer (seven spp.), Melanoblossia Purcell (four spp.), Microblossia Roewer (one sp.), Unguiblossia Roewer (two spp.) Solpugidae – Metasolpuga Roewer (one sp.), Prosolpuga Roewer (one sp.), Solpuguna Roewer Scorpiones The checklist of Lawrence (1955, with additions) contains 104 spp. of scorpions from Southern Africa with many subspecies. More recent checklist was provided by Prendini (2005). The scorpions south of Kunene-Zambezi belong to 14 genera and to the present-day families Bothriuridae (Brandbergia Prendini, Lisposoma Lawrence) and Buthidae. An important review of the scorpions of Namibia belongs to Lamoral (1979). In the monograph find place two families (Buthidae and Scorpionidae) and seven genera (Buthotus, Karasbergia, Parabuthus, Uroplectes, Hadogenes, Lisposoma, Opisthophthalmus). The conclusions of Lamoral are that “The scorpion fauna of Namibia is derived from elements that originated in Laurasia during Pangean times. These elements migrated overland to the North African region of Gondwanaland and ultimately to southern Africa in the wake of the southward shift of the warm tropical belt. …No affinities could be found between the scorpion fauna of either Namibia or southern Africa and South America.” “Namibia has the highest species richness and endemism of scorpions in southern Africa. All four families, seven genera (64%), and 57 sp. (44%) of southern African scorpions occur within its borders, of which one genus and at least 27 species (21%) are endemic” (Prendini, 2003a). After the description of Brandbergia Prendini, 2003, the endemism becomes even higher but still cannot compare with Madagascar (100%). African, Australian, and the only Indian member of Bothriuridae could be considered as palaeoendemics (many authors, generalized by Prendini (2003a)). Most Bothriurids live in South America, so Bothriuridae displays a purely Gondwanan distribution, as do Scorpionoidea as a whole. The survey of Prendini (2005) on the scorpions of Southern Africa contains data on 13 genera and 140 spp. Pseudoscorpiones The checklist of Dippenaar-Schoenman and Harvey (2000) enumerates 135 species from South Africa, belonging to 65 genera and 15 families (Beier 14 papers from 1931 to 1970, Ellingsen 1912; Lawrence 1935; Newlands 1978). Endemic genera in South Africa (incl. Lesotho, Botswana, Namibia, and Swaziland): Fam. Chthoniidae Selachochthonius Chamberlin, 1929 – South Africa, Lesotho (three spp.) Fam. Garypidae Ammogarypus Beier, 1962 – Namibia, South Africa (three spp.) Eremogarypus Beier, 1955 – Namibia (four spp.) Meiogarypus Beier, 1955 – Namibia (one sp.) 694 8 Some Peculiarities of the Distribution of Arachnida Fam. Gymnobisiidae Gymnobisium Beier, 1931 – South Africa, Lesotho (two spp.) Fam. Ideoroncidae Botswanoncus Harvey et Du Preez, 2014 – Botswana (one sp.) Fam. Cheliferidae Aperittochelifer Beier, 1955 – South Africa (five spp.) Litochelifer Beier, 1948 – South Africa (one sp.) Rhopalochelifer Beier, 1964 – South Africa (one sp.) Fam. Chernetidae Myrmochernes Tullgren, 1907 – South Africa (one sp.) Fam. Withidae Afrowithius J.C. Chamberlin, 1931 – South Africa (one sp.) Cyrtowithius Beier, 1955 – Namibia, South Africa (two spp.) Paragoniochernes Beier, 1932 – South Africa (two spp.) Parallowithius Beier, 1955 – Namibia, South Africa (two spp.) Pseudatemnus Beier, 1947 – South Africa (one sp.) Very high endemism on species level: Ninety-seven species (73%) are known only from South Africa, 33 spp. are more widespread in the Afrotropical Region, and 3 spp. are cosmopolitan. In the paper of Harvey and Du Preez (2014) are described a new genus and species of cave pseudoscorpion from Botswana (Botswanoncus ellisi, Ideoroncidae). Opiliones A “preliminary” analysis of the zoogeography of Southern African Opiliones was made by Starêga (1989a). According to him, south of KuneneZambezi live 217 nominal species of Opiliones and “the largest part of this fauna (63,6%) is of Gondwanan origin. Other groups represented belong to the Paleotropical (19,8%) and Mediterrano-Afrotropical element (16%).” Since this analysis, more species have been added (Lotz 2009; de Bivort and Giribet 2010), or now (2017) the species are 225 (222 endemic) from 53 genera and 9 families (Hansen and Soerensen 1904; Kauri 1961; Lawrence 1931, 1933, 1934, 1938, 1963; Rosas Costa 1950; Starêga 1984, 1992, 2008). Starega (1989a) concludes that “the largest part of the southern African harvestmen fauna shows clear relationship with the faunas of other remnants of the former Gondwana.” Cyphophthalmi In the recent paper of de Bivort and Giribet (2010) have been described 7 new species, bringing the number of South African Cyphophthalmi to 15 spp. of 3 endemic genera (Purcellia Hansen et Soerensen, 1904 [syn. Speleosiro Lawrence, 1931], Parapurcellia Rosas Costa, 1950) and 1 family – Pettalidae, shared with Madagascar, Chile, Sri Lanka, Australia, and New Zealand. 8.12 The Cape Arachnofauna 695 Eupnoi. In the area have been recorded 46 spp. of 5 genera of Caddidae, Phalangiidae, and Neopilionidae (Lotz 2009). Endemic genera are: Fam. Caddidae: Caddella Hirst, 1925 – Six spp. Fam. Neopilionidae Neopilioninae – Endemic subfamily (three spp.) Neopilio Lawrence, 1931 – Two spp. Vibone Kauri, 1961 – One sp. Dyspnoi – Not recorded in the described area Laniatores In the area have been recorded 164 spp. from 45 genera and 5 families: Triaenonychidae, Assamiidae, Samoinae, Trionyxellidae, and Biantidae. Family Triaenonychidae (from the six subfamilies, only Adaeinae is endemic in Southern Africa) is “the most important component of the South African fauna” (Staręga 1989a) with 125 spp. (more than 50% of all Opiliones in this area). All South African genera of Triaenonychidae are endemic. Adaeulum Roewer, 1914 (12 spp.); Amatola Lawrence, 1931 (6 spp.); Austromontia Lawrence, 1931 (6 spp.); Austronuntia Lawrence, 1931; Biacumontia Lawrence, 1931 (7 spp.); Cryptadaeum Lawrence, 1931 (1 sp.); Graemontia Lawrence, 1931 (6 spp.); Larifuga Loman, 1898 (10 spp.); Larifugella Lawrence, 1933 (5 spp.); Lawrencella Strand, 1932 (1 sp.); Lizamontia Kury, 2004 (4 spp.); Mensamontia Lawrence, 1931 (2 spp.); Micradaeum Lawrence, 1931 (1 sp.); Micromontia Lawrence, 1939 (1 sp.); Monomontia Lawrence, 1931 (17 spp.); Montadaeum Lawrence, 1931 (1 sp.); Paradaeum Lawrence, 1931 (1 sp.); Paramontia Lawrence, 1934 (2 spp.); Roeweriana Lawrence, 1934 (1 sp.); Rostromontia Lawrence, 1931 (3 spp.); Speleomontia Lawrence, 1931 (1 sp.); Yulella Lawrence, 1939 (1 sp.) Fam. Assamiidae Cryptopygoplus Lawrence, 1931 – Three spp. Polycoryphus Loman, 1902 – Namibia, South Africa (one sp.) Umtaliella Lawrence, 1934 – Zimbabwe (one sp.) Fam. Biantidae Biantessus Roewer, 1949 (two spp.) Cryptobiantes Kauri, 1962 (one spp.) Amblypygi In the catalogue of Lawrence (1955) are listed three species of Amblypygi from Southern Africa: Damon variegates (Perty, 1834) (Damoninae), Phrynichus scullyi (Purcell, 1901), and Ph. bacillifer (Gerstächer, 1873) (Phrynichidae). Weygoldt (1996, 1999) added the endemic genus and species Phrynichodamon scullyi Weygoldt and the new species Damon brachialis, among others. 696 8 Some Peculiarities of the Distribution of Arachnida Prendini et al. (2005) described another new species (Damon sylviae) and provided a table of all 11 spp. in genus Damon and a key with the 5 species from Southern Africa: Damon annulatipes, D. brachialis, D. gracilis, D. variegates, and D. sylviae. Schizomida Endemic subfamily (Megaschizominae), genus (Megaschizomus Lawrence, 1969), and two species – M. mossambicus (Lawrence) and M. zuluanus (Lawrence) – are the only Schizomida south of Kunene-Zambezi (Lawrence 1947, 1955, 1969). Araneae In Southern Africa are recorded spiders of 64 families, 2 of them endemic (Griffin and Dippenaar-Shoeman 1992; Griswold and Platnick 1987; Haddat et al. 2009; Hewitt 1916, 1917; Huber 2003; Jocqué 2001; Lawrence 1938; Purcell 1902; Scharff 1992; Tucker 1923). South of Zambezi-Kunene have been recorded at least 105 endemic genera and 2 endemic families of spiders. Endemic spider supraspecific taxa in Southern Africa: Fam. Anapidae Dippenaaria Wunderlich, 1995 – South Africa (one sp.) Fam. Theraphosidae Brachionopus Pocock, 1897 – South Africa (five spp.) Ceratogyrus Pocock, 1897 – Botswana, South Africa, Zimbabwe, Malawi, Mozambique, Namibia Harpactira Ausserer, 1871 – South Africa, Namibia (16 spp.) [Harpactirella Purcell, 1902 – South Africa (11 spp.), ? Morocco (one sp.) Idiothele Hewith, 1919 – Southern Africa (two spp.) Trichognathiella Gallon, 2004 – South Africa (one sp.) Fam. Chummidae – Endemic family (two spp.) Chumma Jocqué, 2001 – (two spp.) Fam. Microstigmatidae (the other genera are known from South America) Microstigmata Strand, 1932 – South Africa (six spp.) Fam. Gallieniellidae Austrachelas Lawrence, 1938 (nine spp.), Drassodella Hewitt, 1916 (seven spp.) – South Africa 8.12 The Cape Arachnofauna 697 Fam. Orsolobidae Afrilobus Griswold et Platnick, 1987 (three spp.), Azanialobus Griswold et Platnick, 1987 (one sp.) Fam. Amaurobiidae Chresiona Simon, 1903 (three spp.); Obatala Lehtinen, 1967 (one sp.); Pseudauxinum Simon, 1902 – South Africa Fam. Dictynidae Shango Lehtinen, 1967 – South Africa (one sp.) Fam. Cyrtaucheniidae Homostela Simon, 1892 – South Africa (five spp.) Fam. Trachelidae Afroceto Lyle et Haddad, 2010, Southern Africa (16 spp.); Fuchiba Haddad et Lyle, 2008, South Africa, Botswana, Lesotho, Mozambique (6 spp.); Fuchibotulus Haddad et Lyle, 2008, South Africa, Mozambique (3 spp.); Poachelas Haddad et Lyle, 2008, South Africa, Zimbabwe (4 spp.); Spinotrachelas Haddad, 2006, South Africa (5 spp.) Fam. Caponiidae Diplogena Purcell, 1904 – South Africa, Namibia, Botswana (six spp.) Fam. Mysmenidae Isela Griswold, 1985 – South Africa (one sp.) Fam. Oecobiidae Paroecobius Lamoral, 1981 – South Africa, Botswana (two spp.) Uroecobius Kullmann et Zimmermann, 1976 – South Africa (one sp.) Fam. Zodariidae Caesetius Simon, 1893, South Africa, Namibia, Mozambique, Malawi (ten spp.); Cicynethus Simon, 1910, South Africa, Namibia (three spp.); Heradida Simon, 1893, South Africa, Namibia (six spp.); Procydrela Jocqué, 1999, South Africa (two spp.); Psammodnon Jocqué, 1991, South Africa, Namibia (three spp.); Psammorygma Jocqué, 1991, South Africa, Namibia; Rotundrela Jocqué, 1999, South Africa (two spp.); Thaumastochilus Simon, 1897, South Africa (two spp.) Fam. Zoropsidae Griswoldia Dippenaar-Schoeman et Jocqué, 1997 – South Africa (12 spp.) Phanotea Simon, 1896 – South Africa (13 spp.) 698 8 Some Peculiarities of the Distribution of Arachnida Fam. Pisauridae Tapinothelella Strand, 1909, South Africa (one sp.); Voraptipus Roewer, 1955, Mozambique (one sp.); Walrencea Blandin, 1979, South Africa (five spp.) Fam. Oonopidae Australoonops Hewitt, 1915, South Africa (one sp.); Calculus Purcell, 1910, South Africa (one sp.); Dalmasula Platnick et al., 2012, South Africa, Namibia (five spp.); Nephrochirus Simon, 1910, Namibia; Pseudoscaphiella Simon, 1907, South Africa (one sp.) Fam. Eresidae Paradonea Lawrence, 1968 – South Africa, Zimbabwe, Botswana, Namibia (five spp.) Seothyra Purcell, 1903 – Namibia, Botswana, South Africa, Angola (13 spp.) Fam. Araneidae Ideocaira Simon, 1903 (three spp.), Nemospiza Simon, 1903 (one sp.) – South Africa Fam. Ammoxenidae Ammoxenus Simon, 1893 – South Africa, Namibia, Botswana (six spp.) Rastellus Platnick et Griffin, 1990 – South Africa, Namibia, Botswana, Zimbabwe (seven spp.) Fam. Palpimanidae Ikuma Lawrence, 1938 – Namibia (two spp.) Fam. Penestomidae – Endemic family! Penestomus Simon, 1902 – South Africa, Lesotho (three spp.) Fam. Theridiidae Chorizopella Lawrence, 1947 (one sp.), Histagonia Simon, 1895 (one sp.) – South Africa Fam. Liocranidae Coryssiphus Simon, 1903 – South Africa (three spp.) Fam. Clubionidae Carteroniella Strand, 1907 – South Africa (one sp.) Fam. Prodidomidae Austrodomus Lawrence, 1947 (two spp.); Eleleis Simon, 1893 (one sp.); Purcelliana Cooke, 1964 (one sp.) – South Africa Namundra Platnick et Bird, 2007 – Namibia, Angola (four spp.) 8.12 The Cape Arachnofauna 699 Theuma Simon, 1893 – South Africa, Namibia (25 spp.) Fam. Idiopidae Ctenolophus Purcell, 1904, Segregara Tucker, 1917 – South Africa Galeosoma Purcell, 1903, Gorgyrella Purcell, 1902 – Southern Africa Fam. Miturgidae Parapostenus Lessert, 1923 – South Africa, Lesotho (one sp.) Fam. Phyxelididae Lamaika Griswold, 1990 (one sp.); Malaika Lehtinen, 1967 (two spp.); Matundua Lehtinen, 1967 (one sp.); Namaquarachne Griswold, 1990 (five spp.); Pongolania Griswold, 1990 (two spp.); Themacrys Simon, 1906 (five spp.); Vidole Lehtinen, 1967 (five spp.) – South Africa Xevioso Lehtinen, 1967 – South Africa, Malawi, Zimbabwe, Swaziland (nine spp.) Fam. Hersiliidae Tyrotama Foord et Dippenaar-Shoeman, 2005 – South Africa, Namibia, Angola (eight spp.) Fam. Eutrichuridae Lessertina Lawrence, 1942 – South Africa (two spp.) Fam. Microstigmatidae Microstigmata Strand, 1932 – South Africa (six spp.) Fam. Cyatholipidae Ilisoa Griswold, 1987 (three spp.), Ubacisi Griswold, 2001 – South Africa Fam. Nemesiidae Lepthercus Purcell, 1902; Pionothele Purcell, 1902; Spiroctenus Simon, 1889 – South Africa Fam. Linyphiidae Afribactrus Wunderlich, 1995 (one sp.); Lucrinus O.P.-Cambridge, 1904; Erigonops Scharff, 1990 (one sp.) – South Africa Fam. Sparassidae Arandisa Lawrence, 1938 (one sp.); Carparachne Lawrence, 1962 (two spp.); Mierorchestris Lawrence, 1962 (two spp.); Orchestrella Lawrence, 1965 (two spp.); Palystella Lawrence, 1928 (four spp.) – Namibia Panaretella Lawrence, 1937 (five spp.), Parapalystes Croeser, 1996 (five spp.) – South Africa 700 8 Some Peculiarities of the Distribution of Arachnida Fam. Lycosidae Minicosa Alderweireldt et Jocqué, 2007 – South Africa Fam. Gnaphosidae Smionia Dalmas, 1920 – South Africa Titus O.P.-Cambridge, 1901 – Zimbabwe Trephopada Tucker, 1923, Trichothyse Tucker, 1923 – South Africa, Namibia Fam. Philodromidae Hirriusa Strand, 1932 – South Africa, Namibia (three spp.) Fam. Thomisidae Avelis Simon, 1895 – South Africa (one sp.) Stiphropella Lawrence, 1952 – South Africa (one sp.) Fam. Salticidae Araegeus Simon, 1901 – South Africa, Mozambique Ureta Wesolowska et Haddad, 2013 – South Africa Zulunigma Weselowska et Cumming, 2011 – South Africa (one sp.) Fam. Corinnidae Austrophaea Lawrence, 1952; Pronophaea Simon, 1887 (three spp.); Vendaphaea Haddad, 2009 (one sp.) – South Africa Opilioacarida Two species have been described, both in the genus Salfacarus Van der Hammen, common with Tanzania and Madagascar (S. dispar V.d. Hammen and S. lawrencei V.d. Hammen) (Van der Hammen 1977). Acariformes Trombidiformes – Prostigmata Fam. Erythraeidae Endemic genera: Kakamasia Lawrence, 1944 – South Africa Oribatida. Niedbala (2006) presents an outline of the ptyctimous mites in South Africa. Parasitiformes Ixodida From Tanzania to Namibia and South Africa lives one of the strangest ticks in the world, strange enough to form a family of its own, Nuttalliellidae, on the same level with Ixodidae and Argasidae. This is Nuttalliella namaqua Bedford, 1931, described from the Little Namaqualand. 8.14 Madagascar 701 Conclusion – Very high endemism on species level: Ninety-seven species (73%) are known only from South Africa, 33 spp. are more widespread in the Afrotropical Region, and 3 spp. are cosmopolitan (in the paper of Harvey and Du Preez (2014)). 8.13 Arachnofauna of Madagascar and the Adjacent Islands Here is outlined and compared the arachnofauna of Madagascar, the Seychelles, and the island groups Comoro, Aldabra, and Mascarene (Benoit 1977, 1979; Fage 1929, 1954; Heurtault 1986; Hoogstraal 1953; Lawrence 1959; Lourenço 1995, 1996a, b, 1999, 2000a; Lourenço et al. 2006, Millot 1948, Vachon 1979b, Vasquez and Klompen 2010). 8.14 8.14.1 Madagascar Geography, General Zoogeography, and Paleogeography The Big Island (587, 041 km2) is 419 km far from the coast of Mozambique. The highest point of Madagascar is Maromokotro (2876 m.), in the TsaratananaMassif; other higher massifs are Ankaratra (2642 m.), Andringitra (2658 m.), and others. In the last century, 95% of the native forests have been destroyed by logging and slashand-burn cultivation. The population (over 20 million, ten times more than in 1900) is growing very fast, and the unique nature of the island is under severe threat (Badyukov et al. 1989; Embleton and McElhinny 1975; Flores 1970; Furon 1949; Katz and Premoli 1979; McElhinny et al. 1976; Paulian 1951; Rabinowitz et al. 1983; Smith and Hallam 1970). The prehistoric breakup of the Gondwana supercontinent separated the Madagascar-Antarctica-India landmass from the Africa-South America landmass around 135 million years ago. Madagascar later split from India around 88 million years ago, allowing plants and animals on the island to evolve in complete isolation. In the mid-Eocene to the early Miocene, the channel may have been spanned by a land bridge allowing terrestrial mammals to move between Madagascar and the mainland. Now the Mozambique Channel is 3292 m. deep. According to Warren et al. (2010): “Corollary of island biogeographical theory is that islands are largely colonized from their nearest mainland source. Despite Madagascar’s extreme isolation from India and proximity to Africa, a high proportion of the biota of the Madagascar region has Asian affinities. This pattern has 702 8 Some Peculiarities of the Distribution of Arachnida rarely been viewed as surprising, as it is consistent with Gondwanan vicariance. Molecular phylogenetic data provide strong support for such Asian affinities, but often not for their vicariant origin; most divergences between lineages in Asia and the Madagascar region post-date the separation of India and Madagascar considerably (up to 87 Myr), implying a high frequency of dispersal that mirrors colonization of the Hawaiian archipelago in distance. Indian Ocean bathymetry and the magnitude of recent sea-level lowstands support the repeated existence of sizeable islands across the western Indian Ocean, greatly reducing the isolation of Madagascar from Asia.” The general information about Madagascar has been summarized mainly by Paulian (1951); Battistini and Richard-Vindard (1972); and Lourenço (Ed.) (1996). Map 8.5 Madagascar, Comoro Islands, and the Mascarene Arachnogeography The fauna of Madagascar has puzzled the biologists since the beginning of the zoogeography. The endemism of many groups of vertebrate and invertebrate fauna has been described many times (Dippenaar-Schoenman and González Reyes 2006; Hewitt 1919; Lawrence 1955; Purcell 1903; Roewer 1941; Wharton 1981; Millot 1948a, b). No need to repeat it here, but the recent achievement of the knowledge concerning the Arachnofauna of Madagascar is worth analyzing. 8.14 Madagascar 703 In the books on zoogeography or biogeography of the world, Madagascar is treated differently. From phytogeographic point of view, with 9 endemic families and over 300 endemic genera of plants and 85% of endemic species, the island is a region (subkingdom) on equal footing with the Afrotropical or Indomalayan regions (Takhtadjian 1970). The different treatment of the island by zoogeographers is explained further, but now is prevailing the idea of independent region (Krijanovskij 1980). What concerns the insects, Krijanovskiy (2002) accepts a special Madagascar Region, with two subregions: Madagascar Subregion and Mascarene Subregion. He writes that “The fauna of the Madagascar Region is completely comparable in the richness with the faunas of the other regions of Paleotropical Kingdom and is highly peculiar. The level of the species endemism in the majority of the insect groups under consideration is overpassing 90%, the level of generic endemism oscillates between 35 and 90%. There are several endemic taxa of higher rank – tribes and subfamilies. Not less are remarkable also the negative particularities of the fauna under consideration. The distinctions of this fauna with the faunas of the Afrotropical and the Indomalayan regions are not less than the distinction of these regions among themselves. The history of the fauna of Madagascar developed at least since the early Miocene (more probably since the Oligocene) independently from the faunas of the other regions of the Paleotropical Kingdom” (we translated these important lines, written in Russian). On the Big Island are known Palpigradi, Scorpiones, Amblypygi, Pseudoscorpiones, Opiliones, Araneae, Opilioacarida, and many other Acari. Lacking are Ricinulei, Uropygi, Solifugae (most intriguing absence!), and Holothyrida (another puzzling absence). Lacking are also the suborders Palaeoamblypygi, Mesothelae, Eupnoi (another puzzle!), and Dyspnoi. The fauna of Palpigradi (19 spp.) is the richest in the world, and the scorpions are almost 100% endemic. Palpigradi The papers of Rémy (1950, 1960a) are the base and were longtime the only source of information about Palpigradi from Madagascar (13 Eukoenenia Börner and 3 Koeneniodes Silvestri, all Eukoeneniidae). Some are endemic in the “Grande Terre,” so is Triadokoenenia Condé (1991) (Prokoeneniidae, the only endemic genus). Others have been found also on Réunion, Mauritius, and even as far as Sri Lanka (Eukoenenia chartoni, Koeneniodes madecassus) (Condé 1991, 1996). Scorpiones The immense progress in the study of scorpions of Madagascar could be assessed by comparing the three “faunas” – of Fage (1929) and Millot (1948), listing 10 species each, and of Lourenço (1996) with 19 species of 9 genera and 3 families (Buthidae, Scorpionidae, and Hemiscorpiidae – Ischnuridae). Both Fage (1929) and Millot (1948) came to the conclusion that, because of their large size, scorpions are easily collected and “we know all the populations of the island as well as their relative density and distribution” (Fage 1929). Since this optimistic (but premature) statement of the prominent French arachnologist, however, the number of species known from Madagascar has more than doubled and is far 704 8 Some Peculiarities of the Distribution of Arachnida from being exhaustive (Lourenço 1995, 1996, 2001, 2003, 2012, etc.). Microcharminae became a family and later has been downgraded again. Vachon (1969/1979) added two more species. Since the beginning of the studies of Lourenço in 1995 have been described many new species of smaller dimensions, unknown to the former researchers. Also important analysis took place, due to Lourenço (1996b, 1996e, 2000a). According to Lourenço (1996b), “The most remarkable characteristic of the scorpion fauna of Madagascar is the impressive level of endemicity, both in species and in genera. This supports the hypothesis of the very early isolation of the island from other land masses.” Fet, Gantenbein et al. (2003) concluded from molecular analysis that the endemic genus Grosphus is cladistically close to the South American genera Centruroides and Rhopalurus. The present-day (2017) list of the scorpions of Madagascar contains 52 sp., 13 genera, and 3 families (Gervais 1844; Kraepelin 1895, 1901; Fage 1929, 1946; Vachon 1969, 1979; Lourenço 1995, 1996b, 1996e, 1997, 1999, 2000a, c, 2001, 2003c, 2012c; Lourenço et al. 2004, 2006; Lourenço and Goodman 2002, 2004, 2009; Lourenço and Ythier 2010; Pocock 1889, 1890, 1893, 1894, 2002, and others). Fam. Buthidae G. ankarafantsika Lourenço, 2003 G. ankarana Lourenço et Goodman, 2003 G. annulatus Fage, 1929 G. bistriatus Kraepelin, 1901 G. darainensis Lourenço, Goodman et Ramilijaona, 2004 G. feti Lourenço, 1996 G. flavopiceus Kraepelin, 1901 G. garciai Lourenço, 2001 G. grandidieri Kraepelin, 1901 G. hirtus Kraepelin, 1901 Grosphus Simon, 1880 T. guillaumeti Lourenço, 1995 T. ivohibe Lourenço et Goodman, 1999 T. judsoni Lourenço, 1996 T. lucileae Lourenço, 1995 T. manonae Lourenço, 2000 T. monodi Lourenço, 2000 T. parriloi Lourenço, 1996 T. petrae Lourenço, 1996 T. pococki Lourenço, 1995 T. racotondravonyi Lourenço et Goodman, 2003 G. intertidalis Lourenço, 1999 Troglotityobuthus Lourenço, 2000 G. limbatus Pocock, 1889 T. gracilis (Fage, 1946) G. madagascariensis (Gervais, 1844) [Fam. Microcharmidae] G. mahafaliensis Lourenço, Goodman et Microcharmus Lourenço, 1995 Ramilijaona, 2004 G. olgae Lourenço, 2004 M. cloudsleythompsoni Lourenço, 1995 G. simoni Lourenço, Goodman et M. hauseri Lourenço, 1996 Ramilijaona, 2004 Neogrosphus Lourenço, 1995 M. jussarae Lourenço, 1996 N. blanci Lourenço, 1996 M. sabineae Lourenço, 1996 N. griveaudi (Vachon, 1969) Neoprotobuthus Lourenço, 2000 Isometrus Hemprich et Ehrenberg, 1829 N. intermedius Lourenço, 2000 I. maculatus (DeGeer, 1778) – Alien Fam. Heteroscorpiidae 8.14 Madagascar Pseudouroplectus Lourenço, 1995 P. betschi Lourenço, 1995 P. lalyae Lourenço et Ythier, 2010 P. pidgeoni Lourenço et Goodman, 1999 Palaeogrosphus Lourenço, 2000 P. copalensis (Lourenço, 1996) Tityobuthus Pocock, 1893 T. baroni (Pocock, 1890) T. darainensis Lourenço et Goodman, 2002 T. dastychi Lourenço, 1997 T. griswoldi Lourenço, 2000 705 Heteroscorpion Birula, 1903 H. goodmani Lourenço, 1996 H. kaii Lourenço et Goodman, 2009 H. magnus Lourenço et Goodman, 2002 H. opistacanthoides (Kraepelin, 1895) H. raselimananai Lourenço et Goodman, 2004 Fam. Hormuridae (Liochelidae) Opisthacanthus Peters, 1861 O. madagascariensis Kraepelin, 1894 Iomachus Pocock, 1893 I. punctulatus Pocock, 1897 Paleocheloctonus Lourenco, 1996 P. pauliani Lourenço, 1996 Endemism: Except the alien Isometrus maculatus, all 12 native genera (Grosphus, Neogrosphus, Pseudouroplectus, Tiyobuthus, Palaeogrosphus, Microcharmus, Neoprotobuthus, Pseudouroplectes, Heteroscorpion, Opisthacanthus, Paleocheloctonus, Troglotityobuthus) and 100% of the species are endemic in Madagascar, partly with the Comoros. After Goodman and Benstead (2005), the scorpions known from Madagascar are 40 spp. The family Microcharmidae, considered endemic, was synonymized with Buthidae by Volschenk et al. (2008). Despite the high endemism, several affinities can be detected. They have been hinted by the older authors (Pocock, Vachon) but analyzed in details by Lourenço (1996b, 1996e, 1999, 2000a, 2000d) and Lourenço et al. (2006). Pseudoscorpiones According to Millot (1948), in Madagascar are known 11 spp. of Pseudoscorpions of 5 families. Vachon (1951, 1960) added also the presence of Chthoniidae, Tridenchthoniidae (Dithidae), and Faella sp. (Faellidae). Now (2017, partial information from M. Harvey) from Madagascar are known 14 spp. of Pseudoscorpions of 10 genera and the families Hyidae, Feaellidae, Garypidae, Olpiidae, Pseudochiridiidae, Atemnidae, Chernetidae, and Withiidae. Endemic genera of Pseudoscorpiones of Madagascar: Fam. Garypidae Paragarypus Vachon, 1937 – One sp. Fam. Pseudochiridiidae Paracheiridium Vachon, 1938 – Two spp. (also Iles Glorieuses) The endemism is relatively low, compared to other groups. 706 8 Some Peculiarities of the Distribution of Arachnida Opiliones According to the Fauna of Lawrence (1959) and the supplements, since this time in Madagascar have been recorded Opiliones of 17 genera and 5 families: Pettalidae, Triaenonychidae, Biantidae, Samoidae, and Zalmoxidae (Fage 1945; Loman 1898; Mendes and Kury 2012; Pocock 1902, 1903; Roewer 1914, 1949; Shear and Gruber 1996; Starêga 1992). Remarkable is the lack of families like Assamiidae, widely distributed in Africa and other tropical countries of the Eastern Hemisphere (Lawrence 1959). Even more intriguing is the lack of Dyspnoi and, even more, of Eupnoi, so widespread in continental Africa. Cyphophthalmi From Madagascar have been described two (endemic and monotypic) genera: Fam. Pettalidae. Manangotria Shear et Gruber, 1996 – One sp. Incertae sedis: ? Ankaratra Shear et Gruber, 1996 – One sp. Eupnoi and Dyspnoi No native “Palpatores” are known in Madagascar. Laniatores From Madagascar are known Laniatores of 15 endemic genera and the families Biantidae, Triaenonychidae, Samoidae, Podoctidae, and Zalmoxidae. Endemic genera of Opiliones Laniatores in Madagascar are: Fam. Triaenonychidae Acumontia Loman, 1898 (syn. Triaenomontia Roewer, 1914; syn. Tanalaius Roewer, 1914; syn. Triacumontia Roewer, 1915; syn. Spinimontia Roewer, 1915; syn. Bezavonia Roewer, 1949) (22 spp.); Ankaratrix Lawrence, 1959 (two spp.); Antongila Roewer, 1931 (one sp.); Decarynella Fage, 1945 (one sp.); Hovanuncia Lawrence, 1959 (syn. Flavonuncia Lawrence, 1959) (three spp.); Ivohibea Lawrence, 1959 (one sp.); Millomontia Lawrence, 1959 (two spp.); Millotonyx Lawrence, 1959 (one sp.); Paulianyx Lawrence, 1959 (two spp.) Fam. Biantidae: Anaceros Lawrence, 1959 (four spp.), Fageibiantes Roewer, 1949 (two spp.) Fam. Samoidae: Hovanoceros Lawrence, 1959, Malgaceros Lawrence, 1959 Fam. Zalmoxidae: Bunofagea Lawrence, 1959, Remyus Roewer, 1949 Schizomida Lawrence (1969). Three (endemic?) species of Schizomida have been recorded from Madagascar (Lawrence, 1931), referred as genus Bamazomus by Harvey (2001). 8.14 Madagascar 707 Amblypygi Fam. Charinidae Charinus madagascariensis Fage, 1954 (endemic species), is the only representative of Amblypygi from the Big Island. Araneae On Madagascar are known about 400 spp. of spiders of 46 families (Griswold 1997; Blandin 1979; Emerit 1974; Griswold and Ledford 2001; Haddad et al. 2009; Huber 2000, 2003; Jocqué 1994; Lawrence 1938; Lehtinen 1967; Millot 1946, 1947; Platnick 1984, 1995; Pocock 1895; Simon 1889, 1902, 1903), but according to Alderweireldt and Jocqué (1994), this figure is only 10% of the real Malagasy spider fauna. According to Goodman and Benstead (2005), in Madagascar are known 459 spp. of spiders, 390 of which are endemic (85%). The opinion of Emerit (1978) is the same – that the “Prétendues lacunes faunistiques de la Région Malgache” are only due to the insufficient collection. We may say that Madagascar is very rich in endemics (seen well from the vertebrates), but one should be careful to judge the endemism in the invertebrates, taking into account the intensity of collecting in a given area. Emerit (1997) recorded for Madagascar the Asian subfamily Chorizopinae (Araneidae); in 1980 he recorded the family Mimetidae. Some endemic genera of spiders for Madagascar: Fam. Barychelidae (found on Madagascar, but not in Southern Africa; the genus Tigidia Simon is known from Madagascar, seven spp.; Mauritius, one; and India, four) Zophoryctes Simon, 1902 (one species) Fam. Zoropsidae (the other genera are known from America and New Zealand) Calamistrula Dahl, 1901 (one sp.) Fam. Gallieniellidae Gallieniella Millot, 1947 (three spp. in Madagascar, one sp. in Comoro Islands) Legendrena Platnick, 1984 (seven spp.) Fam. Ctenidae Mahafelytenus Silva-Dávila, 2007 (seven spp.) Fam. Hersiliidae Prima Foord, 2008 (one sp.) Fam. Theraphosidae Encyocrates Simon, 1892 (one sp.) 708 8 Some Peculiarities of the Distribution of Arachnida Fam. Ubudidae Ubuda Simon, 1880 (three spp.) Zorodictyna Strand, 1907 (two spp.) Fam. Cyatholipidae Alaranea Griswold, 1997 (four spp.) Vazaha Griswold, 1997 (two spp.) Fam. Pisauridae (incl. Halidae Jocqué, 1994, and the genera Hala Jocqué, 1994, two spp., and Tolma Jocqué, 1994, one sp.) Caripetella Strand, 1928 – Madagascar, Comoro Islands (one sp.) Paracladycnis Blandin, 1979 (one sp.) Ransonia Blandin, 1979 (one sp.) Tallonia Simon, 1889 (one sp.) Thalassiopsis Roewer, 1955 (one sp.) Fam. Synaphridae Africepheia Miller, 2007 (one sp.) Fam. Araneidae Augusta O.P.-Cambridge, 1877 (one sp.); Exechocentrus Simon, 1889 (two spp.); Parmatergus Emerit, 1994 (three spp.) Fam. Phyxelididae Ambohima Griswold, 1990 (ten spp.) Manampoka Griswold et al., 2012 (one sp.) Rahavavy Griswold et al., 2012 (three spp.) Fam. Theridiidae Asygyna Agnarsson, 2006 (two spp.) Fam. Idiopidae Genysinae Genysa Simon, 1889 (two spp.), Hiboka Fage, 1922 (one sp.) Fam. Migidae Paramiginae Micromesomma Pocock, 1895 (1 sp.); Madacantha Emerit, 1970 (1 sp.); Paramigas Pocock, 1895 (11 spp.); Thyropoeus Pocock, 1895 (2 sp.) Fam. Filistatidae Andoharano Lehtinen, 1967 8.14 Madagascar 709 Fam. Selenopidae Garcorops Corronca, 2003 – Madagascar, Comoro Islands (three spp.) Hovops Benoit, 1968 (nine spp.) Fam. Liocranidae Donuea Strand, 1932 (two spp.) Fam. Lycosidae Katableps Jocqué, Russell-Smith et Alderweireldt, 2011 (three spp.) Fam. Linyphiidae Thapsagus Simon, 1894 (one sp.), Thyreobaeus Simon, 1889 (one sp.) Tmeticides Strand, 1907 (one sp.) Fam. Oxyopidae Hostus Simon, 1898 (one sp.) Fam. Pholcidae Paramicromerys Millot, 1946 (14 spp.), Zatavua Huber, 2003a (17 spp.) Fam. Sparassidae Chrosioderma Simon, 1897 (nine spp.); Megaloremmius Simon, 1903 (one sp.); Staianus Simon, 1889 (one sp.) Fam. Zodariidae Aschema Jocque, 1991 (two spp.), Omucukia Kocak et Kemal, 2008 (two spp.) Omucutia Kocak et Kemal, 2008 (two spp.) Fam. Thomisidae Apyretina Strand, 1929 (five spp.) Cyriogonus Simon, 1886 (six spp.) Diplotychus Simon, 1903 (one sp.) Emplesiogonus Simon, 1903 (two spp.) Herbessus Simon, 1903 (one sp.) Iphoctesis Simon, 1903 (one sp.) Lampertia Strand, 1907 (one sp.) Plastonomus Simon, 1903 (one sp.) Pseudoporrhopis Simon, 1886 (one sp.) Soelteria Dahl, 1907 (one sp.) Fam. Salticidae Echinussa Simon, 1901 (three spp.) Goleta Peckham et Peckham, 1894 710 8 Some Peculiarities of the Distribution of Arachnida Macopaeus Simon, 1900 Myrmanu Prósz., 2016 Myrmele Prósz., 2016 Phaulostyleus Simon, 1902 (four spp.) Poessa Simon, 1902 Tomobella Szüts et Scharff, 2009 (two spp.) Tomocyrba Simon, 1900 (six spp.) Opilioacarida Vásquez and Klompen (2010) contributed to the study of Opilioacarids of Madagascar with another four species to the two already described by Van der Hammen (1977), all in his genus Salfacarus. Other members of this genus live on the African continent (South Africa and Tanzania). The six species are all endemic. Acariformes Oribatida In the fauna of Oribatida of Madagascar, 80% of the species are endemic. The genera are distributed evenly between the Afrotropical and the “Oriental” Regions (Schatz, 2004). Other sources: Mahunca (2002) and nine other papers; Niedbala 2004. Fauna of ptyctimous mites from Madagascar and neighbouring islands Seychelles, Mauritius, Comoros and Reunion (MAG) comprises 111 species for which morphological diagnoses have been made. …The similarity of the fauna of MAG (111 species) to the fauna of Afrotropical Region (200 species) is much higher (15 % common species) than to the fauna of Oriental Region (262 species) (4 % common species). The fauna of MAG shows no similarity to that of India. In the years 2010–2015 new soil samples (171) were collected on Madagascar and they contained 78 species. The high number of endemic species among the dominants and high-frequency species confirm the unique character of the fauna of Madagascar. (Niedbala 2017) Parasitiformes Ixodida The base of our knowledge is the paper of Hoogstraal (1953), in which find place 11 endemic or presumably endemic species of Ixodida (9 Haemaphysalis, 1 Ixodes, and 1 Argasidae indet). Here we quat the conclusions of Hoogstraal concerning the origin of the Malagasy tick fauna: Millot (1948, 1952) stated that Malagasy ticks are certainly all of African origin. This is entirely or almost entirely true for introduced species. The endemic species show some affinities with African fauna, but many more with that of southern Asia. …As in Madagascar, haemaphysalid species form the bulk of the southern Asiatic ixodid tick fauna….The very important African genus Rhipicephalus, consisting of over forty species, is entirely unrepresented by endemic species in the Malagasy Faunal Region. Amblyomma ticks, repre- 8.15 Mascarene Islands 711 sented in Africa by many more species than in Asia, are lacking in endemic species in the Malagasy Region except in the Seychelles. According to Klompen (2003) and Goodman and Benstead (2005), there are 27 spp. of Ixodida, among them 25 are endemic. 8.15 8.15.1 Mascarene Islands Geography, General Zoogeography, and Paleogeography The Mascarene Islands (or Mascarenhas Archipelago) is a group of islands in the Indian Ocean east of Madagascar comprising Mauritius, Réunion, Rodrigues, Cargados Carajos shoals, etc. The islands share a common geologic origin in the volcanism of the Réunion hot spot beneath the Mascarene Plateau and form a distinct ecoregion with a unique flora and fauna. The youngest islands to form were Mauritius (7–10 ma), the oldest of the existing islands, created along with the undersea Rodrigues ridge. The islands of Rodrigues and Réunion were created in the last 2 million years. The Mascarene Plateau is an undersea plateau that extends approximately 2000 km, from the Seychelles to Réunion. The plateau covers an area of over 115,000 km² of shallow water, with depths ranging from 8 to 150 m. The islands have never been connected to the mainland, so the flora and fauna of the Mascarenes arrived from over the sea. Prehistoric islands of the Mascarene Plateau, now disappeared under the sea, may have served as “stepping stones” which allowed species to island-hop from the Seychelles or Madagascar. Réunion – A French island, 200 km southwest of Mauritius, the nearest island. Area 2512 km2, 63 km long, 45 km wide. Highest point is Piton des Neiges (3070 m), also the shield volcano Piton de la Fournaise, more than 2631 m. Mauritius – An island 900 km. east of Madagascar, 570 km. from Rodrigues. Area – 2040 km2, highest point 828 m Rodrigues – An island, part of the Republic of Mauritius, 109 km2, highest point 355 m Arachnogeography These small strange islands with fantastic (already gone) vertebrate fauna have several orders of Arachnida, but no endemisms higher than genus. They are Palpigradi, Scorpiones (one endemic species), Pseudoscorpiones (six endemic species, two endemic genera), Opiliones (seven end. species of Laniatores, one endemic genus), Araneae (one endemic genus), and different Acari, of which most interesting is the presence of Holothyrida (three sp., incl. one endemic genus on Mauritius). Nothing similar to the fauna of Africa or Madagascar. From the Mascarene are missing the orders Ricinulei, Thelyphonida, Solifugae, and Opilioacarida and the suborders Palaeoamblypygi, Mesothelae, Cyphophthalmi, and Dyspnoi (Maureemootoo, 2000). 712 8 Some Peculiarities of the Distribution of Arachnida Palpigradi The information about the Palpigradi on these islands comes from Remy (1952b – Réunion, 1958 - Mauritius), in total three spp. (one of Eukoenenia, two of Koeneniodes), all of them belonging to Eukoeneniidae. On Réunion have been collected Eukoenenia hanseni (Silvestri), Koeneniodes frondiger Remy, and K. madecassus Remy, all found also on Madagascar and Mauritius and in Indonesia. Scorpiones Endemic sp. on Mauritius is Lychas serratus (Buthidae). Chiromachus ochropus (Hormuridae) is shared with the Seychelles and Zanzibar (Probst 1973). Pseudoscorpiones On the islands (Réunion, Mauritius) have been recorded eight species of Pseudoscorpions (Chthoniidae, Cheliferidae, Chernetidae, and Withiidae), six of them are endemic in the Archipelago, and two endemic genera (Beier 1935, 1940; Mahnert 1975). Thaumatowithius Beier, 1940 – Réunion, Mauritius (two spp.) Pilochelifer Beier, 1935 – Réunion, Mauritius (one sp.) Opiliones From Mascarene Islands are known seven genera of Opiliones (Roewer 1927, 1936, 1949, 1954; Simon 1879; Staręga 1989b). Cyphophthalmi – None Eupnoi Fam. Sclerosomatidae – Gagrella Stoliczka Dyspnoi – None Laniatores Fam. Assamiidae – Maccabeesa Roewer (one sp.) Fam. Biantidae – Hinzuanius Karsch (one sp.) Fam. Podoctidae – Erecanana Strand (one sp.), Sitalcicus Roewer (one sp.), Ibalonius Karsch (one sp.) Fam. Zalmoxidae – Zalmoxis Sørensen (Zalmoxomma Roewer) (one sp.) Endemic Opiliones on the islands are: Fam. Biantidae Hinzuanius mauriticus Roewer, 1927 – Mauritius Fam. Assamiidae Maccabeesa lawrencei Roewer, 1936 – Mauritius 8.16 Seychelles (incl. Aldabra), Comores 713 Fam. Podoctidae Erecanana insulana Roewer, 1949 – Réunion Sitalcicus novemtuberculatus Simon, 1879 – Réunion Ibalonius breoni Simon, 1879 – Réunion Fam. Zalmoxidae Zalmoxis [Zalmoxomma] occidentalis (Roewer, 1949) – Mauritius Fam. Sclerosomatidae Gagrella reunionis (Roewer, 1954) – Réunion Amblypygi From Mauritius is known Phrynichus scaber (Gervais, 1844) (Phrynichidae), known also from the Seychelles. Holothyrida From Mauritius have been described two endemic species: Holothyrus coccinella Gervais (1842), the first holothyrid ever described, and H. legendrei Van der Hammen (1983). According to the modern system of Holothyrida (Lehtinen 1995, Beron 2014), the genus Holothyrus remains endemic in Mauritius. The order Holothyrida is known also from Rodrigues. 8.16 8.16.1 Seychelles (incl. Aldabra), Comores Geography, General Zoogeography, and Paleogeography The Republic of Seychelles consists of four archipelagoes (Seychelles, Amirantes, Farquhar, and Aldabra), with some 155 islands of total surface area of 451 km2. The most interesting for arachnology are the 42 granite islands, remnants of a much bigger landmass. They lay to the northeast of Madagascar and c. 1600 km. east of Kenya (the African coast). Some scientists claim that 18 000 years ago, the sea level was 120 m lower and today’s Seychelles was part of a landmass covering c. 32 000 km2. The granite islands are very old (the oldest islands of any ocean), and careful research is being done on their flora and fauna (Benoit 1978; Badyukov et al. 1989; Gerlach et al. 2013; Scott 1958). The granite islands are situated in a radius of 56 km from the main island Mae (142 km2, highest point 914 m.). They are part of the so-called Inner islands, on which live 82 500 inhabitants. Some other bigger The Outer Islands (211 km2, 2% of the population) are coral (Amirante, Aldabra, Farcouar, and others) but are also interesting. These granitic islands are part of the Mascarene Plateau, detached from the Indian Plate 65 million years ago. 714 8 Some Peculiarities of the Distribution of Arachnida The system of national parks covers 42% of the land and 260 km2 of the sea around it. The flora of the Seychelles includes 1170 species of higher plants (75 endemic species); among the animals 15 bird species, 3 mammal species, 30 species of amphibians and reptiles, and hundreds of species of invertebrates are not found elsewhere. Starting with the words of Jeannel (1942), “Dans tous les groupes d’Insects, les monographes s’accordent pour affirmer que la faune des Seychelles a un caractère nettement indo-malais, les éléments malgaches s’y trouvant en majorité, tout élément africain faisant défaut,” we should check the rightness of this conclusion on what concerns the arachnids. The very important contribution to the fauna of the Seychelles (Séchelles) was the series of papers as a result of the mission of P.L.G. Benoit and J.J. Van Mol in 1972. The Arachnida were treated by Benoit (1978, series of papers on spiders), Mahnert (1978, pseudoscorpions), and Benoit (1979, scorpions). The results were published in Belgium under the editorship of P. Benoit, who wrote the introduction of this important series. Many other specialists have explored the strange islands and their non-insect inhabitants (see Gerlach, J. & J. Marusik, Eds, 2010). Arachnogeography The amazing fauna of the granitic islands, combining African, Indian, and endemic elements, received attention by very qualified arachnologists, and now we know that: The Seychelles are home of the orders Palpigradi, Scorpiones, Pseudoscorpiones, Opiliones (four endemic genera), Amblypygi, Schizomida, Araneae, and some groups of Acari (Gerlach 2003; Gerlach and Marusik (Eds) 2010; Hirst 1913; Harvey 2001). Absent are the orders Ricinulei, Solifugae, and Opilioacarida and the suborders Palaeoamblypygi, Mesothelae, Cyphophthalmi, Dyspnoi, and Eupnoi. To be noticed is the presence of Holothyrida (three endemic genera of the family Holothyridae), the big poisonous creatures, absent in Madagascar and in the nearby Africa. Schizomida are also very well represented by seven genera (two endemic) and seven species (five endemic). Among the 228 spp. of spiders of 40 families (Saaristo 2003), there are at least 12 endemic genera. Despite this interesting fauna, the level of endemism is only to genus, and in some orders, there are only endemic species. Palpigradi Koeneniodes madecassus is recorded from La Digue (known also from Madagascar and Indonesia) (Harvey 2001). Scorpiones From the Seychelles have been reported the species Isometrus maculatus, Lychas braueri (Scorpionidae), and Chiromachus [Ischnurus] ochropus (Hormuridae, known from Zanzibar and Mauritius). Lychas braueri (Kraepelin, 1896) is endemic in the Seychelles (Benoit 1979; Harvey 2001; Kraepelin 1896; Lourenço and Goodman 2009; Prendini 2010). Pseudoscorpiones The 9 species of Pseudoscorpiones, mentioned by Beier (1940) from the Seychelles, have been added up in 1978 to 4 more, so the figures, quoted by Mahnert (1978) and Harvey (2001, 2009, 2011), are 23 species of 18 genera and 8.16 Seychelles (incl. Aldabra), Comores 715 12 families: Chthoniidae, Tridenchthoniidae, Syarinidae, Garypidae, Garypinidae, Geogarypidae, Pseudochiridiidae, Olpiidae, Atemnidae, Feaellidae, Chernetidae, and Withiidae (Beier 1974; Chamberlin 1930; Hirst 1911). No endemic genera, seven endemic species: Fam. Tridenchthoniidae Compsaditha seychellensis Beier, 1974 – Mahé Fam. Feaellidae Feaella affinis Hirst, 1911 – Praslin, Silhouette Fam. Geogarypidae Afrogarypus seychellensis (Beier, 1940) – Praslin, Mahé Geogarypus ocellatus Mahnert, 1978 – Praslin Fam. Olpiidae Beierolpium benoiti Mahnert, 1978 – Curieuse, Silhouette (one endemic sp.) Fam. Syarinidae Ideoblothrus seychellesensis (Chamberlin, 1930) – Silhouette Fam. Atemnidae Oratemnus brevidigitatus Beier, 1940 – Mahé, Silhouette, Aride The fauna has affinities with both the Afrotropical and Indomalayan faunas. From the other ten spp. in the family Feaellidae (the only genus Feaella), eight live on the African continent (incl. four in Southern Africa), one in India, and one in Australia. Compsaditha has 12 spp. (5 from Africa, 5 from Asia and Oceania, 1 from the Seychelles, and 1 from South America). From 14 spp. of Afrogarypus, 1 is from the Seychelles and 13 from the African continent (A. impressus is known from Aldabra, Mozambique, and South Africa). Beierolpium has also 14 known species, from Africa (8), Australia, Oceania, Malaysia, and Venezuela. Oratemnus is typically Oriental – 14 spp. from Asia, 4 from Australia, 1 from the Seychelles, and 1 from Oceania and the Caribbean. Geogarypus and Ideoblothrus are widespread. Opiliones Cyphophthalmi, Eupnoi, and Dyspnoi – Not known on the Seychelles Laniatores The paper of Rambla (1984) summarizes the knowledge on the Opiliones of the Seychelles – 17 spp., all of Laniatores of 9 genera and the families Phalangodidae, Biantidae, Podoctidae, Samoidae, and Zalmoxidae (Loman 1902; Hirst 1911; Roewer 1912, 1949). 716 8 Some Peculiarities of the Distribution of Arachnida Fam. Podoctidae Centrobunus braueri Loman, 1902; Holozoster ovalis Loman, 1902; Ibalonius inscriptus Loman, 1902; I. lornani Hirst, 1911; l. bimaculatus Loman, 1902; I. karschi Loman, 1902; l. flavopictus Hirst, 1911; Sitalcicus gardineri Hirst, 1911; S. incertus Rambla, 1984 Fam. Samoidae Benoitinus elegans Rambla, 1984; Mitraceras crassipalpum Loman, 1902; M. pulchra Rambla, 1984; Samoa sechellana Rambla, 1984 Fam. Biantidae Biantes parvulus (Hirst, 1911), B. albimanus (Loman, 1902), B. minimus (Rambla, 1984) Fam. Zalmoxidae Metazalmoxis ferruginea Roewer, 1912 Endemic genera for the Seychelles are: Fam. Podoctidae Centrobunus Loman, 1902 (one sp.), Holozoster Loman, 1902 (one sp.) Peromona Roewer, 1949 (one sp.) Fam. Samoidae Benoitinus Rambla, 1984 (one sp.), Mitraceras Loman, 1902 (one sp.) Fam. Zalmoxidae Metazalmoxis Roewer, 1912 (one sp.) Amblypygi. From the Seychelles are known Phrynichus scaber (Gervais, 1844) (Phrynichidae), known also from Mauritius, and Charinus seychellarum Kraepelin, 1898 (Benoit 1979, Harvey 2001, Kraepelin 1898) (Charinidae) (endemic). Schizomida. On the Seychelles are found seven species of Schizomida. Harvey (2001) describes four new monotypic and partly endemic genera: Fam. Hubbardiidae Mahezomus apicoporus Harvey, 2001 – Endemic genus and species Secozomus latipes (Hansen, 1905) – Endemic species Zomus bagnalii (Jackson, 1908) – Known from Southeast Asia Anepsiozomus sobrinus Harvey, 2001 – Endemic genus and species Ovozomus lunatus (Gravely, 1911) – Known from the Seychelles, Cook Islands, Mayotte and other islands, Ceylon, India Bamazomus aviculus Harvey, 2001 – Endemic species Apozomus gerlahi Harvey, 2001 – Endemic species 8.16 Seychelles (incl. Aldabra), Comores 717 Three species of Schizomida, described from Madagascar, are transferred to Bamazomus Harvey: B. madagassus (Lawrence), B. milloti (Lawrence), and B. vadoni (Lawrence). As a result, on the Seychelles are known seven spp. of Schizomida of seven genera, including four endemic genera and five end. species. Araneae Many specialists (Simon, Benoit, Brignoli, and others) contributed to the study of the spiders of the fascinating islands. Gerlach (2003) quoted the figure of 198 spp. on the islands. After a series of papers (1978, 1998, 2001, 2002) on the spiders of the Seychelles, Saaristo (2003) published online a list of 228 spp. from 40 families, living on the islands (198 spp. from the Granitic Islands and 30 more species on the reefs and atolls) (Benoit 1978 – several papers, 1979; Grasshoff 1980; Wanless 1984). Endemic genera of spiders: Fam. Theraphosidae Nesiergus Simon, 1903 (three spp.) Fam. Tetrablemmidae Mariblemma Lehtinen, 1981 Fam. Ochyroceratidae Euso Saaristo, 2001, Roche Saaristo, 1998 Fam. Oonopidae – Gamasomorphinae Diblemma O. P.- Cambridge, 1908 (one sp.); Ischnothyrella Saaristo, 2001 (one sp.); Lionneta Benoit, 1979 (eight spp.); Lisna Saaristo, 2001 (one sp.); Matyotia Saaristo, 2001 (one sp.); Patri Saaristo, 2001 (one sp.); Prida Saaristo, 2001 (one sp.); Aridella Saaristo, 2002 (one sp.); Cousinea Saaristo, 2001 (one sp.); Farqua Saaristo, 2001 – Farquhar Islands – (one sp.) Fam. Gnaphosidae Microdrassus Dalmas, 1919 Fam. Tetragnathidae Pholcipes Schmidt et Krause, 1993 – Comoro Islands (one sp.) Fam. Theridiidae Argyrodella Saaristo, 2006 Bardala Saaristo, 2006 – Aldabra (one sp.) Nanume Saaristo, 2006 – Aldabra (one sp.) Sesato Saaristo, 2006 (one sp.) Seycellesa Kocak et Kemal, 2008 (one sp.) Spinembolia Saaristo, 2006 (one sp.) Stoda Saaristo, 2006 (one sp.) 718 8 Some Peculiarities of the Distribution of Arachnida Fam. Ctenidae Apolania Simon, 1898 (one sp.) Fam. Scytodidae Soeuria Saaristo, 1997 Fam. Sparassidae Pleurotus Simon, 1898 (one sp.) Rhacocnemis Simon, 1897 (one sp.) Stipax Simon, 1898 (one sp.) Thomasettia Hirst, 1911 (one sp.) Fam. Salticidae Baviola Simon, 1896 Holothyrida There are three endemic genera (Sternothyrus Lehtinen, 1995; Michaelothyrus Gerlah, Lehtinen et Madl, 2010; and Dicrognatus Gerlah, Lehtinen et Madl, 2010) in the Seychelles, one genus (Hammenius Lehtinen, 1981) is shared with New Guinea, and one more species is incertae sedis. The presence of these peculiar mites on the Seychelles and the Mascarene Islands is as curious as their absence in Africa and Madagascar (Beron 2014; Gerlah 1995; Gerlah et al. 2010; Hammen 1983; Lehtinen 1981, 1995; Thon 1906). Discussion concerning the affinities and differences in the Arachnid faunas of Madagascar and the adjacent islands: We may start with the words of Millot (1952): “Parmi les Invertébrés, les Arachnides fournissent des documents particulierement significatifs. Les espèces de la Réunion aussi bien que celles de Maurice ou de Mahé sont tout à fait différentes de celles de Madagascar: Araignées, Opilions, Scorpions, Pseudoscorpions sont presque aussi dissemblables que s’il s’agissait de régions fort separées – et alors que les formes malgaches sont d’affinités nettement africaines, les espèces des Séchelles et des Mascaraignes sont de parenté non moins nettement indo-malaise”. Palpigradi Four genera are known in the Big Island, including the endemic genus and species Triadokoenenia Condé, 1991 (Prokoeneniidae). The other genera (Eukoenenia, Prokoenenia, and Koeneniodes) are represented also on Mauritius, Réunion, and even as far as Sri Lanka, with the same species. On the Seychelles (La Digue) has been recorded Koeneniodes madecassus. Solifugae – Absent on all these islands, but numerous in East and South Africa Scorpiones Several attempts to analyze the origins and affinities of the scorpions in Madagascar and the surrounding islands (Fage 1929; Millot 1948; Legendre 1972; Vachon 1979b) were crowned by the contribution and analysis of Lourenço (1996). After this paper, many more new species have been published by Lourenço and his coauthors, but the main conclusions remain valid. To quote Lourenço (1996b): 8.16 Seychelles (incl. Aldabra), Comores 719 “…the present representatives of the Madagascar scorpion fauna are derived from protoelements of both the families Buthidae and Ischnuridae which were already present in Gondwanaland, previous to fragmentation and continental drift” The endemicity of scorpions in Madagascar is the highest in the world: 100% of the species known and in the native genera. All 12 native genera are endemic, Grosphus also on the Comores. There is also an endemic (contested) family (Microcharmidae). This high endemicity (and not only of scorpions) “supports the hypothesis of the very early isolation of the island of other land masses (Lourenço 1996). Meanwhile, in their subdivision of the world, based on scorpions, Nenilin and Fet (1992) include Madagascar, the Mascarene, and the Seychelles in the Afrotropical Region as Madagascar Subregion, divided into Madagascar Rayon and Mascarene – Seychelles Rayon. If we compare the scorpion fauna of Madagascar with Southern Africa, we may find similarities only on family level. Pseudoscorpiones From the total area under study (Madagascar, Seychelles, Aldabra, Comores, Mascarene Islands), there are relatively few species of pseudoscorpions recorded (24 genera and 14 families). No endemic families, only two endemic genera (Mauritius, Réunion). If we compare the generic lists of the various island groups, we get the following distribution: Fam. Chthoniidae Tyrannochthonius Chamberlin, 1929 – Seychelles Chthonius C.L. Koch, 1843 – Seychelles Fam. Tridenchthoniidae Compsaditha Chamberlin, 1929 – Seychelles (endemic sp.) Fam. Syarinidae Ideoblothrus Balzan, 1872 – Seychelles (one endemic sp.) Fam. Hyidae Indohya Beier, 1974 – Madagascar, India Fam. Garypidae Paragarypus Vachon, 1937 – Madagascar (one sp., endemic) Fam. Geogarypidae Afrogarypus Beier, 1931 – Aldabra, Seychelles (endemic sp.) Geogarypus Chamberlin, 1930 – Seychelles (endemic sp.) Fam. Garypinidae Aldabrinus Chamberlin, 1930 – Aldabra (one endemic sp.) Fam. Feaellidae – Madagascar Feaella Ellingsen, 1906 – Seychelles (one endemic sp.) 720 8 Some Peculiarities of the Distribution of Arachnida Fam. Olpiidae Beierolpium Heurtault, 1976 – Seychelles Xenolpium Chamberlin, 1930 – Madagascar, Seychelles, Aldabra Fam. Pseudochiridiidae Paracheiridium Vachon, 1938 – Madagascar (two spp., endemic, but also Iles Glorieuses) Pseudochiridium With, 1906 – Seychelles Fam. Atemnidae Anatemnus Beier, 1932 – Madagascar, Seychelles Oratemnus Beier, 1932 – Seychelles Paratemnoides Harvey, 1990 – Madagascar Fam. Chernetidae Haplochernes Beier, 1932 – Madagascar (one endemic sp.) Lamprochernes Tomosvary, 1882 – Seychelles Metagoniochernes Vachon, 1939 – Madagascar (endemic sp.) Fam. Withiidae Nesowithius Beier, 1940 – Seychelles Thaumatowithius Beier, 1940 – Réunion, Mauritius (two spp.) Withius Kew, 1911 – Madagascar, Seychelles Fam. Cheliferidae Pilochelifer Beier, 1935 – Réunion, Mauritius (one sp.) Heurtault (1986) writes that on Madagascar live members of 12 endemic genera of Pseudoscorpiones but did not provide a list of these genera. According to the information available, there is a striking difference between the huge endemicity in Madagascar by the scorpions (and many other groups) and the very low level by the pseudoscorpions. Opiliones According to Lawrence (1959), the most striking feature in the fauna of Opiliones is its total absence in Madagascar and the quasi total on the adjacent islands of native “Palpatores.” If we complete the table in the monograph of Lawrence (1959) with the data of Rambla (1984) and others, it will take shape as follows: Area rMadagascar Seychelles Mauritius Réunion Area Madagascar Cyphophthalmi + − − − Biantidae + Triaenonychidae + − − − Podoctidae − Phalangodidae − + − − Samoidae + Assamiidae − − + − Zalmoxidae + 721 8.16 Seychelles (incl. Aldabra), Comores Seychelles − + Mauritius + + Réunion − + Genera in common: Madagascar – Seychelles: none Seychelles – Mauritius: none Seychelles – Réunion: Ibalonius, Sitalcicus (Podoctidae) Madagascar – the Mascarene: none Réunion – Mauritius: none + − − + + − Schizomida Madagascar: Three endemic Bamazomus species Seychelles: Seven spp. known, five of them endemic, three endemic genera (Mahezomus, Anepsiozomus, Secozomus). Zomus bagnalii (Jackson) is known from Malaysia (incl. Sarawak), Singapore, and England (Kew Gardens). Mauritius: Zomus bagnallii (Jackson), Schizomus vinsoni Lawrence Rodrigues: Schizomus tenuipes Lawrence, Schizomus virescens Lawrence Réunion: Ovozomus lunatus (Gravely) Comoro Islands: Ovozomus lunatus (Gravely) (Mayotte) Amblypygi Madagascar: Charinus madagascariensis Fage, 1954 (endemic species) Mauritius: Phrynichus scaber (Gervais, 1844) Seychelles: Charinus seychellarum (Kraepelin, 1898), Phrynichus scaber (Gervais, 1844) Araneae The only family considered endemic in Madagascar (Halidae) has been merged with Pisauridae in 2006. Opilioacarida Known only from Madagascar (six endemic species of genus Salfacarus Hammen, living also in Southern Africa) Holothyrida Four species on the Seychelles (incl. endemic genus Sternothyrus Lehtinen), two endemic species of Holothyrus Gervais, endemic genus for Mauritius. The order is not known from Madagascar and Africa but is found in Sri Lanka, India, New Guinea, etc. (Table 8.5) 722 8 Some Peculiarities of the Distribution of Arachnida Table 8.5 Distribution of the arachnids in the areas concerned Group Order Palpigradi Fam. Eukoeneniidae Order Ricinulei Fam. Ricinuididae Order Solifugae Fam. Galeodidae Fam. Daesiidae Fam. Solpugidae Fam. Ceromidae Fam. Melanoblosiidae Fam. Gylippidae Fam. Hexisopodidae Order Scorpiones Fam. Bothriuridae Fam. Buthidae Fam. [Microcharmidae] Fam. Heteroscorpionidae Fam. Hormuridae Fam. Scorpionidae Fam. Lisposomidae Order Schizomida Fam. Hubbardiidae Order Uropygi Fam. Hypoctonidae Order Amblypygi Suborder Neoamblypygi Fam. Charinidae Fam. Phrynichidae Suborder Palaeoamblypygi Order Opiliones Suborder Cyphophthalmi Fam. Ogoveidae Fam. Neogoveidae Fam. Pettalidae Suborder Eupnoi Afrotropical (Southern Africa) Present Present Absent in South Africa Absent in South Africa Present Present Present Present Present Present Present Present Present Present Present Absent Absent Present Present Present Present Present Absent in Southern Africa Absent in Southern Africa Present Present Present Present Absent in Southern Africa Present Present Absent in Southern Africa Absent in Southern Africa Present Present Madagascar Present Present Absent Seychelles Present Present Absent Mascarene Islands Present Present Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Present Present Present Present Present Absent Absent Present Present Absent Absent Absent Absent Absent Absent Absent Absent Absent Present Absent Present Absent Absent Present Absent Absent Present Present Absent Absent Absent Absent Absent Absent Absent Absent Absent Present Absent Present Absent Absent Present Present Absent Present Present Absent Absent Absent Absent Present Present Present Present Absent Present Present Present Present Absent Present Present Absent Present Absent Present Present Absent Present Present Absent Present Absent Absent Absent Absent Absent Present Present Absent Present Absent Present (continued) 8.16 Seychelles (incl. Aldabra), Comores 723 Table 8.5 (continued) Group Fam. Caddidae Fam. Phalangiidae Fam. Sclerosomatidae Fam. Neopilionidae Suborder Dyspnoi Suborder Laniatores Fam. Triaenonychidae Fam. Assamiidae Fam. Biantidae Fam. Podoctidae Fam. Pyramidopidae Fam. Samoidae Fam. Zalmoxidae Order Pseudoscorpiones Suborder Epiocheirata Fam. Chthoniidae Fam. Tridenchthoniidae Fam. Pseudotyrannochthoniidae Fam. Lechytiidae Fam. Feaellidae Suborder Iocheirata Fam. Ideoroncidae Fam. Hyidae Fam. Neobisiidae Fam. Syarinidae Fam. Garypidae Fam. Geogarypidae Fam. Cheiridiidae Fam. Pseudochiridiidae Fam. Olpiidae Fam. Garypinidae Fam. Menthidae Fam. Sternophoridae Fam. Withiidae Fam. Cheliferidae Fam. Atemnidae Fam. Chernetidae Order Araneae Suborder Mesothelae Afrotropical (Southern Africa) Present Present Present Present Absent Present Present Present Present Present Present Present Absent Present Present Present Present Present Present Present Present Present Absent Present Present Present Present Present Present Present Present Absent in Southern Africa Present Present Present Present Present Present Absent Madagascar Absent Absent Absent Absent Absent Present Present Absent Present Present Absent Absent Absent Present Present Present Absent Absent Seychelles Absent Present Present Absent Absent Present Absent Absent Present Present Absent Present Present Present Present Present Present Absent Mascarene Islands Absent Absent Present Absent Absent Present Absent Present Present Present Absent Absent Present Present Present Present Absent Absent Absent Present Present Absent Present Absent Absent Absent Absent Absent Present Present Absent Absent Absent Absent Present Absent Absent Absent Present Present Present Present Absent Present Absent Absent Absent Absent Present Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Present Absent Present Present Present Absent Absent Present Present Present Present Present Absent Absent Present Present Absent Present Present Absent (continued) 724 8 Some Peculiarities of the Distribution of Arachnida Table 8.5 (continued) Group Suborder Orthothelae Infraorder Mygalomorphae Fam. Microstigmatidae Fam. Hexathelidae Fam. Dipluridae Fam. Nemesiidae Fam. Atypidae Fam. Cyrtaucheniidae Fam. Idiopidae Fam. Ctenizidae Fam. Migidae Infraorder Araneomorphae Fam. Archaeidae Fam. Hypochilidae am. Austrochilidae Fam. Filistatidae Fam. Drymusidae Fam. Scytodidae Fam. Sicariidae Fam. Leptonetidae Fam. Ochyroceratidae Fam. Telemidae Fam. Pholcidae Fam. Caponiidae Fam. Tetrablemmidae Fam. Dysderidae Fam. Oonopidae Fam. Orsolobidae Fam. Segestriidae Fam. Eresidae Fam. Hersiliidae Fam. Oecobiidae Fam. Palpimanidae Fam. Mimetidae Fam. Deinopidae Fam. Uloboridae Fam. Udubidae Fam. Caponiidae Fam. Dysderidae Fam. Oonopidae Fam. Segestriidae Afrotropical (Southern Africa) Present Present Present Present Present Present Present Present Present Present Present Present Present Absent Present Present Present Present Present Absent Present Present Present Present Present Present Present Present Present Present Present Present Present Present Present Present Absent Present Present Present Present Madagascar Present Present Absent Absent Present Present Absent Absent Present Absent Absent Present Present Absent Absent Present Absent Present Absent Absent Absent Absent Present Absent Present Absent Absent Absent Present Present Present Absent Absent Present Present Present Present Absent Absent Absent Present Seychelles Present Present Absent Present Present Present Present Present Absent Present Absent Present Absent Present Absent Present Absent Present Present Present Present Present Present Absent Present Absent Present Absent Present Present Present Present Present Present Present Present Absent Absent Absent Present Present Mascarene Islands Present Present Absent Absent Absent Absent Absent Absent Absent Absent Absent Present Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Present Absent Absent Absent Present Present Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent (continued) 8.16 Seychelles (incl. Aldabra), Comores 725 Table 8.5 (continued) Group Fam. Eresidae Fam. Anapidae Fam. Araneidae Fam. Cyatholipidae Fam. Linyphiidae Fam. Symphytognathidae Fam. Synaphridae Fam. Tetragnathidae Fam. Theridiidae Fam. Theraphosidae Fam. Theridiosomatidae Fam. Ctenidae Fam. Lycosidae Fam. Oxyopidae Fam. Pisauridae Fam. Psechridae Fam. Trechaleidae Fam. Zoropsidae Fam. Agelenidae Fam. Amaurobiidae Fam. Anyphaenidae Fam. Cybaeidae Fam. Desidae Fam. Dictynidae Fam. Hahniidae Fam. Sparassidae Fam. Selenopidae Fam. Zodariidae Fam. Chummidae Fam. Clubionidae Fam. Miturgidae Fam. Phyxelididae Fam. Titanoecidae Fam. Ammoxenidae Fam. Cithaeronidae Fam. Gallielienillidae Fam. Gnaphosidae Fam. Prodidomidae Fam. Trochanteriidae Fam. Philodromidae Fam. Thomisidae Afrotropical (Southern Africa) Present Present Present Present Present Present Absent Present Present Present Present Present Present Present Present Absent Absent Present Present Present Present Absent Present Present Present Present Present Present Present Present Present Present Absent Present Present Present Present Present Present Present Present Madagascar Present Absent Present Present Present Present Present Present Present Absent Absent Present Present Present Present Absent Absent Absent Absent Absent Absent Absent Present Absent Absent Present Present Present Absent Present Present Present Absent Absent Absent Present Present Absent Present Present Present Seychelles Present Present Present Present Present Present Present Present Present Present Present Present Present Present Present Present Present Present Present Present Present Present Present Present Present Present Present Present Absent Present Present Present Present Absent Present Absent Present Present Present Present Present Mascarene Islands Absent Absent Present Present Present Absent Absent Absent Present Present Absent Present Present Present Present Absent Absent Absent Present Absent Absent Absent Absent Absent Absent Absent Present Absent Absent Present Absent Absent Absent Absent Present Absent Present Absent Present (continued) 726 8 Some Peculiarities of the Distribution of Arachnida Table 8.5 (continued) Group Fam. Salticidae Fam. Corinnidae Fam. Liocranidae Order Opilioacarida Fam. Opilioacaridae Order Holothyrida Fam. Holothyridae Order Ixodida Fam. Ixodidae Fam. Argasidae Fam. Nuttalliellidae Order Mesostigmata Order Sarcoptiformes Order Trombidiformes Afrotropical (Southern Africa) Present Present Present Present Present Absent Absent Present Present Present Present Present Present Present Madagascar Present Present Present Present Present Absent Absent Present Present Present Absent Present Present Present Seychelles Present Present Present Absent Absent Present Present Present Present Present Absent Present Present Present Mascarene Islands Present Absent Absent Absent Absent Present Present Present Absent Present Present Present Oribatida On the island of Madagascar and five neighbouring islands: Aldabra, the Comoros, Mauritius, Réunion and the Seychelles, the number of hitherto found Ptyctima species is 57, including 37 Phthiracaroidea, 18 Euphthiracaroidea, and two Mesoplophoroidea. Of those 57 species, 30 are endemic (53%), ten Ethiopian species (17%), 12 pantropical species (21%) three oriental (5%) and two semicosmopolitan (3%). Endemism of Phthiracaroidea (62%) is similar to Euphthiracaroidea (59%). Similarity between the ptyctimous fauna of Madagascar and the neighbouring islands is very small - only one or two species are common. Moreover, the fauna of Mauritius is completely different from that of Madagascar. The ptyctimous fauna of Madagascar and the near islands must have been shaped before separation of Madagascar from Africa (Niedbala 2004). 8.16.2 Remarkable Distributions and Absences Southern Africa – Missing orders or suborders: Orders Holothyrida, Ricinulei, Thelyphonida (Uropygi); suborders Dyspnoi (Opiliones), Mesothelae (Araneae), Palaeoamblypygi (Amblypygi); present orders Opilioacarida (1 genus in common with Madagascar), Schizomida (endemic genus and subfamily), Cyphophthalmi (3 endemic genera of a family shared with Madagascar and other countries), Eupnoi (3 endemic genera, 1 endemic subfamily), Laniatores (1 endemic subfamily), Pseudoscorpiones (14 endemic genera, no endemic families), Scorpiones, Solifugae (endemic families), Araneae (48 endemic genera, 2 endemic families), Acari Ixodida (1 endemic family) 8.17 Evaluation of Wallace’s Line and the Other Lines in Southeast Asia… 727 Madagascar – Missing orders or suborders: Holothyrida, Solifugae, Ricinulei, Uropygi; suborders Eupnoi and Dyspnoi (Opiliones), Mesothelae (Araneae), Palaeoamblypygi (Amblypygi); recent orders Palpigradi (1 endemic genus), Opilioacarida (1 genus in common with Southern Africa), Scorpiones (12 endemic native genera, 100% native species, 1 endemic family, sharp difference with Southern Africa or the other islands), Pseudoscorpiones (no endemic families), Araneae (no endemic families) Seychelles – Missing orders or suborders: Orders Opilioacarida, Ricinulei, Uropygi; suborders Cyphophthalmi, Eupnoi and Dyspnoi (Opiliones), Mesothelae (Araneae), Palaeoamblypygi (Amblypygi); present orders Holothyrida (three endemic genera), Schizomida (four endemic genera) Mascarene – Missing orders or suborders: Orders Opilioacarida, Ricinulei, Uropygi; suborders Cyphophthalmi and Dyspnoi (Opiliones), Mesothelae (Araneae), Palaeoamblypygi (Amblypygi); present orders Holothyrida (endemic family), Pseudoscorpiones (two endemic genera), Eupnoi (Opiliones) On all smaller islands is missing the Opilion family Triaenonychidae, having in Madagascar 15 endemic genera. There are almost no genera of Opiliones in common between Seychelles and Mascarene and no genera in common between Madagascar and the other island groups. Groups of no special zoogeographic interest (no endemisms above species): Southern Africa – Palpigradi Madagascar – Amblypygi, Opilioacarida, Schizomida, Pseudoscorpiones Seychelles – Amblypygi Mascarene - Amblypygi, Eupnoi (Opiliones), Scorpiones Conclusion: Arachnologically Madagascar, the Seychelles, and the Mascarene are so different from the African continent that they deserve the rank of region pairing with Africa. There is also considerable difference between Madagascar and the smaller islands, which are often called subregions. Important is the absence of many groups (orders and families) abundant in Africa but lacking in the islands. 8.17 Evaluation of Wallace’s Line and the Other Lines in Southeast Asia According to the Arachnida. “La frontière, en biogéographie, n’est pas, en général, une ligne, mais une zone complexe don’t l’emplacement et la topographie ont subi de multiples variations au cours des temps géologiques... On peut cependant affirmer que la ligne Wallace – pour certaines formes de Scorpions est une frontière réellement existante.” Max Vachon (1953) 728 8.17.1 8 Some Peculiarities of the Distribution of Arachnida Wallace’s Line In result of his studies of the nature of Australasia, Wallace (1876) concluded that “We may consider it established that the Strait of Lombok [between Bali and Lombok] (only 15 miles wide) marks the limit and abruptly separates two of the great zoological regions of the globe.” This famous line, called by Huxley (1868) “Wallace’s line”, runs through Makassar Strait between Borneo and Sulawesi and then the line was based mostly on mammals, but Mayr (1944b) made the remark that “An equally pronounced faunal difference exists among birds, insects, and other groups of animals in the two regions.” When crossing the famous line in 1994 and 1995, I asked myself: “Is this statement true also for such zoogeographically interesting animals as Arachnida?” Meanwhile, a lot of new information was accumulated on the taxonomy of such groups as Schizomida, Opiliones, Scorpiones, and others, and time has come to check how this zoogeographical “rift” (the term belongs to Prof. Vachon) reflects the past and present distribution of the various Arachnida. Some orders (Solifugae, Opilioacarida, Ricinulei) are not known in this area. 8.17.2 Lydekker’s Line The line proposed in 1895 by the British naturalist Richard Lydekker (1849–1915) is separating Wallacea from the shelf of Australia – New Guinea. 8.17.3 Weber’s Line Max Weber (1852–1937) proposed a line passing between Sulawesi and Halmahera and marks the balance of the Indomalayan and Australian elements in the fauna. It is a line based on mammals and should be tested what concerns the invertebrates (Arachnida and others). Wallacea Observing longtime the strange transition in the animal world of the islands of Malay Archipelago, Wallace presumed, “such facts could be explained only by major changes in the Earth’s surface.” Now we know that in the Pleistocene, the major continental islands Borneo, Java, Sumatra, and Bali have been connected 8.17 Evaluation of Wallace’s Line and the Other Lines in Southeast Asia… 729 with the Asian mainland. The rainforest existed by this time, and the sea level was lower by 180 m. That is why on these islands, we find the same animals like in Malaya (sometimes other subspecies) – elephants, rhinos, tapirs, tigers, leopards, and primates. Some of them meanwhile have disappeared on the continent but still live on the islands (the orangutans). Crossing the narrow (only 25 km) Lombok Chanel between Bali and Lombok, all these animals disappear. Appear cockatoo parrots, some marsupials, and different reptiles. During his research between 1854 and 1862, Wallace found that the birds on Bali and Java are almost the same (97%), and as soon as we cross the 25 km straight, the picture changes abruptly – Bali and Lombok have only 50% of their bird species in common; one may think that the others were not capable to fly over this distance. Starts Wallacea, most interesting territory of a mix of fauna. The ancestors of the present-day animals and plants of Wallacea as a whole originate either from Asia or from Australia – New Guinea – but on the bigger islands occurred an active autochthonous speciation, and there are many endemics. Some of the emblematic Indonesian endemics are actually inhabiting Wallacea (anoa, babyrussa, “Comodo dragon”). Most islands have not been interconnected, they are separated by deep straights, and this was important to explain the high endemism. According to Conservation International, Wallacea has more than 10 000 species of plants, including ca. 1500 (15%) endemic. Among the terrestrial vertebrates, the endemism is even higher: from 1142 species, almost half (529) are endemics. In Wallacea live 223 species of native mammals, 126 of them endemics. Only bats count 124 species – almost one tenth of the world’s bats. The birds of Wallacea are extremely diverse (650 species, including 265 еndemic). More than half of them live in Sulawesi alone (356 species, 96 endemic). In Wallace’s time, the area has been covered by lush tropical forests. Only small fraction of them remains – 45% of the surface is covered by some sort of forest, but only in 52,017 km², or 15% – the forest is in its pristine state. From the total surface of Wallacea (347,000 km²), some 20,000 km² is protected (at least legally). The remaining forest is most often under concessions of powerful Western or Japanese companies, which destroy not only the irreplaceable rainforest but also hundreds of its described, semidescribed, and undescribed inhabitants. 730 8 Some Peculiarities of the Distribution of Arachnida Map 8.6 Location map of Torres Strait in relation to the regional Quaternary shoreline changes (Adapted from Rowe 2007) 8.17.4 Development of Wallacea and the Surrounding Lands of Southeast Asia and the Malayan Archipelago The analysis of Moss and Wilson (1998) concerning the biogeographic implications of the Tertiary paleogeographic evolution of Sulawesi and Borneo is worth quoting. Wallacea, according to them, is a biogeographic region, situated between the areas with Asiatic and Australian floras and faunas, where organisms show a high degree of endemism. A land connection between Borneo and mainland Southeast Asia may have existed throughout much of the Tertiary and would have allowed migration of terrestrial biota. Western Sulawesi had been connected to Eastern Borneo by the Late Cretaceous and by the Early Eocene with possibilities of dispersal of fauna between Borneo and Western Sulawesi. The East Sulawesi ophiolite was accreted onto Sulawesi during or after the late Oligocene and resulted in the formation of more extensive land areas of Sulawesi. “Microcontinental fragments accreted onto eastern Sulawesi in the Miocene to Pleistocene may have been emergent as they drifted towards Sulawesi and allowed island hopping or rafting for biota of Australian affinity. Island hopping routes for the dispersal of organisms between Borneo-Sulawesi and the Philippines may have existed along volcanic arcs, such as the long-lived North Sulawesi arc, the Sulu and Sangihe arcs, and the Cagayan arc” (Moss and Wilson 1998). 8.17 Evaluation of Wallace’s Line and the Other Lines in Southeast Asia… 731 Remarks Concerning Araneae From the three suborders of spider, one (Mesothelae) does not live east of Wallace’s Line but is known from Southeast Asia. From 112 families of spiders in the world, 19 are known from Australasia east of this line, but not from Southeast Asia. Some are endemic for Australia, New Zealand, or New Caledonia; others live also in faraway parts of the world, mostly in South Africa, Chile, or Argentina. Typical are Austrochilidae, Orsolobidae, Malkaridae, Cyatholipidae, Amphinectidae, Zoropsidae, and Ammoxenidae, also subfam. Prithinae of fam. Filistatidae (one genus in New Guinea, two in Australia, seven in Africa, North and South America). Some subfamilies of widespread families are endemic for the area east of Wallace’s Line: Hexathelidae (Hexathelinae), Dipluridae (Masteriinae), Cyrtaucheniidae (Aporoptychinae), Idiopidae (Arbanitinae), Miturgidae (Diaprograptinae), Gallieniellidae (Meedoinae), and others. 8.17.5 The “Lines” and Some Other Groups In their Atlas of Diplopoda, Shelley and Golovatch (2011) conclude that From present records, the hypothetical borders separating Asian and Australian faunas in Indonesia (Wallace’s, Weber’s, and Lydekker’s ‘lines’) hold little relevance for diplopods. Of the four exclusively Laurasian orders, only Glomerida and Platydesmida (…) expand into the East Indies/Indonesia, as Julida and Callipodida (…) terminate on the southeast Asian continent. In the north, Glomerida spread eastward to Weber’s line, between Sulawesi and Halmahera, while the southern edge conforms roughly to Wallace’s line. Platydesmida, however, extend to Wallace’s line between Borneo and Sulawesi while the southern border passes between Sumatra/Java and corresponds to nothing. We would not expect these ‘lines’ to be operative for the Gondwanan representatives, which were carried passively to southeast Asia by the terranes, and this is indeed the case. The only taxon conforming precisely to one of these hypothetical boundaries is Spirostreptidea, derived from Gondwana I, whose eastern border matches Lydekker’s line completely. While future discoveries may alter East Indian taxon borders, hypotheses like these lines may not apply to low vagility organisms like millipeds, whose geographies are so dependent on geological events; conformity of Spirostreptidea may therefore be coincidence. In a recent analysis of the caddisflies (Trichoptera) of Lombok, Bali, and Java, Malicky et al. (2014) conclude that “The caddisfly fauna of the three islands is of Asiatic origin, no Australian influence was noted. The well-known Wallace’s line does not act as a faunistic border between Bali and Lombok for Trichoptera.” It is interesting to compare this analysis with our data concerning the Arachnida – other non-insect group of relatively small animals with (mostly) low vagility. Of course, there are differences – almost all Arachnida are predators, not living so much in rotten logs, some are ballooning, and others are small and relatively easy to be dispersed by air or otherwise (at least, it seems so). According to Lehtinen (1980), “Various lines limiting and dividing the Wallacea seem to have no equivalents in the arachnological zoogeography.” It seems that Gressitt (1974) arrived to a similar conclusion on what concerns the zoogeography of insects. 732 8 Some Peculiarities of the Distribution of Arachnida Map 8.7 Australasia and the lines (After Hall 2002, modified) 8.17.6 Arachnida and the Lines Palpigradi Two families (Eukoeneniidae and Prokoeneniidae) are represented in the area. West of Wallace’s Line: Fam. Prokoeneniidae and the genus Prokoenenia (one sp. on Sulawesi) – Thailand, Java, Sulawesi East of Wallace’s Line: None Both sides: Fam. Eukoeneniidae (genera Eukoenenia and Koeneniodes) Amblypygi Two families are represented in the area, Charinidae and Charontidae, both of the two sides of the Wallace’s Line. Special case is the only species of Phrynus (from the American family Phrynidae) from a cave on Flores (Indonesia). Only West of Wallace’s Line: Catageus Thorell, 1889 – Burma Only East of Wallace’s Line – None 8.17 Evaluation of Wallace’s Line and the Other Lines in Southeast Asia… 733 Both sides: Charinus Simon, 1892 (Charinides Gravely) – Australia, Andaman Islands, India, W. Samoa, Vanuatu, Indonesia (Java, Borneo), Singapore, Malaysia, Papua New Guinea, New Caledonia, Solomon Islands Sarax Simon, 1892 (Phrynichosarax Gravely) – Solomon Islands, Malaysia, Singapore, the Philippines (Luzon), Indonesia (Java, Kalimantan), India, Andaman Islands, Papua New Guinea, New Britain, Vietnam, Laos, Cambodia, Borneo Charon Karsch, 1879 – Australia (Northern Territory, Queensland, Christmas Island), Indonesia (Java, Maluku, Sumbawa), Malaysia (incl. Borneo), Palau, Papua New Guinea, the Philippines, Singapore, Solomon Islands Stygophrynus Kraepelin, 1895 – Solomon Islands (Guadalcanal), Burma, Thailand, Vietnam, Malaysia, Indonesia (Java, Sumatra, Kalimantan) Thelyphonida (Uropygi) Uropygi (Thelyphonidae, incl. Hypoctoninae) are known from the area (from both sides of Wallace’s Line). West of Wallace’s Line: Ginosigma Speijer, 1936 – Sunda Islands, Thailand Hypoctonus Thorell, 1889 – Burma (Myanmar), South China, Malaysia, Thailand, Bangladesh, Java, India East of Wallace’s Line: Chajnus Speijer, 1936 – Sunda Islands (Lombok) Glyptogluteus Rowland, 1973 – Philippines (Panay) Thelyphonoides Krehenwinkel et al., 2009 – Philippines (Panay) Both sides: Thelyphonus Latreille, 1802 (syn. Abaliella Strand, 1928, syn. Minbosius Speijer, 1936, syn. Tetrabalius Thorell, 1889, fide Haupt, 2009a) – Indonesia, Singapore, Philippines, Vanuatu (New Hebrides), Fiji, Rennell, Solomon Islands, New Guinea, New Britain, Burma, Sri Lanka, India, Thailand, W. Samoa, Borneo, Moluccas Typopeltis Pocock, 1894 (syn. Teltus Speijer, 1936) – China, ? Russia, Taiwan, Hainan, Japan, Thailand, Vietnam Schizomida One family (Hubbardiidae) is represented in the area. West of Wallace’s Line: Schizomus Packard-Cambridge, 1972 Clavizomus Reddell et Cokendolpher, 1995 – Java, West Malaysia, Singapore Trithyreus Kraepelin, 1899 – Burma (Myanmar) Zomus Reddell et Cokendolpher, 1995 – Malaysia (incl. Sarawak), Singapore, England (Kew Gardens) 734 8 Some Peculiarities of the Distribution of Arachnida East of Wallace’s Line: Attenuizomus Harvey, 2000 – Australia, NT Brignolizomus Harvey, 2000 – Australia Draculoides Harvey, 1992 – Western Australia Julattenius Harvey, 1992 – Queensland (Australia) Notozomus Harvey, 1992 - Queensland (Australia) Orientzomus Cokendopher et Tsurusaki, 1994 – Philippines (Luzon), Japan, Bonin Island Paradraculoides Harvey et al., 2008 – Western Australia Hubbardiidae indet. – New Guinea, New Britain, New Ireland; Schizomus modestus Both sides: Apozomus Harvey, 1992 – Australia, Japan, Papua New Guinea, Taiwan, Sabah (Borneo), W. Malaysia, Vietnam Bamazomus Harvey, 1992 – Thailand, Hong Kong, W. Malaysia, Ryukyu Islands, Papua New Guinea, Australia, Hawaii Ovozomus Harvey, 2001 – Seychelles, Comoro Islands, Cook Islands, Ceylon, India, Christmas Island, Réunion Scorpiones Seven families of scorpions are known from the area. West of Wallace’s Line: Fam. Chaerilidae – Bangladesh, India, Indonesia, Malaysia, Nepal, Singapore, Sri Lanka, Vietnam Fam. Pseudochactidae – Caves of Laos and Vietnam Fam. Scorpiopsidae – Southeast Asia, India, Indonesia, Malaysia Fam. Scorpionidae – South and Southeast Asia, Indonesia East of Wallace’s Line: Fam. Urodacidae – Australia Both sides: Fam. Buthidae – All continents, tropical and subtropical countries and in some countries with moderate climate Fam. Hormuridae – Australia, Southeast Asia, Pacific islands, Indian Ocean Islands Pseudoscorpiones In the area have been registered representatives of 20 families. Most of them (18) are represented from both sides of Wallace’s Line and usually also on other continents or worldwide. West of Wallace’s Line: Fam. Ideoroncidae – Malaysia, India, Cambodia, Thailand, Vietnam, Indonesia (Sumatra) 8.17 Evaluation of Wallace’s Line and the Other Lines in Southeast Asia… 735 East of Wallace’s Line: Fam. Pseudogarypidae – Tasmania (one extant species) Both sides: Fam. Chthoniidae – Cosmopolitic Fam. Tridenchthoniidae (Dithidae) – Australia, New Caledonia, Lord Howe Island, Caroline Islands, India, Vietnam, Philippines, Indonesia (Moluccas, Sumatra, Java), New Guinea, Japan, Palau Island, Bhutan, Nepal, Laos, Marcus Island, Malaysia Fam. Lechytiidae – Vietnam, Marshall Islands, Marcus Island Fam. Hyidae – Philippine Islands, Indonesia (Java, Sumatra), Australia Fam. Neobisiidae – Philippines, Vietnam, Thailand, Japan, Burma, China, Taiwan Fam. Syarinidae – Bhutan, India, Sri Lanka, Nepal, Tuvalu, Philippines, Solomon Island, New Zealand, New Caledonia, New Guinea Fam. Parahyidae – Caroline Islands, Singapore Fam. Garypidae (Synsphyronidae) – India, Japan, Indonesia (Krakatau, Nicobar Island), Maldives Island, Sri Lanka-Marshall Islands, Australia (incl. Tasmania), New Zealand Fam. Menthidae – Socotra, Australia Fam. Cheiridiidae – China, Taiwan, Nepal, Philippines, Japan, Hawaii-Papua New Guinea, Salomon Islands, Mariana Islands, Australia, New Zealand Fam. Olpiidae – Australia, New Zealand, Norfolk Island, Indonesia, India, Burma, Japan, Cambodia, Vietnam, Papua New Guinea, New Caledonia, Salomon Islands, Tuvalu, Samoa, Caroline Islands, Mariana Islands, Marshall Islands Fam. Garypinidae – Solomon Islands, Australia, New Guinea, Hawaii, Indonesia, Thailand, Cambodia, Vietnam Fam. Sternophoridae – Australia, Papua New Guinea, India, Sri Lanka, Laos, Cambodia, Vietnam Fam. Withiidae – New Zealand, Lord Howe Island, Vietnam, Indonesia, Christmas Island, India, Cambodia, Malaysia, Thailand, Philippines, Solomon Islands, Nepal, China, Taiwan, Mariana Islands, Marshall Islands, Papua New Guinea Fam. Cheliferidae – Indochina, India, China, Indonesia-Australia, New Zealand, Papua New Guinea Fam. Atemnidae (Miratemnidae) – South East Asia, Philippines, Bhutan, Nepal, India, Nicobar Islands, Solomon Islands, Sri Lanka, Indonesia, China-New Caledonia, Lord Howe Island, Caroline Islands, Mariana Islands, Christmas Island, Marquesas Islands, Marshall Islands, Papua New Guinea Fam. Chernetidae (Myrmochernetidae) – New Guinea, Solomon Islands, etc. Opiliones All suborders are represented in the area. Cyphophthalmi (three fam.) Only west of Wallace’s Line: None Only east of Wallace’s Line: 736 8 Some Peculiarities of the Distribution of Arachnida Fam. Troglosironidae – New Caledonia Both sides: Fam. Stylocellidae – Southeast Asia (from India to New Guinea) Fam. Pettalidae – New Zealand, Australia-Sri Lanka Eupnoi West of Wallace’s Line: Fam. Phalangiidae – Mitopiella Banks, Borneo East of Wallace’s Line: Fam. Caddidae – Australia (incl. Tasmania), New Zealand (but also in North America, South America, and South Africa, not found in Southeast Asia) Fam. Sclerosomatidae – Europe, Asia, Africa, South, Central and North America, Antilleans, Australia, New Guinea, Solomon Island (not found in Southeast Asia) Fam. Neopilionidae – Australia, New Zealand (but also in South America and South Africa, not found in Southeast Asia) Both sides: None Dyspnoi West of Wallace’s Line: Fam. Nemastomatidae (Ortholasmatinae, Cladolasma Suzuki) – Thailand, China, Japan (three spp.) East of Wallace’s Line: None Both sides: None Laniatores West of Wallace’s Line: Fam. Stygnommatidae – ? Neotropical family with two doubtful spp. from Malaya of genus Stygnomimus Roewer, 1927 Fam. Biantidae (Biantes, Probiantes) – Nepal, India, Burma, Sumatra, Malaya Fam. Pyramidopidae – Africa, Central America East of Wallace’s Line: Fam. Synthetonychiidae – New Zealand (14 spp.) Fam. Triaenonychidae – Australia, Tasmania, New Zealand, New Caledonia (but also in Madagascar, South Africa, South America) Both sides: Fam. Assamiidae – Southern Asia, subfam. Dampetrinae to Indonesia, New Guinea, and Australia Fam. Samoidae – Australia (New South Wales), Samoa, Indonesia 8.17 Evaluation of Wallace’s Line and the Other Lines in Southeast Asia… 737 Fam. Podoctidae (incl. Erecananinae and Ibaloniinae) – Palau, Taiwan, Indonesia, Malaysia, New Guinea, India, Philippines, Vanuatu Fam. Sandokanidae [Oncopodidae, praeoccup.] – Philippines, Sumatra, Borneo, Burma, Nepal, India, Malaysia, Singapore, Thailand Fam. Epedanidae (incl. Dibuninae, Acrobuninae, Sarasiniciinae, Sarasinellinae) – Philippines, New Guinea, Thailand, Sulawesi, Borneo, Sumatra, Japan, Malaysia, India, Taiwan, China, Vietnam Fam. Zalmoxidae (Stygnoleptinae) – New Guinea, Solomon Islands, Fiji, Bismarck Archipelago, Philippines, Java, Sulawesi, Moluccas, Australia, Marianas, New Caledonia, Marshall Islands, Caroline Islands, Polynesia Remarks Concerning Araneae From the three suborders of spider, one (Mesothelae) does not live east of Wallace’s Line but is known from Southeast Asia. From 112 families of spiders in the world, 19 are known from Australasia east of this line, but not from Southeast Asia. Some are endemic for Australia, New Zealand, or New Caledonia; others live also in faraway parts of the world, mostly in South Africa, Chile, or Argentina. Typical are family Austrochilidae, Orsolobidae, Malkaridae, Cyatholipidae, Amphinectidae, Zoropsidae, and Ammoxenidae, also subfam. Prithinae of fam. Filistatidae (one genus in New Guinea, two in Australia, seven in Africa, North and South America). Some subfamilies of widespread families are endemic for the area east of Wallace’s Line: Hexathelidae (Hexathelinae), Dipluridae (Masteriinae), Cyrtaucheniidae (Aporoptychinae), Idiopidae (Arbanitinae), Miturgidae (Diaprograptinae), Gallieniellidae (Meedoinae), and others. Araneae Only West of Wallace’s Line (Especially in Southeast Asia) Suborder Mesothelae – Southeast Asia, China, Japan Fam. Liphistiidae – Southeast Asia, China, Japan Suborder Orthognathi Infraorder Mygalomorphae – Both sides Fam. Atypidae Fam. Hexathelidae – New Zealand, Australia, Tasmania, Asia (Macrothele) Subfam. Hexathelinae – Australia, New Zealand Subfam. Macrothelinae – Asia, New Zealand Fam. Dipluridae – Australia, Oceania, Taiwan, Thailand, New Caledonia Subfam. Euagrinae – Australia, Taiwan, New Caledonia Subfam. Masteriinae – Oceania, Australia Fam. Nemesiidae – Burma, India, China, Australia, New Zealand, Ryukyu Islands, Southeast Asia Fam. Theraphosidae – Australia, South and Southeast Asia, New Guinea Fam. Barychelidae – Australia, New Caledonia, Southeast Asia, New Guinea, Sri Lanka, Oceania, Fiji 738 8 Some Peculiarities of the Distribution of Arachnida Subfam. Barychelinae – Australia, New Caledonia Subfam. Sasoninae – Australasia Fam. Atypidae – Asia Fam. Cyrtaucheniidae – Australia, Thailand Subfam. Aporoptychinae – Australia Fam. Idiopidae – Australia, New Zealand, India, Sri Lanka, South Asia, Thailand Subfam. Arbanitinae – Australia, New Zealand Fam. Ctenizidae – Taiwan, China, Thailand, Australian region, New Guinea Subfam. Ctenizinae – Thailand, China, Taiwan, Australian region Subfam. Pachylomerinae – Australian region Fam. Migidae – Australia, New Zealand Subfam. Calathotarsinae – Australia Subfam. Migidae – New Zealand, Australia Fam. Actinopodidae – Australia Suborder Araneomorphae Fam. Austrochilidae – Tasmania Subfamily Hickmaniinae (as family Hickmaniidae) Fam. Gradungulidae – New Zealand, Australia (A.C.T., Queensland, New South Wales, Victoria) Fam. Filistatidae – Australia, New Guinea Subfam. Prithinae – New Guinea Fam. Periegopidae – Australia (Queensland), New Zealand Fam. Scytodidae – Worldwide, except of the Far North Fam. Sicariidae – Worldwide, except of the Far North Fam. Ochyroceratidae – South Asia, China, Pacific Islands, New Guinea Fam. Telemidae – Sumatra, Malaysia, China, Vietnam, New Caledonia, New Guinea Fam. Pholcidae – Worldwide Fam. Tetrablemmidae – Southeast Asia, Borneo, India, Indonesia (Sulawesi), Samoa, Caroline Islands Fam. Dysderidae – Dysdera, worldwide Subfam. Dysderinae – Dysdera, worldwide Fam. Oonopidae – Philippines, Southeast Asia, Australia, Tasmania, New Zealand, New Caledonia, Hawaii, Sri Lanka Subfam. Gamasomorphinae – Bhutan, Nepal, China, Philippines, Southeast Asia, Australia, New Zealand, Hawaii Subfam. Oonopidae – New Caledonia, Tasmania, Sri Lanka 8.17 Evaluation of Wallace’s Line and the Other Lines in Southeast Asia… 739 Fam. Orsolobidae – Australia, New Zealand, Tasmania Fam. Segestriidae – Asia, Australia, New Zealand Fam. Lamponidae – Australia, New Zealand, New Caledonia (genus Centrocalia) Subfam. Centrothelinae – Australia, New Caledonia Subfam. Lamponinae – Australia, New Zealand Subfam. Pseudolamponinae – Australia Fam. Prodidomidae – Australia Fam. Eresidae – Eurasia Fam. Hersiliidae – Australasia, India, Sri Lanka, Australia, Borneo Fam. Oecobiidae – Asia; Oecobius is cosmopolitan Fam. Holarchaeidae – Tasmania, New Zealand Fam. Mecysmaucheniidae – New Zealand Subfam. Mecysmaucheniinae Simon, 1895 – New Zealand Subfam. Zearchaeinae Forster et Platnick, 1984 – New Zealand Fam. Micropholcommatidae – New Zealand, Australia, Tasmania, New Guinea Fam. Pararchaeidae – Australia (incl. Tasmania), New Zealand, New Caledonia Fam. Huttoniidae – New Zealand Fam. Palpimanidae – South Asia, Sri Lanka Subfam. Chediminae – South Asia, Sri Lanka Fam. Stenochilidae – Thailand, Malaysia, Singapore, Burma, Philippines, Bali, New Guinea, Fiji, Borneo, Vietnam, India Fam. Malkaridae – Australia Fam. Mimetidae – Palearctic, Africa, Principe, Central and South America, USA, Asia, Australia: Ero and Mimetus worldwide Fam. Deinopidae – All continents, except Europe and Antarctica Fam. Uloboridae – Philippines, Samoa, Fiji, Vanuatu, New Guinea, Oceania, Australia, New Zealand; Uloborus is spread worldwide; Zosis is pantropical Fam. Anapidae – New Caledonia, Australia, Tasmania, New Zealand, New Guinea, China, Taiwan Fam. Araneidae – Worldwide Fam. Cyatholipidae – Australia Fam. Linyphiidae – Worldwide Fam. Mysmenidae – Samoa, New Caledonia, Tasmania Fam. Nesticidae – Worldwide Fam. Pimoidae – North America, Asia, Europe, Japan The Pimoidae form a relictual group along the Western coast of North America, Europe (Alps, Apennines, and Cantabrian Mountains of northern Spain), and the Himalaya. This makes a Holarctic predecessor probable. In 2003, a species was found in Japan. 740 8 Some Peculiarities of the Distribution of Arachnida Fam. Sinopimoidae – (one sp.) Fam. Symphytognathidae – In the tropics of Central and South America and the Australian region (with Oceania), three species (Anapistula benoiti, A. caecula, Symphytognatha imbulunga) found in Africa and one (Anapistula ishikawai) in Japan. Anapistula jerai occurs in Southeast Asia. Fam. Synotaxidae – Australia, Tasmania, New Zealand Subfam. Pahorinae – Endemic to New Zealand Subfam. Physogleninae – Australia, Tasmania, New Zealand Subfam. Synotaxinae – Australia Fam. Nephilidae – Pantropical Fam. Theridiidae – Cosmopolitic Fam. Theridiosomatidae – Australia, China, Malaysia, Sri Lanka, Philippines, Sumatra, Samoa, Taiwan Fam. Ctenidae – Australia, China, Sumatra, New Guinea Fam. Lycosidae – Cosmopolitic Fam. Oxyopidae – South East Asia Fam. Pisauridae – Africa, South America, North America, Asia, New Caledonia, Canary Islands, Madagascar, New Guinea, Cuba, Chatham Islands, Australia, etc. Fam. Psechridae – China, India, Nepal, Sri Lanka, Southeast Asia, Nicobar Islands, Philippines, Australia Fam. Stiphidiidae – Australia, Tasmania, New Zealand, Madagascar, Mauritius Fam. Trechaleidae – South and Central America, Mexico, USA Fam. Udubidae – Sri Lanka, Madagascar, Africa, USA to Panama Fam. Zoropsidae – Australia, New Zealand, Sri Lanka, China, Korea, Japan, Cyprus, Mediterranean, South Africa Fam. Agelenidae – Mediterranean, Europe to Central Asia, USA to Chile, New Zealand, Socotra, Himalaya, Cuba, Bahamas, Cyprus, China; Tegenaria, worldwide Fam. Amphinectidae – Australia, Tasmania, New Zealand, Chile, South America Fam. Amaurobiidae – Russia, Korea, China, Japan, Argentina, Paraguay, Chile, Taiwan, Australia, New Zealand, USA, Canada, Falkland Islands, India, Laos, Thailand, Vietnam, Balkans, Greece, Crete, Germany, Switzerland, Italy, Micronesia, Ethiopia, Libya, Nepal, Bhutan, Pakistan, Tajikistan, etc. Fam. Anyphaenidae – Cosmopolitic Fam. Cybaeidae – America, Europe, Japan, Korea, China, Turkmenistan, Tajikistan, Uzbekistan, Sumatra, Venezuela, Colombia; Argyroneta, Palearctic Fam. Desidae – Australia, Tasmania, New Zealand, New Caledonia, Oceania, Korea, Japan, the USA, Paraguay, Chile Fam. Dictynidae – Europe, China, Mongolia, India, Algeria, Kazakhstan, Canary Islands, New Zealand, Australia, Sri Lanka, South Africa, USA, Canada, Mexico, Kyrgyzstan, West Indies, Hawaii, South America, St. Helena, Andaman Islands, Galapagos Island, New Caledonia, etc. 8.17 Evaluation of Wallace’s Line and the Other Lines in Southeast Asia… 741 Fam. Hahniidae – Oceania, Australia, Tasmania, New Zealand, India, Philippines, Sumatra Fam. Nicodamidae – Australia, Tasmania, New Zealand, New Guinea Fam. Sparassidae – Worldwide, except of the Far North Fam. Selenopidae – Asia Fam. Zodariidae – Australia, New Zealand, New Guinea, Southeast Asia Fam. Zoropsidae – New Zealand Fam. Clubionidae – Lord Howe Island, Asia, Australia, Oceania, Sri Lanka Fam. Cycloctenidae – New Zealand, Australia, Indonesia (Java) Fam. Miturgidae – Thailand, Indonesia, Australia, Tasmania, New Zealand, New Guinea, Central and South America, Israel; Zora, Palearctic Subfam. Eutichurinae – Australia, Thailand, Indonesia Subfam. Miturginae – Australia, New Guinea Subfam. Systariinae – Southeast Asia Subfam. Diaprograptinae – Timor, Australia, New Zealand Subfam. Zorinae – Central and South America, Australia, Tasmania, New Zealand, Israel; Zora, Palearctic Fam. Phyxelididae – Sumatra, Borneo Fam. Titanoecidae – India, Sri Lanka to China, New Guinea, Marquesas Islands, Europe, Mexico, South America; Titanoeca, Holarctic Fam. Ammoxenidae – Australia, Tasmania, Southern Africa Fam. Cithaeronidae – Africa, India, Malaysia, Greece, introduced to Australia and Brazil Fam. Gallieniellidae – Australia, Argentina, Madagascar, South Africa, Kenya Subfam. Meedoinae – Australia (five gen.) Fam. Gnaphosidae – Worldwide Fam. Trochanteriidae – Australia, Sulawesi, Christmas Island, China, India, New Caledonia Fam. Philodromidae – India, New Guinea, Southeast Asia, Hawaii Fam. Thomisidae – Worldwide Fam. Salticidae – Worldwide Fam. Corinnidae – Worldwide Fam. Liocranidae – Burma, Thailand, Borneo, Sumatra, Australia, Sri Lanka, New Guinea Opilioacarida (both sides of the “Line”) Ref.: Beron (1914), Das and Bastawade (2007), Leclerc (1989), Walter and Proctor (1998) Only one family (Opiliocaridae) in the world. Only west side of Wallace’s Line: Endemic genus Indiacarus Das et Bastawade, 2007 – India Endemic genera Siamacarus Leclerc, 1989 and Vanderhammenacarus Leclerc, 1989 – Thailand (caves) 742 8 Some Peculiarities of the Distribution of Arachnida East side of Wallace’s Line: Indet. Opilioacarida from Australia (Walter and Proctor 1998) Holothyrida (both sides of the Line) Ref.: Berlese (1923), Beron (1914), Domrow (1955), Lehtinen (1981, 1991, 1995), Thorell (1882), Van der Hammen (1961, 1983), Womersley (1935) Only west side of Wallace’s Line: None in Southeast Asia East side of Wallace’s Line: Fam. Allothyridae – Australia, New Zealand (two gen., three spp.) Both sides: Fam. Holothyridae – New Guinea, Seychelles, New Caledonia, Mauritius, Sri Lanka, Lord Howe Island (23 spp.) Remarks: It seems interesting that in New Caledonia and Lord Howe Island live holothyrids of the genera Lindothyrus Lehtinen, 1995, and Haplothyrus Lehtinen, 1995, of fam. Holothyridae and not of Allothyridae. Some Arachnida and the Lydekker’s Line Palpigradi – Both sides of Lydekker’s Line (LL) Amblypygi – Charinidae and Charontidae live on both sides (usually the same genera), the only species of Phrynidae from Flores is west of LL. Thelyphonida (Uropygi) – Only one widespread genus (Thelyphonus Latreille s. lato) is found east of LL (in New Guinea), species of seven genera live in Southeast Asia, Indonesia, or the Philippines. Schizomida – The only family (Hubbardiidae) of LL, with some endemic genera on either side. Pseudoscorpiones – As with the Wallace’s Line, only one family (Ideoroncidae) is known from the west side, one family (Pseudogarypidae, relict in Tasmania LL.) only is from the east side, and 18 families of Pseudoscorpiones live on both sides of Opiliones Cyphophthalmi – Only the relict family Troglosironidae is known from the east side of LL (New Caledonia), no family is known only from the Western side, and two families (Stylocellidae and Pettalidae) have been recorded from both sides (as with Wallace’s Line). Opiliones Dyspnoi – Practically missing in the area (one sp. in Thailand) Opiliones Eupnoi – Fam. Neopilionidae is found in Australia, New Zealand (but, similarly to Triaenonychidae, also in South Africa and South America) Opiliones Laniatores – Synthetonychiidae is endemic in New Zealand; Triaenonychidae is known east of LL but has representatives also in South Africa, South America, and Madagascar. Six families are distributed on both sides of LL. Scorpiones – As with the Wallace’s Line, only one family (Urodacidae) is typical for the Australian side, four families are known only in the west, and two (Buthidae and Hormuridae) live on both sides. Araneae – Suborder Mesothelae lives only West of all “lines.” Mygalomorphae and 8.17 Evaluation of Wallace’s Line and the Other Lines in Southeast Asia… 743 Araneomorphae are crossing all “lines”; many families are restricted to the land east of Lydekker’s Line (see the comments above). Acari – All orders are found on both sides of Lydekker’s Line. Allothyridae (Holothyrida) is confined to Australia and New Zealand. According to Lehtinen (1980), “Various lines limiting and dividing the Wallacea seem to have no equivalents in the arachnological zoogeography.” It seems that Gressitt (1974) arrived to a similar conclusion on what concerns the zoogeography of insects. 8.17.7 Conclusions From the orders of Arachnida, some (Ricinulei and almost all Solifugae) are missing from the analyzed area (Indomalayan and Australian regions). The situation with the other orders is as follows. Palpigradi – Fam. Eukoeneniidae is found on both sides (the same genera), fam. Prokoeneniidae is found in Thailand, Java, and Sulawesi, almost not living east of Wallace’s Line. Amblypygi – Endemism on species level, all families and almost all genera are found on both sides. Thelyphonida (Uropygi) – Low level of endemism (only genera), one family on both sides. Schizomida – Low level of endemism (only genera), one family on both sides. Scorpiones – Four families live only west of the “Line,” only one (Urodacidae) is known only east of it. Two families are known on both sides. Solifugae – Represented only by one species (Dinorhax rostrumpsittaci Simon) in Vietnam and Maluku Islands (both sides of Wallace’s Line). Pseudoscorpiones – One family (Ideoroncidae) only west of Wallace’s Line, 1 family (Pseudogarypidae) only east of the Line (relict in Tasmania), 18 families on both sides. Opiliones – Cyphophthalmi. Two families live on both sides, one (the relict Troglosironidae in New Caledonia) only on the east side, none on the west side. Opiliones – Eupnoi. One widespread family (Phalangiidae) is known on the west side (Borneo); three families live on the east side but are recorded also from faraway countries. Opiliones – Dyspnoi. One family (Nemastomatidae) is represented in Thailand with one species; none is known from the east side or from both sides (practically the suborder is absent in the whole area). 744 8 Some Peculiarities of the Distribution of Arachnida Opiliones – Laniatores. Two families are known only from the west side, and two others are known only from the east side, but one of them (Triaenonychidae) is recorded also from Madagascar, South Africa, and South America (but not found in Southeast Asia). The other (Synthetonychiidae) is endemic in New Zealand. Six families live on both sides. Araneae Mesothelae – Only west of the “Lines” Mygalomorphae – Both sides of the “Lines” Araneomorphae – Worldwide distributed suborder, living on both sides of the “Lines” Only on the west side of Wallace’s Line are known five families. Only on the east side are known 20 families. Some are endemic for Australia, New Zealand, or New Caledonia; others live also in faraway parts of the world, mostly in South Africa, Chile, or Argentina, but not in Southeast Asia. On both sides are recorded at least 70 families (out of the 114 families of spiders in the world), mostly widespread. Opilioacarida – The only family of the order Opilioacaridae is known from both sides of the “Line.” Two endemic genera live in India and Thailand; from Australia the order is recorded, but without further data. Parasitiformes – We shall consider here only the zoogeographically interesting suborder Holothyrida, living on both sides of the “Line.” The family Holothyridae is known with endemic genera from both sides of the “Line” (not recorded in Southeast Asia, but living in India, Seychelles, Mauritius, and Sri Lanka). Only East of the “Line” (Australia and New Zealand) is known the family Allothyridae. None of the higher groups of Acari (Opilioacarida, Sarcoptiformes, Prostigmata, Acaridida, Oribatida, Parasitiformes, Mesostigmata, Ixodida, and Holothyrida) is limited by the “Lines.” In general, the analysis of Arachnida on both sides of Wallace’s Line confirms the conclusion of Shelley and Golovatch (2011) that “… the hypothetical borders separating Asian and Australian faunas in Indonesia (Wallace’s, Weber’s, and Lydekker’s ‘lines’) hold little relevance for diplopods.” The “lines” seem to “hold little relevance” also for Arachnida. There is no order of Arachnida living only on one side of the “Lines.” Only the spider suborder Mesothelae seems limited to the Western side. Some confirmation of the Vachon’s opinion (1953) (“On peut cependant affirmer que la ligne Wallace – pour certaines formes de Scorpions – est une 8.17 Evaluation of Wallace’s Line and the Other Lines in Southeast Asia… 745 frontière réellement existante”) could be confirmed what concerns the scorpions. There are families and genera, known only on one side, but the level of endemism is relatively low. Within Arachnida we cannot detect anything similar to the endemic orders and even subclasses in the vertebrates which are the base of the construction of the “Lines.” Conclusion of Lehtinen (1980): “Arachnological data suggests a rearrangement of the classical Oriental and Australian regions into a big, fairly homogenous IndoPacific region and a South-Gondwanan region, the latter extending also to southern parts of other continents of the southern hemisphere. The limitation of their subregions is largely different from the classical limitations based on the distribution of vertebrates. Various lines limiting and dividing the Wallacea seem to have no equivalents in the arachnological zoogeography.” This very experienced author thinks also that “The spider fauna of Melanesia, Micronesia, and Polynesia, with the exception of New Caledonia, Lord Howe Island and surrounding small archipelagoes, is simply an impoverished Oriental fauna.” Principal geological events important to Wallace’s Line (Partly after Audley-Charles 1981) Million Q 10 20 30 40 50 60 70 Period Pleistocene Pliocene Miocene Oligocene Eocene Palaeocene Q – Quaternary 1. Australia/New Guinea splits from Antarctica (c. 53 Ma) 2. Formation of the Philippines by collision of Asian continental fragment with an island arc (Oligocene). 3. Possible land connection(s) across Makassar Strait (mid-Miocene) 4. Collision between New Guinea and a Tertiary island arc (c.15 Ma) 5. Collision between Gondwana (Sula Peninsula) and Laurasia at or near east Sulawesi (c. 15 Ma) but submarine 6. Island chain established between east Sulawesi and Australia (Late Miocene to Late Pliocene) 7. Collision between parts of Gondwanan Outer Banda Arc and Laurasian (volcanic) Inner Banda Arc (Latest Miocene to Early Pliocene) 8. Probable land connection (s) across south Makassar Strait (from Late Pliocene) 746 8.18 8 Some Peculiarities of the Distribution of Arachnida The Arachnida of the Papuan Subregion 8.18.1 New Guinea and Melanesia 8.18.1.1 Geography, General Zoogeography, and Paleogeography Melanesia consists of the island of New Guinea, the Bismarck Archipelago, d’Entrecasteaux Islands, Louisiade Archipelago, Maluku Islands (not included here), Fiji, Norfolk Island, Solomon Islands, Schouten Islands, Torres Strait Islands, Trobriand Islands, Vanuatu, Woodlark Island, and some other islands. Central is the huge island of New Guinea (829 000 km2, maximal height 4884 m.). It is actually a small continent – only the surface of the state of Papua New Guinea is 462 840 km2. Within this state are also the “smaller” islands – New Britain (35.600 km2, more than Belgium), New Ireland (8.600 km2, comparable to Corsica), Bougainville (10.500 km2), and others. About 25% of New Guinea is situated higher than 1000 m. The giant mountain chain, stretching along the entire island, is long more than 2000 m. and is higher than the Alps (4884 m). The other islands are also mountainous (New Britain 2440 m, Bougainville 2740 m). The rivers of the Big Island (fed by up to 6500 mm rain annually) are impressive. Sepik, Fly, Ramu, and Baliem are comparable to Rhine or Don. In the Papuan phytogeographic Province (of the Malesian Region) are known more than 9000 species and 1400 genera of higher plants, from which 8500 species and 140 genera are endemic. “When New Guinea, the leading edge of the Australian plate, was first elevated to form an extensive land area in the late Oligocene, it was colonized largely from the adjacent rich tropical lowlands of Malaysia. Biogeographic affinities with Australia are poorly developed, but there has been a spectacular late Tertiary radiation of many groups…Land connections between Australia and New Guinea are also suggested by the mid-Pliocene appearance of the marsupials…The island as a whole is a region of faunal and floral mixing, survival, and evolution in the middle to late Tertiary” (Raven and Axelrod 1972). “The consensus of contemporary zoogeographic studies is that though New Guinea and its shelf islands were joined to Australia during several phases of the Pleistocene, New Guinea has a basic, rich biota, much more ancient in origin as well as evolutionary history, and independent of Australian faunal or floral influence” (Udvardy 1975). Some data on the glaciation and climate change in New Guinea we find in Löffler (1982): The Pleistocene glacial history of New Guinea may date back to about 700000 -+ 100 000 years BP when lava erupted under ice on Mt Giluwe....The last glaciation lasted until about 15 000 years BP when the ice receded rather rapidly to completely disappear by about 9000 years BP…The snowline during the last glaciation was at about 3550 – 3600 m altitude indicating a lowering of the snowline by about 1000 – 1100 m. This would correspond to a temperature depression of about 5–6 oC assuming that precipitation was similar to the present. 8.18 The Arachnida of the Papuan Subregion 747 On the summit of Jaya (Carstensz), the glacial ice covers 6.9 km2 (16.4 km2 in the middle of the nineteenth century), retreating rapidly. Its thickness is ca. 40 m. Some sources: Axelrod and Raven (1982), Chase (1971), Coleman (1980), Doutch (1972), Gressitt (1956, 1958, 1967, 1971, 1974, 1982a, 1982b), Heads (2002, 2009), Kemp (1984), Krijanovskiy (1980), Mitchell and Warden (1971), Raven and Axelrod (1972), Robbins (1971), Solem (1958), Smith (1990), Taylor, Goodliffe and Martinez (1999), Thompson (1967), Whitmore (1981). Arachnogeography New Guinea, Bismarck, and Salomon islands are with different positions in the biogeographical subdivision of Australasia, view by botanists, mammalogists, ornithologists, and entomologists. According to the botanists and entomologists, they are clearly part of the Indomalayan region. Krijanovskiy (1980) also includes them into his Paleotropical dominion, together with tropical Africa, Madagascar, and the Indomalayan region. The peculiar high-ranking vertebrates send Papuan area into the Notogea (Australian Realm). What concerns the Arachnida, our conclusion, is that the level of endemism is much lower than in the mammals or birds. In the Papuan area (still very understudied) are found the orders Palpigradi, Scorpiones, Pseudoscorpiones, Opiliones, Amblypygi, Uropygi, Schizomida, Araneae, and many groups of Acari. Most interesting is the order Holothyrida, represented in New Guinea by 11 species of 3 endemic genera of the family Holothyridae. Absent from New Guinean area are the orders Ricinulei, Solifugae, and Opilioacarida and the suborders Amblypygi, Mesothelae, and Dyspnoi. Palpigradi Recorded are only two spp.: Eukoenenia cf. lawrencei and Koeneniodes cf. frondiger. (Eukoeneniidae) (Condé 1980, 1981). Scorpiones According to the monograph of Koch (1977) and the older papers of Takashima (1948, 1950), in New Guinea (incl. Aru and Bougainville) live only six widespread species of the genera Isometrus, Lychas (Buthidae), and Liocheles (Hormuridae). They are shared with Australia. From Rennell Island (the Solomon Islands) have been found three spp., including the newly described species (endemic) Liocheles penta Francke et Lourenço (1991) (Browning and Hyatt 1957). Pseudoscorpiones Beier (1940, 1965b, 1967, 1968, 1982) brought the pseudoscorpions from Papua New Guinea (incl. Bismarck Islands to 60 species of 36 genera and 12 families) (Chthoniidae, Tridenchthoniidae, Syarinidae, Cheiridiidae, Garypinidae, Geogarypidae, Atemnidae, Olpiidae, Sternophoridae, Cheliferidae, Chernetidae, Withiidae). Other data are provided by Beier (1940, 1982) and Morikawa (1963). Beier (1966b) lists 24 genera and 8 families of pseudoscorpions from the Solomon Islands (incl. Bougainville and Buka Islands, which belong to Papua New Guinea), mostly from Guadalcanal (26 spp.). Now the known species of 748 8 Some Peculiarities of the Distribution of Arachnida Pseudoscorpiones in the islands are 38, including some endemics. Species living on both Solomon Islands and New Guinea and the Bismarcks are Acanthicochernes biseriatus Beier (1964), and the common genus of both is Gelachernes Beier (1940) – four spp. (Chernetidae). Endemic genus for New Guinea and the Bismarck Archipelago: Fam. Cheliferidae Papuchelifer Beier, 1965 – Indonesia (Papua), Papua New Guinea (three spp.) Endemic genus for the Solomon Islands: Fam. Chernetidae Cyclochernes Beier, 1970 – Guadalcanal (one sp.) The known pseudoscorpions from Vanuatu (New Hebrides) are only four species of four genera and the families Atemnidae, Cheliferidae, and Chernetidae. Lissochelifer insularis (Beier) is common in New Guinea and Paratemnoides salomonis and Haplochernes hebridicus in the Solomon Islands. Only Hebridochernes paradoxus is an endemic species to Vanuatu. On Fiji are registered three spp. of the genera Geogarypus (Geogarypidae) and Haplochernes (Chernetidae). Geogarypus longidigitatus (Rainbow, 1897) and Haplochernes funafutensis (With, 1907) were described from Funafuti (Tuvalu) and Haplochernes ellenae Chamberlin (1938) from Viti Levu (Fiji). Opiliones From Melanesia have been recorded Opiliones of the families Stylocellidae, Sclerosomatidae, Assamiidae, Podoctidae, Epedanidae, and Samoidae, following Clouse and Giribet (2007), Forster (1949b), Goodnight and Goodnight (1947), Loman (1906), Roewer (1911, 1912, 1913, 1915b, 1935–1936, 1949, 1963), and Suzuki (1982). The collection of P. Beron from New Guinea, New Britain, and New Ireland is still under study. Cyphophthalmi Until recently, Cyphophthalmi were not registered east of the Lyddeker’s Line. However, two species of Stylocellus Westwood, 1874, genus known from Malaysia, Indonesia, Singapore, and Palawan (Philippines), have been found in Papua (Indonesian New Guinea) (Clouse and Giribet 2007). These animals don’t cross saltwater areas, so a problem arises about their establishment in New Guinea. Eupnoi Fam. Sclerosomatidae – Gagrella Stoliczka Dyspnoi – Not recorded from Melanesia Laniatores In Melanesia are recorded Laniatores of the families Assamiidae, Podoctidae, Epedanidae, Samoidae, and Zalmoxidae. Endemic genera are: Fam. Assamiidae Granobunus Roewer, 1912 – New Guinea (one sp.) Heteropygoplus Roewer, 1915 – New Guinea (one sp.) 8.18 The Arachnida of the Papuan Subregion 749 Apygoplus Roewer, 1912 – New Guinea (three spp.) Macrodampetrus Roewer, 1915 – New Guinea (two spp.) Metadampetrus Roewer, 1915 – New Guinea (two spp.) Metamosoia Roewer, 1915 – New Guinea (one sp.) Mosoia Roewer, 1912 – New Guinea (one sp.) Neonothippus Roewer, 1912 – New Guinea (one sp.) Fam. Podoctidae (incl. Erecananinae and Ibaloniinae) Asproleria Roewer, 1949 – New Guinea (one sp.) Heteroibalonius C.J. Goodnight et M.L. Goodnight, 1947 – New Guinea (one sp.) Heteropodoctis Roewer, 1912 – New Guinea (one sp.) Orobunus Goodnight et Goodnight, 1947 – (one sp.) Paramesoceras Roewer, 1915 – New Guinea (one sp.) Podoctinus Roewer, 1923 – New Britain (one sp.) Proholozoster Roewer, 1915 – New Guinea (one sp.) Santobius Roewer, 1949 (Mesoceras Soerensen, 1886, preoccup.,=Mesoceratula Roewer, 1949) – Vanuatu, Fiji (two spp.) Fam. Samoidae Badessa Sørensen, in L. Koch 1886 – Fiji (one sp.) Fijicolana Roewer, 1963 – Melanesia (one sp.) Fam. Zalmoxidae – In New Guinea and the nearby islands live 26 spp. of the genus Zalmoxis Sorensen (the only Zalmoxis in the Eastern Hemisphere). Amblypygi In Melanesia are represented four genera of Amblypygi (Dunn 1949; Gravely 1915; Rahmadi and Kojima 2010; Thorell 1888): Fam. Charinidae Charinus Simon – Ch. australianus (L. Koch) (Samoa, Fiji), Ch. pescotti Dunn (Solomon Island) Sarax Simon – S. monodenticulatus Rahmadi et Kojima (Papua, Waigeo), S. newbritainensis Rahmadi et Kojima (New Britain, New Ireland), S. sarawakensis (Thorell) (New Guinea, Solomon Island), S. willeyi Gravely (New Britain), S. subterraneus Thorell (papuanus) (New Guinea, Maluku). Fam. Charontidae Charon Karsch – Ch. grayi (Gervais) (? = papuanus Thorell) (New Guinea) Stygophrynus Kraepelin – S. (Neocharon) forsteri Dunn (Solomon Island, endemic for Guadalcanal) Thelyphonida (Uropygi) On the island of New Guinea and nearby islands of Fergusson, New Ireland, and New Britain live three species of Abaliella Strand. One Thelyphonus was described by Rowland (1973a) from Guadalcanal (Solomon Islands), added to the other species from the Solomon’s Thelyphonus leucurus 750 8 Some Peculiarities of the Distribution of Arachnida Pocock (1898). The widespread Minbosius manilanus (C.L. Koch) lives on the Moluccas, in New Guinea, and the Philippines. All of them belong to Thelyphonidae and Thelyphoninae, typical for the Papuan area, Samoa, the Philippines, and Southeast Asia. Haupt (2009a) synonymized Abaliella Strand, Minbosius Speijer, and Tetrabalius Thorell with Thelyphonus Latreille. Schizomida Schizomus modestus Hansen, 1905, was described from New Guinea and recorded also in New Britain and W. Malaysia (unverified record of Buxton (1917)). Hubbardiidae indet. are known from New Guinea, New Britain, and New Ireland (Reddell and Cokendolpher, 1995). In my collection in Sofia, there are schizomids, collected by me in New Ireland in 1975. Araneae The Spiders of New Guinea In New Guinea and the Bismarck Archipelago (incl. Buka and Bougainville) have been recorded spiders of at least 50 families (Balogh 1936; Baehr and Baehr 1993; Baert 1979, 1980, 1982, 1984; Bourne 1980; Chrysanthus, many papers since 1958; Lehtinen and Saaristo 1980; Main 1982; Robinson 1982; Thorell 1881; Versteirt et al. 2010; York 1982). The collection, brought by me (43 spp. of 15 families), was identified only partially by Brignoli (1981), but his untimely death did not allow him to describe the many new taxa in this collection. It has noted that some species related to groups believed “typical” of New Zealand (Stiphidiidae). The preliminary conclusion of Brignoli (1981) is “It is still too early to appreciate the value of the proposal of Lehtinen (1980), of abandoning the traditional Oriental and Austral regions in favour of an Indo-Pacific and a South Gondwanan region, but I would like to observe that, if it is probably possible to accept the limited value of Wallace’s and Weber’s lines for spiders, I do not see every definite border between the two new regions proposed by Lehtinen.” Several families of spiders are known from Australia (Zoropsidae, Synotaxidae), but not from New Guinea. Others (Telemidae, Ochyroceratidae, Stenochilidae), on the contrary, have been recorded from New Guinea, but not from Australia, despite the proximity of both territories. None of the many Zoropsidae described in Queensland had ventured across the Torres Strait. The list of Araneae in Fiji contains 122 spp. of spiders (part of them indet.), of 35 families (Evenhuis 2006). Lakarobius Berry et al. (Salticidae) is an endemic genus. Endemic genera of spiders in the “Papuan area” are: Fam. Telemidae Jocquella Baert, 1980 – New Guinea 8.18 The Arachnida of the Papuan Subregion Fam. Theridiidae (Hadrotarsinae) Yoroa Baert, 1984 – New Guinea Fam. Barychelidae Fijocrypta Raven, 1994 – Fiji Fam. Anapidae Guiniella Rix et Harvey, 2010 – New Guinea (one sp.) Fam. Philodromidae Pseudopsellonus Balogh, 1936 – New Guinea (one sp.) Senoculifer Balogh, 1936 – New Guinea (three spp.) Fam. Clubionidae Arabellata Baert, Versteirt et Jocqué, 2010 – New Guinea Invexillata Versteirt, Baert and Jocqué, 2010 – New Guinea Fam. Mysmenidae Kekenboschiella Baert, 1982 – New Guinea (four spp.) Fam. Pisauridae Campostichommides Strand, 1911 – New Guinea (Kei Island) Hesydrimorpha Strand, 1911 – New Guinea Fam. Selenopidae Godumops Crews et Harvey, 2011 – New Guinea (one sp.) Fam. Uloboridae Lubinella Opell, 1984 – New Guinea Fam. Pisauridae Hesydrimorpha Strand, 1911 – New Guinea Papakula Strand, 1911 – New Guinea (one sp.) Fam. Sparassidae Clastes Walckenaer, 1837 – Moluccas, New Guinea (one sp.) Exopalistes Hogg, 1914 – New Guinea Strandiellum Kolosváry, 1934 – New Guinea Fam. Stiphidiidae Asmea Gray et Smith, 2008 – New Guinea (four spp.) 751 752 8 Some Peculiarities of the Distribution of Arachnida Fam. Lycosidae Satta Lehtinen et Hippa, 1979 – New Guinea Fam. Thomisidae Cetratus Kulcziński, 1911 – New Guinea Latifrons Kulcziński, 1911 – New Guinea Rhaebobates Thorell, 1881 – New Guinea (two spp.) Fam. Salticidae Allococalodes Wanless, 1982 – New Guinea Aruana Strand, 1911 – New Guinea, Aru Bulolia Zabka, 1996 – New Guinea Chalcolemia Zhang et Maddison, 2012 – New Guinea [Coccorchestes Thorell, 1881 – 38 spp. from New Guinea, 1 from New Britain, 1 from Queensland Cucudeta Maddison, 2009 – New Guinea Diolemus Thorell, 1870 – Sixteen spp. in New Guinea, two in Moluccas Furculattus Balogh, 1980 – New Guinea, New Britain (one sp.) Leptathamas Balogh, 1980 – New Guinea Opistoncana Strand, 1913 – New Ireland Papuaneon Maddison, 2016 – New Guinea Paraharmochirus Szombathy, 1915 – New Guinea Porius Thorell, 1892 – New Guinea (two spp.) Saitissus Roewer, 1938 – New Guinea Tabuina Maddison, 2009 – New Guinea (three spp.) Tarodus Pocock, 1899 – New Britain Variratina Zhang et Maddison, 2012 – New Guinea Viribestus Zhang et Maddison, 2012 – New Guinea Zabkattus Zhang et Maddison, 2012 – New Guinea Fam. Corinnidae Melanesotypus Raven, 2015 – Solomon Islands Holothyrida With 11 out of 29 species in the order Holothyrida, the island of New Guinea looks like the center of speciation of these strange arachnids. All species are endemic to New Guinea: Hammenius fujuge fujuge, H. fujuge iora, H. ingii, H. insularis, Hammenius grandjeani grandjeani, Leiothyrus armatus, L. holthuisi, L. nitidissimus, Thonius berlesei, Th. longipes, Th. montanus, and Th. mendi. So are the genera Hammenius Lehtinen and Thonius Lehtinen. Hammenius niger (Thon) lives on Silhouette Island (the Seychelles), a highly interesting distribution for these conservative animals (Thorell 1882, Beron 2014; Canestrini 1897; Lehtinen 1981, 1995; Van der Hammen 1983). 8.19 Arachnogeography of Australia and Tasmania 8.19 753 Arachnogeography of Australia and Tasmania 8.19.1 Australia 8.19.1.1 Geography, General Zoogeography, and Paleogeography The continent of Australia has an area of 7 692 000 km2 (without Tasmania but including the offshore islands). The highest point is Mount Kosciuszko (2228 m.). The continent was detached from Antarctic some 95–80 Ma (Late Cretaceous) and drifted northward, coming close to the equator and to Sundaland, the presentday Indochina, and the Greater Sunda islands. The present-day Australia consists of several subregions of very different landscape. Huge areas in Western and central Australia are deserts and form the so-called Eremial. The northernmost part of the continent, on Torres Strait, consists of dry savanna (in the Western part) and of rainforests similar to the forests in New Guinea (in the Eastern part). Udvardy (1975) notes that Usinger (1963) attached Cape York (the northern peninsula of Queensland) to his Papuan subdivision of the Oriental region, but Gressitt (1961) treated it as a clearly transitional area together with southern New Guinea and in 1975 still opines (in a letter to Udvardy) that “the overlap of Australian and Oriental in southern New Guinea and Northern Australia needs to be shown as an overlap zone with dominance of Oriental elements….” Prof. Gressitt told the same personally to the present author during a visit in Wau in the same year (1975). However, this remark concerns mostly to the flora and the invertebrates, especially the insects. Speaking of vertebrates, Papuan Subregion is clearly part of the Australian region (the differences are between kingdoms!). To quote partly Keast (1959): New Guinea and Tasmania, lying on the Australian continental shelf, would be brought into contact by a fall in sea-level of about 10 m and 50 m, respectively. They are known to have been joined to Australia during part of the Tertiary and apparently twice during the Pleistocene…This being so, it would seem surprising that the flora of New Guinea bears little resemblance to that of Australia…and much of its invertebrate fauna is likewise IndoMalayan…In explanation of this it has been suggested that the geologically ancient northern part of New Guinea was once, presumably in the Mesozoic, in direct land contact with Asia … Another proposal is that Australia did not always lie in close proximity to New Guinea but ‘drifted’ into it. According to Keast (1959), “approximately one-third of Australian continent lies within the tropics and the rest is either temperate or sub-temperate.” Special adaptations to cold are little developed in Australian animals. Also after this author, Australia has not been in direct land contact with Asia since the beginning of Tertiary, a period of over 50 Ma. Actually, after the modern concepts, Australia has never been in direct contact with Asia, as it drifted from the south. If for the final phase of extinction of the giant animals in Australia (Diprotodon, Palorchestes, Thylacoleo) could be blamed the combination of aridity and the arrival 754 8 Some Peculiarities of the Distribution of Arachnida of aboriginal man and the dingo, eventual changes in the composition of the arachnofauna are to be explained only with the aridization. Some sources: Brown, Campbell and Crook (1968), Embleton (1973), Filewood (1984), Keast (Ed) (1981), McElhinny et al. (1976), Veevers and McElhinny (1976), Walker (Ed.)(1972), Weissel et al. (1976). Several attempts have been made to subdivide Australia zoogeographically (Hedley 1894; Spencer 1896; Sloane 1915; Campbell 1943; Udvardy 1975). The history of the regionalization has been narrated by Ebach (2012). A recent subdivision is due to Ebach, Gill, Kwan, Anyong, Murphy, and Cassis (2013). On the mainland are outlined 15 regions; Tasmania is a subregion. The zoogeographical subdivision of Ebach et al. (2013) of the “Australian Sub-realm”: Sub-realm Australia Bassian Region Subregions: Southeastern Forest, McPherson-MacPherson-Macleay, Adelaide, Tasmania Torresian-Bassian Region Subregions: Atherton Plateau, Cape York Peninsula, Arnhem Land, Kimberley Plateau Eyrean Region Subregions: Northern Desert, Eastern Desert, Pilbara, Western Desert, Eyre Peninsula, South-West Australia, Riverina “Breakup between Australia-Antarctica and the northern part of Greater India commenced ca. = 130 Ma, and between Australia and Antarctica around 96 Ma. At the beginning of the Palaeogene, Australia commenced its northward drift towards its present position. Seafloor spreading between Australia and Antarctica was at first slow, but increased to ca. 5 cm. per year around 45 Ma. By 35 Ma, the Antarctic Circumpolar Current became established, thereby triggering glaciation in Antarctica. Northern Australia reached the tropics by the beginning of the Miocene, and Australia has progressively moved northwards at 7–8 cm per year since” (Li and Powell 2001). Arachnogeography Concerning the vertebrates, Australia is a continent with many endemic high-ranking taxa, and many groups are lacking in the native fauna. About Arachnida there are no endemic orders or suborders, and lacking are only a few: the orders Ricinulei, Thelyphonida (Uropygi), and Solifugae and the suborders Paleamblypygi, Dyspnoi, and Mesothelae. Some of the 11 orders, represented on the continent, contain endemic taxa, but of low rank, incomparable with the situation with the vertebrates. Palpigradi, Amblypygi, and Opilioacarida are represented with only several species. The scorpions are of four families (1 family is endemic, 42 spp.); Pseudoscorpions (150 spp.) and Schizomida (53 spp., 9 genera, mostly endemic) are well studied, thanks, besides Beier and other older authors, mainly to the energetic researcher M. Harvey and his team. On the continent there are also good specialists of opilions, spiders, and many groups of Acari. Still, very few high- 8.19 Arachnogeography of Australia and Tasmania 755 ranking endemic taxa have been found. There is 1 subendemic (shared with New Zealand) family of Holothyrida; 5 endemic genera of Pseudoscorpiones; 20 end. genera of Opiliones; 338 genera of Araneae, of which 88 (26%) are endemic; and many endemic genera of Acari. Yeates, Harvey, and Austin (2003) tried to make an actual (2003) estimate of the number of Arachnida in Australia and to predict the real number of different orders, living there. Here is their table (after 13 years, this should be modified somehow (added sp. in brackets)). Taxa Arachnida total Palpigrada Amblypygi Schizomida Pseudoscorpiones Opiliones Acarina Described Australian species 5705 1 (native) 4 46 (now 52) 181 194 2871 Estimated total Australian species 42 195 5 10 80 750 1200 20 000 Palpigradi Besides Eukoenenia mirabilis (imported), from Western Australia was described an indigenous member of Palpigradi: Eukoenenia guzikae Barranco et Harvey, 2008. Scorpiones In his monograph, Koch (1977) enumerates for the “Australo-Papuan Region” 29 spp. of scorpions from the genera Cercophonius (Bothriuridae), Lychas, Isometroides, Isometrus (Buthidae), Liocheles (Hormuridae), and Urodacus (Urodacidae). Endemic family is Urodacidae and end. genera are Urodacus Peters with 19 species (incl. 7 newly described by L.E. Koch) and Isometroides Keyserling. Cercophonius Peters with one sp. (Australia and Tasmania) has been recently found in the Himalaya. As there are no strictly Papuan species (not found in Australia), this is the final figure 32, to add the new genus and sp. Aops oncodactylus Volschenk et Prendini, 2008, and the two new Urodacus, described by Volschenk, Smith, and Harvey (2000) and Volschenk, Harvey, and Prendini (2012) of species of scorpions on the Australian mainland or in Tasmania (the last has no endemic scorpion and shares with the mainland Cercophonius squama). We should extract from the 32 spp. Lychas variatus (also in Fiji) and the widespread Isometrus maculatus, I. melanodactylus, Liocheles australasiae, L. waigiensis, and L. karschii. Remain the 26 species, endemic to Australia (Cercophonius squama also in Tasmania). Australia and New Guinea actually share only the six widespread species of Isometrus, Lychas, and Liocheles. Now (2017) some of the family names are different. To quote from the analysis of Koch (1977): “The highly speciated urodacine scorpionid genus Urodacus is widespread in Australia but absent from both New Guinea and Tasmania. Because of the time required for the evolution of a genus, there are two possibilities: either the genus Urodacus has always been confined to inland Australia, or there have been species outside Australia and these became extinct. The exclusion 756 8 Some Peculiarities of the Distribution of Arachnida from Tasmania of the genera Urodacus and Lychas…could be due to cold. There is however an absence of factors…which might exclude Urodacus from New Guinea. I consider therefore that either its species in the more peripheral northern areas of the Australian mainland have reached there relatively recently (since the last major transgression of the sea) or that species in New Guinea have been eliminated.” In his article on the zoogeography of Australian scorpions, Koch (1981) distributed the scorpion taxa in three “levels of endemism”: 1. Cosmopolitan species or non-endemic species shared with Asia: Isometrus maculatus, Liocheles australasiae, L. waigiensis, L. karschii 2. Species that have arisen in Australo-Papua from non-endemic genera: Lychas (three spp.), Isometrus (one sp.) 3. Species that have arisen in Australo-Papua: Isometroides (one sp.), Cercophonius (one sp.), Urodacus (20 spp.) He also classified the genera regarding the continental drift: 1. Genera derived from an ancient stock with related genera only in South America: Cercophonius 2. Australian genus with less certain relationships and probably ultimately derived from Asia: Urodacus 3. Genera in common with Asia: Isometrus, Lychas, Liocheles 4. Genus clearly derived from Lychas in Australia: Isometroides Further, the scorpions are distributed into three groups: Southern, Central, and Northern species. Endemic family is Urodacidae. Pseudoscorpiones In Australia (Tasmania excluded) have been registered 181 spp. of Pseudoscorpions of 59 genera and 20 families (Beier 1948, 1954, 1966, 1975; Edwards and Harvey 2008; Harvey 1992, 2006, and many others, Harvey and Edward 2007; Harvey and Mei Chen Leng 2008a, b; Harvey and Muchmore 1990; Hoff 1947; Harvey 2013f, actualized). Endemic genera of pseudoscorpions in Australia (without Tasmania) are: Fam. Menthidae Theхmus Harvey, 1990 (two spp.) Fam. Olpiidae Austrohorus Beier, 1966 – (one sp.) Barbaraella Harvey, 1005 – (one sp.) Linnaeolpium Harvey et Leng, 2008 – (one sp.) Fam. Cheliferidae Australochelifer Beier, 1975 – (one sp.) Fam. Chernetidae 8.19 Arachnogeography of Australia and Tasmania 757 Conicochernes Beier, 1948 – (four spp.) Marachernes Harvey, 1992 – (three spp.) Opiliones In Australia (without Tasmania) are known ten indigenous families of Opiliones. Cyphophthalmi In Australia have been registered nine spp. of Cyphophthalmi of two genera of the family Pettalidae (Boyer and Giribet 2007; Clouse and Giribet 2007; Giribet 2003; Juberthie 1988b). The genera are endemic in Queensland (Austropurcellia Juberthie, 1988, six spp.) and Western Australia (Karripurcellia Giribet, 2003, three spp.). Eupnoi (Forster 1949a, 1952b, 1955; Hunt 1971, 1985, 1992; Hunt and Cokendolpher 1991; Hunt and Hickman 1993; Kauri 1954; Roewer 1914–1949; Shear 1996, 2001; Taylor 2009, 2011; Hickman 1957, 1958; Hogg 1909; Hunt 1990, 1995) Fam. Neopilionidae – Arrallaba Hunt et Cokendolpher (one sp.), Ballarra Hunt et Cokendolpher (six spp.), Plesioballarra Hunt et Cokendolpher (one sp.), Hypomegalopsalis Taylor, 2011 (one sp.) Fam. Sclerosomatidae – Gagrella Stoliczka (one sp.) Genus Hesperopilio Shear (WA, one sp.) was nested in Phalangioidea by Groh and Giribet (2015) but with unclear family belonging. Dyspnoi – Fam. Acropsopilionidae-Acropsopilio Silvestri, 1905, Queensland (one sp.) [Austropsopilio Forster, 1955 (syn. Tasmanopilio Hickman, 1957), NSW, Queensland (two endemic spp., the genus known also from Tasmania)] Laniatores In Australia (without Tasmania) are known Laniatores of 54 genera (39 genera in Triaenonychidae) and 6 families (Assamiidae, Phalangodidae (misplaced?), Podoctidae, Samoidae, Zalmoxidae, Triaenonychidae). Endemic genera of Opiliones Laniatores for Australia (Tasmania excluded) are: Fam. Triaenonychidae Breviacantha Kauri, 1954 (one sp.); Callihamina Roewer, 1942 (one sp.); Callihamus Roewer, 1931 (one sp.); Cluniella Forster, 1955 (three spp.); Heteronuncia Roewer, 1920 (one sp., Queensland); Holonuncia Forster, 1955 (13 spp., ACT, NSW, Victoria); Perthacantha Roewer, 1931 (one sp.); Yatala Roewer, 1942 (SA, one sp.); Conoculus Forster, 1949 (one sp.); Dingupa Forster, 1952 (one sp., WA); Dipristes Roewer, 1931 (one sp.) Amblypygi In Australia (the Northern Territory, Queensland, and the Australian Christmas Island) are known six spp. of Amblypygi of the genera Charon Karsch (three spp.) and Charinus Simon (three spp.) (Charinidae). They are proper only to the tropical part of the continent (Dunn (1949), Harvey (1985), Harvey et al. (2012), Harvey and West (1998), Monteith (1965). All species are (so far) endemic to the areas of description. From Western Australia has been recorded another amblypygid (Harvey et al. 2012). 758 8 Some Peculiarities of the Distribution of Arachnida Map 8.8 Distribution of Amblypygi and Schizomida in Australia Amblypygi Charon – ▲ Charinus – ▼ Schizomida Draculoides – ■ Apozomus – □ Bamazomus – ◊ Notozomus – ♠ Julattenius – Brignolizomus – ◄ Ovozomus – ► Paradraculoides – Ω Attenuizomus – ♦ 8.19 Arachnogeography of Australia and Tasmania 759 Thelyphonida (Uropygi) – Not recorded in Australia. Schizomida The first member of order Schizomida from Australia (a troglobitic one) was described by Harvey (1988), despite the fact that the order was recorded from this continent as early as 1963. In his important paper in 1992, Harvey described 5 new genera (Draculoides, Apozomus, Bamazomus, Notozomus, and Julattenius) and 24 new species of Schizomida. From the map (p. 79), it is clear that all these species are known from the periphery of the continent: the far west, far north, and easternmost parts of Australia. Later Harvey (2000) added two more genera Brignolizomus and Attenuizomus, with three new species. New descriptions followed (Harvey (1992, 2000a, 2000b, 2001b, 2013g); Harvey and Humphreys (1995); Harvey et al. (2008). Now (2017) in Australia are known 56 spp. of the genera Draculoides (6), Apozomus (11), Bamazomus (5), Notozomus (17), Julattenius (2), Brignolizomus (3), Ovozomus (1), Paradraculoides (3), and Attenuizomus (4) (endemic genera in bold). Araneae Australia (continental) Following the table of Main (1981) in Australia are registered 46 families of spiders. Without Salticidae, they contained 276 indigenous genera, of which 77 (27.9%) are endemic; including the Salticidae with 338 genera of which 88 (26%) are endemic. The family Salticidae contained 62 genera, of which 11 are endemic. Other families with many genera are Araneidae (33, 6 end.), Lycosidae (25, 2 end.), and Thomisidae (23, 5 end.). Since 1981 many new records of spiders in Australia have changed these numbers. Now (2017) in Australia are registered 73 families. The families Gradungulidae, Periegopidae, Holarchaeidae (Tasmania), Malkaridae-Pararchaeidae (also New Caledonia), and Nicodamidae (also New Guinea) are shared only (or almost only) with New Zealand; the families Hexathelidae (Hexathelinae), Migidae (Calathotarsinidae), Malkaridae, and Amphinectidae are shared with southern South America (Argentina and Chile) (Baehr 2003, b, c, 2004a, b; Baehr and Baehr 1987; Baehr and Churchill 2003; Baehr and Jocqué 2000; Balogh 1980; Crews and Harvey 2011; Davies 1995a, b, 1998, 2003a, b; Dunn 1951; Framenau 2006a, b; Gardziñska 1996; Gardzinska and Zabka 2010; Gray 1973; Harms and Harvey 2009, 2013; Harvey 1995; Hawkeswood 2003; Huber 2001; Jocqué 1993; Koch 1871–1881; Main 1969, 1976, 1981a, 1981b, 1987; Patoleta 2002; Patoleta and Zabka 1999; Platnick 1991, 2000, 2002, 2004; Platnick and Baehr 2006; Platnick and Forster 1989; Raven (1976, 1978, 1979, 1980a, 1980b, 1981a, 1981b, 1985, 1994, 2009), Raven and Stumkat (2005); Richardson and Zabka 2004; Richardson et al. 2006; Rix 2005, 2006; Rix and Harvey 2010, 2010b, 2011, 2012a, 2012b, 2012c; Rix et al. 2010; Thorell 1881; Wanless 1988; Zabka 1987a, b, 1988, 1989, 1990a, b 1991a, b, 1992a, b, 1993b, 1994, 1995b, 1999, 2000, 2001, 2002, 2003, 2006a, b, 2009, 2012, 2014). 760 8 Some Peculiarities of the Distribution of Arachnida Endemic genera of spiders in Australia (Tasmania generally excluded) are: Fam. Hexathelidae Hexathelinae – Atrax O. P-Cambridge, 1877 (3 spp.); Bymainiella Raven, 1978 (4 spp.); Hadronyche L. Koch, 1873 (31 spp., incl. 1 from Tasmania); Paraembolides Raven, 1980 (8 spp.); Teranodes Raven, 1985 (2 spp.) Fam. Dipluridae Subfam. Euagrinae – Australothele Raven, 1984 (7 spp.); Carrai Raven, 1984 (1 sp.); Cethegus Thorell, 1881 (12 spp.); Namirea Raven, 1984 (7 spp.) incertae sedis Troglodiplura Main, 1969 (one sp.) Fam. Anapidae – Chasmocephalon O. P.-Cambridge, 1889 (eight spp.); Maxanapis Platnick et Forster, 1989 (nine spp.); Nortanapis Platnick et Forster, 1989 (one sp.); Octanapis Platnick et Forster, 1989 (two spp.); Queenslanapis Platnick et Forster, 1989 (one sp.); Risdonius Hickman, 1939 (three spp.); Spinanapis Platnick et Forster, 1989 (nine spp.); Victanapis Platnick et Forster, 1989 (one sp.) Subfam. Micropholcommatinae: Micropholcomma Crosby et Bishop, 1927 (8 spp., with Tasmania) Fam. Archaeidae – Austrarchaea Forster et Platnick, 1984 (27 spp.) Fam. Malkaridae – Malkara Davies, 1980 (1 sp.), Perissopmeros Butler, 1932 (7 spp.); syn. Pararchaeidae: Anarchaea Rix, 2006 (4 spp.); Flavarchaea Rix, 2006 (7 spp.); Nanarchaea Rix, 2006 (2 spp.); Ozarchaea (16 spp.); Westrarchaea Rix, 2006 (3 spp.) Fam. Nemesiidae – Aname L. Koch, 1873; Chenistonia Hogg, 1901; Ixamatus Simon, 1887; Kwonkan Main, 1983; Merredinia Main, 1983; Namea Raven, 1984; Pseudoteyl Main, 1985; Teyl Main, 1975; Teyloides Main, 1985; Xamiatus Raven, 1981; Yilgarnia Main, 1986 Fam. Synotaxidae – Calcarsynotaxus Wunderlich, 1995 (two spp.); Microsynotaxus Wunderlich, 2008 (two spp.); Paratupua Platnick, 1990 (one sp.) Fam. Cyatholipidae – Forstera Koçak et Kemal, 2008 (one sp.); Matilda Forster, 1988 (one sp.); Teemenaarus Davies, 1978 (one sp.); Tekellatus Wunderlich, 1978 (one sp.) Fam. Theridiidae – Magnopholcomma Wunderlich, 2008 (one sp.) Fam Ctenidae – Bengalla Gray et Thompson, 2001 (one sp.), Janusia Gray, 1973 (one sp.) Fam. Lycosidae – Anomalosa Roewer, 1960 (two spp.); Costacosa Framenau et Leung, 2013 (two spp.); Hoggicosa Roewer, 1960 (ten spp.); Kangarosa Framenau, 2010 (ten spp., one in Tasmania); Knoelle Framenau, 2006 (one sp.); Mainosa Framenau, 2006 (one sp.); Tapetosa Framenau et al., 2009 (one sp.); Tuberculosa Framenau et Yoo, 2006 (four spp.); Venator Hogg, 1900 (one sp.) Fam. Arkyidae – Arkys Walck., 18370, Demendians Strand, 1929 8.19 Arachnogeography of Australia and Tasmania 761 Fam. Pholcidae – Wugigarra Huber, 2001 (20 spp.); Trichocyclus Simon, 1908 (23 spp.); Micromerys Bradley, 1877(7 spp.); Panjange Deeleman-Reinhold et Deeleman, 1983 (6 spp.) Fam. Oxyopidae – Pseudohostus Rainbow, 1915 (one sp.) Fam. Pisauridae – Inola Davies, 1982 (three spp.) Fam. Psechridae – Baiami Lehtinen, 1967 (9 spp.); Barahna Davies, 2003 (8 spp.); Borrala Gray et Smith, 2004 (4 spp.); Carbinea Davies, 1999 (4 spp.); Corasoides Butler, 1929 (1 spp.); Couranga Gray et Smith, 2008 (2 spp.); Elleguna Gray et Smith, 2008 (2 spp.); Jamberoo Gray et Smith, 2008 (4 spp.); Kababina Davies, 1995 (9 spp.); Karriella Gray et Smith, 2008 (2 spp.); Malarina Davies et Lambkin, 2000 (4 spp.); Malarina Davies et Lambkin, 2000 (4 spp.); Procambridgea Forster et Wilton, 1973 (12 spp., one of them also in NZ – transported?); Tartarus Gray, 1973 (4 spp.); Therlinya Gray et Smith, 2002 (11 spp.); Wabua Davies, 2000 (11 spp.) Fam. Miturgidae (Zorinae) – Elassoctenus Simon, 1909 (one sp.); Hestimodema Simon, 1909 (two spp.); Simonus Ritsema, 1881 (one sp.); Thasyraea L. Koch, 1878 (two spp.); Thasyraea L. Koch, 1878 (three spp.); Tuxoctenus Raven, 2008 (three spp.) Fam. Zoropsidae – Birrana Raven et Stumkat, 2005 (one sp.); Huntia Gray et Thompson, 2001 (two spp.); Kilyana Raven et Stumkat, 2005 (ten sp.); Krukt Raven et Stumkat, 2005 (five spp.); Megateg Raven et Stumkat, 2005 (eight spp.) Fam. Amphinectidae – Austmusia Gray, 1983 (three spp.); Buyina Davies, 1998 (two spp.); Cunnawarra Davies, 1998 (two spp.); Jalkaraburra Davies, 1998 (one sp.); Keera Davies, 1998 (one sp.); Magua Davies, 1998 (one sp.); Penaoola Davies, 1998 (two spp.); Quemusia Davies, 1998 Fam. Amaurobiidae – Bakala Davies, 1990 (1 sp.); Dardurus Davies, 1976 (6 spp.); Daviesa Koçak et Kemal, 2008 (2 spp.); Jamara Davies, 1995 (1 sp.); Manjala Davies, 1990 (3 spp.); Midgee Davies, 1995 (9 spp.); Storenosoma Hogg, 1900 (13 spp., 1 of them in Tasmania); Wabarra Davies, 1996 (2 spp.) Fam. Desidae – Colcarteria Gray, 1992 (three spp.); Forsterina Lehtinen, 1967 (eight spp. in mainland Australia, one in New Caledonia); Lathyarcha Simon, 1908 (three spp.); Paramatachia Dalmas, 1918 (five spp., one of them shared with Tasmania); Phryganoporus Simon, 1908 (five spp., one of them shared with Tasmania); Pitonga Davies, 1984 (one sp.) Fam. Dictynidae – Callevophthalmus Simon, 1906 (two spp., one shared with Lord Howe Island) Fam. Nicodamidae – Ambicodamus Harvey, 1995 (11 spp., three of them shared with Tasmania); Durodamus Harvey, 1995 (3 spp.); Litodamus Harvey, 1995 (3 spp.); Nicodamus Simon, 1887 (2 spp.); Novodamus Harvey, 1995 (2 spp.); Oncodamus Harvey, 1995 (2 spp.) Fam. Sparassidae – Anchognatha Thorell, 1881 (1 sp.); Holconia Thorell, 1877 (9 spp.); Irileka Hirst, 1998 (1 sp.); Keilira Hirst, 1989 (2 spp.); Neosparassus Hogg, 1903 (19 spp., 2 of them also in Tasmania); Yiinthi Davies, 1994 (8 spp., one of them shared with New Guinea); Zachria L. Koch, 1875 (2 spp.) 762 8 Some Peculiarities of the Distribution of Arachnida Fam. Selenopidae – Karaops Crews et Harvey, 2011 (37 spp.) Fam. Zodariidae – Asteron Jocqué, 1991 (8 spp., 2 also in Tasmania); Cavasteron Baehr et Jocqué, 2000 (12 spp.); Chilumena Jocqué, 1995 (2 spp.); Euasteron Baehr, 2003 (17 spp.); Hetaerica Rainbow, 1916 (2 spp.); Holasteron Baehr, 2004 (16 spp.); Leptasteron Baehr et Jocqué, 2001 (2 spp.); Masasteron Baehr, 2004 (21 spp.); Minasteron Baehr et Jocqué, 2000 (3 spp.); Neostorena Rainbow, 1914 (7 spp.); Nostera Jocqué, 1991 (2 spp., 1 of them shared with Lord Howe Island); Notasteron Baehr, 2005 (2 spp.); Phenasteron Baehr et Jocqué, 2001 (2 spp.); Pseudasteron Jocqué et Baehr, 2001(1 sp.); Spinasteron Baehr, 2003 (19 spp.); Storosa Jocqué, 1991 (2 spp.); Subasteron Baehr et Jocqué, 2001(one sp.); Tropasteron Baehr, 2003 (22 spp.); Zillimata Jocqué, 1995 (one sp.) Fam. Zoropsidae – Austrotengella Raven, 2012 (six spp.) Fam. Miturgidae – Diaprograpta Simon, 1909 (five spp.); Eupograpta Raven, 2009 (two spp.); Mitzoruga Raven, 2009 (three spp.); Nuliodon Raven, 2009 (one sp.) Fam. Ammoxenidae – Austrammo Platnick, 2002 (four spp.), Barrowammo Platnick, 2002 (one sp.) Fam. Gallienelliidae – Meedo Main, 1987 (13 spp.); Neato Platnick, 2002 (7 spp.); Oreo Platnick, 2002 (5 spp.); Peeto Platnick, 2002 (1 sp.); Questo Platnick, 2002 (1 sp.) Fam. Gnaphosidae – Battalus Karsch, 1878 (one sp.); Ceryerda Simon, 1909 (one sp.); Encoptarthria Main, 1954 (five spp.); Epicharitus Rainbow, 1916 (one sp.); Homoeothele Simon, 1908 (one sp.); Montebello Hogg, 1914 (one sp.); Symphanodes Rainbow, 1916 (one sp.) Fam. Lamponidae – Asadipus Simon, 1897 (20 spp.); Bigenditia Platnick, 2000 (2 spp.); Centroina Platnick, 2002 (11 spp.); Centrothele L. Koch, 1873 (10 spp.); Centsymplia Platnick, 2000 (1 sp.); Graycassis Platnick, 2000 (10 spp.); Lamponata Platnick, 2000 (2 spp.); Lamponega Platnick, 2000 (3 spp.); Lamponicta Platnick, 2000 (1 sp.); Lamponina Strand, 1913 (6 spp.); Lamponoides Platnick, 2000 (1 sp.); Lamponusa Platnick, 2000 (1 sp.); Longepi Platnick, 2000 (8 spp.); Notsodipus Platnick, 2000 (18 spp.); Paralampona Platnick, 2000 (8 spp.); Platylampona Platnick, 2004 (1 sp.); Pseudolampona Platnick, 2000 (12 spp.); Queenvic Platnick, 2000 (4 spp.) Fam. Prodidomidae – Cryptoerithus Rainbow, 1915 (19 spp.); Molycria Simon, 1887 (36 spp.); Myandra Simon, 1887 (4 spp., 1 of them shared with Tanzania); Nomindra Platnick et Baehr, 2006 (16 spp.); Wesmaldra Platnick et Baehr, 2006 (14 spp.); Wydundra Platnick et Baehr, 2006 (45 spp. in Australia, 1 in Malaysia, Moluccas) Fam. Trochanteriidae – Bolathana Platnick, 2002 (2 spp.); Desognanops Platnick, 2008 (1 sp.); Desognaphosa Platnick, 2002 (26 spp., 1 on Solomon Islands); Fissarena Henschel, Davies et Dickman, 1995 (9 spp.); Hemicloeina Simon, 1893 (9 spp., 1 in New Guinea); Longrita Platnick, 2002 (10 spp.); Morebilus Platnick, 2002 (13 spp.); Platorish Platnick, 2002 (5 spp.); Rebilus Simon, 1880 (17 spp.); Tinytrema Platnick, 2002 (5 spp., 1 also in Tasmania); Trachycosmus Simon, 1893 (4 spp.); Trachyspina Platnick, 2002 (8 spp.); Trachytrema Simon, 1909 (2 spp.) 8.19 Arachnogeography of Australia and Tasmania 763 Fam. Thomisidae – Corynethrix L. Koch, 1876 (one sp.); Isala L. Koch, 1876 (one sp.); Poecilothomisus Simon, 1895 (one sp.); Saccodomus Rainbow, 1900 (one sp.) Fam. Salticidae – Abracadabrella Zabka, 1991 (3 spp.); Allococalodes Wanless, 1982 (3 spp.); Ananeon Richardson, 2013 (1 sp.); Allococalodes Wanless, 1982 (3 spp.); Aruana Strand, 1911 (2 spp., 1 from Aru Island); Astia L. Koch, 1879 (1 sp.); Astilodes Zabk, 2009 (1 sp.); Barraina Richardson, 2013 (1 sp.); Coccorchestes Thorell, 1881 (39 spp. In New Guinea, 1 sp. in Queensland); Damoetas Peckham et Peckham, 1886 (1 sp.); Frewena Richardson, 2013 (1 sp.); Furculattus Balogh, 1980 (1 sp.); Grayenulla Zabka, 1992 (7 sp.); Huntiglennia Zabka et Gray, 2004 (1 sp.); Jacksonoides Wanless, 1988 (7 sp.); Judalana Rix, 1999 (1 sp.); Leptathamas Balogh, 1980 (1 sp.); Maratus Karsch, 1878 (37 spp. in Australia, 1 sp. in China); Maddisonia Zabka, 2014 (3 sp.); Megaloastia Zabka, 1995 (1 sp.); Paraharnochirus Szombathy, 1915 (2 sp.); Parahelpis Gardzinska et Zabka, 2010 (2 sp.); Paraphilaeus Zabka, 2003 (1 sp.); Porius Thorell, 1892 (2 sp.); Pseudosynagelides Zabka, 1991 (6 sp.); Pungalina Richardson, 2013 (1 sp.); Rhombonotus L. Koch, 1879 (1 sp.); Simaethula Simon, 1902 (7 spp.); Sondra Wanless, 1988 (15 spp.); Tauala Wanless, 1988 (7 spp. in Australia, 1 in Taiwan); Udvardya Prószyński, 1992 (1 sp.); Urogelides Zabka, 2009 (1 sp.); Viroqua Peckham et Peckham, 1901 (1 sp.) Fam. Corinnidae – Leichhardteus Raven et Baehr, 2013 (eight spp.); Poecilipta Simon, 1897 (two spp.) Fam. Liocranidae – Liparochrysis Simon, 1909 (one sp.) Opilioacarida No species has been published, but there are indications for the presence of Opilioacarida in Australia (Walter and Proctor 1998). Acariformes Beron (2008), Domrow (1992), Domrow and Lester (1985), Haliday (1998) Trombidiformes Suborder Prostigmata Fam. Smarididae – Endemic genus Sphaerotarsus Womersley, 1936 Fam. Erythraeidae Endemic genera: Erythrellus Southcott, 1946; Erythrites Southcott, 1946; Erythroides Southcott, 1946; Rainbowia Southcott, 1961; Pussardia Southcott, 1961; Mypongia Southcott, 1961; Wartookia Southcott, 1961 Sarcoptiformes Suborder Oribatida According to the Catalogue of Australian Oribatida (Colloff and Halliday, 1998), in Australia are registered 340 named species (256 endemic) and 237 unnamed species or a total of 577 species. There are no endemic superfamilies indigenous to Australia. 764 8 Some Peculiarities of the Distribution of Arachnida Parasitiformes (Ixodida and Mesostigmata) Order Ixodida (Domrow 1987, Haliday 1998, Roberts 1970) Fam. Argasidae – Six spp. of Argas Latreille and one of Otobius Banks Fam. Ixodidae – Sixty-eight spp., no endemic genera, some endemic species Holothyrida Known are two species of the genus Allothyrus (fam. Allothiridae, confined to Australia and New Zealand) (Domrow, 1955; Van der Hammen, 1961, 1983; Womersley, 1935). 8.20 8.20.1 Tasmania Geography, General Zoogeography, and Paleogeography Tasmania is an Australian island and state. It is 240 km south of the continent, separated by Bass Strait. The state includes the island of Tasmania and some smaller islands (state area 68,401 km2), of which the main island covers 62,409 km2. The highest point is Mount Ossa (1614 m.). Situated at 42oS, Tasmania has a cool temperate climate with four distinct seasons. The island was joined to the mainland of Australia until the end of the last glacial period approximately 10,000 years ago. Much of Tasmania is still densely forested, with the Southwest National Park and neighboring areas holding some of the last temperate rain forests in the Southern Hemisphere. The Tarkine, located in the island’s far North West, is the largest temperate rainforest area in Australia covering approximately 3800 square kilometers. According to Darlington (1960), the fauna of Tasmania is depauperate, both in diversity of basic stocks and in total number of species. Many groups, both vertebrate (Varanus monitor lizards, pythons, Typhlopidae, and many others) and invertebrates (Uropygi, Amblypygi, Cyphophthalmi, Dyspnoi), are missing in the island. Still, in Tasmania are (or were) preserved animals extinct in mainland Australia. “Tasmania has long been regarded as a biological treasure trove, supporting an astounding range of species found nowhere else, many of which have survived the perturbations of climate change during the Pleistocene” (Harvey 1998). Arachnogeography No records for the order Palpigradi, Ricinulei, Solifugae, Amblypygi, Uropygi, Schizomida, Opilioacarida, and Holothyrida and the suborders Paleamblypygi, Cyphophthalmi, and Dyspnoi. Almost no scorpions, many endemic genera of Triaenonychidae (Laniatores), 2 endemic subfamilies, and 29 endemic genera of spiders, many endemic genera of Acari (Erythraeidae and others). Scorpiones From Tasmania has been recorded only Cercophonius squama (Bothriuridae), shared with mainland Australia. Also on Flinders I. and King I. (Koch, 1977). 8.20 Tasmania 765 Pseudoscorpiones In Tasmania are registered Pseudoscorpiones from several families (Chthoniidae, Pseudogarypidae, Pseudotyrannochthoniidae, Garypidae, Syarinidae, Garypinidae, etc.) (Benedict and Malcolm 1978; Dartnall 1970; Harvey 1998; Morris 1948). Endemic genera for Tasmania are Neopseudogarypus Morris, 1948 (Pseudogarypidae), and Oreolpium Harvey et Štáhlavský (Garypinidae). Opiliones. Ref.: Hickman (1957, 1958), Hogg (1909), Hunt (1990, 1995), Roewer (1914) In Tasmania have been registered the following Opiliones: Cyphophthalmi – Not recorded in Tasmania Eupnoi – Fam. Neopilionidae (incl. Monoscutidae) (Spinicrus Forster, 1949) Dyspnoi – Fam. Acropsopilionidae: Austropsopilio Forster, 1955 (Tasmanopilio Hickman, 1957) In Tasmania are registered Laniatores of 27 endemic genera of the family Triaenonychidae. Endemic genera are: Fam. Triaenonychidae Allonuncia Hickman, 1958 (one sp.); Ankylonuncia Hickman, 1958 (three spp.); Bryonuncia Hickman, 1958 (one sp.); Calliuncus Roewer, 1931 (five spp.); Cluniella Forster, 1955 (three spp.); Hickmanoxyomma Hunt, 1990 (seven spp.); Leionuncia Hickman, 1958 (one sp.); Notonuncia Hickman, 1958 (three spp.); Nucina Hickman, 1958 (two spp.); Nuncioides Hickman, 1958 (two spp.); Odontonuncia Hickman, 1958 (one sp.); Parattahia Roewer, 1914 (one sp.); Rhynchobunus Hickman, 1958 (one sp.); Stylonuncia Hickman, 1958 (one sp.); Tasmanonyx Hickman, 1958 (one sp.); Pyenganella Hickman, 1958 (one sp.); Tasmanonuncia Hickman, 1958 (one sp.); Thelbunus Hickman, 1958 (one sp.); Allobunus Hickman, 1958 (one sp.); Chilobunus Hickman, 1958 (one sp.); Chrestobunus Roewer, 1914 (three spp.); Eubunus Hickman, 1958 (one sp.); Glyptobunus Roewer, 1914 (two spp.); Mestonia Hickman, 1958 (one sp.); Miobunus Roewer, 1915 (six spp., Tasmania); Phanerobunus Roewer, 1915 (four spp.); Phoxobunus Hickman, 1958 (two spp.) Araneae In Tasmania have been recorded 2 endemic subfamilies (Plesiothelinae and Hickmanniinae) and 29 endemic genera of spiders (Musgrave 1947; Gertsch 1958; Hickman 1957, 1958, 1969; Platnick and Forster 1989; Raven 1978). Endemic genera and subfamilies: Fam. Hexathelidae Plesiothelinae Plesiothele Raven, 1978 – One sp. 766 8 Some Peculiarities of the Distribution of Arachnida Fam. Anapidae Acrobleps Hickman, 1979 (one sp.); Epigastrina Rix et Harvey, 2010 (three spp.); Hickmanapis Platnick et Forster, 1989 (two spp.); Olgania Hickman, 1979 (five spp.); Tasmanapis Platnick et Forster, 1989 (one sp.) Fam. Austrochilidae – Chile, Argentina, Tasmania (nine spp.) Hickmaniinae (as family Hickmaniidae) Hickmania Gertsch, 1958 – One sp.: Hickmania troglodytes (Higgins et Petterd, 1883), Cave Fam. Orsolobidae Cornifalx Hickman, 1979 (one sp.); Hickmanolobus Forster et Platnick, 1985; Olgania Hickman, 1979; Carathea Moran, 1986; Tupua Platnick in Forster, Platnick et Coddington, 1990; Cicirra Simon, 1876; Namandia Lehtinen, 1967; Ommatauxesis Simon, 1903; Toxops Hickman, 1940 Fam. Malkaridae – Carathea Moran, 1986 (two spp.) Fam. Anapidae (subfam. Micropholcommatinae) – Epigastrina Rix et Harvey, 2010 (three spp.) Fam. Synotaxidae – Tupua Platnick, 1990 (four spp.) Fam. Lycosidae – Tasmanicosa Roewer, 1959 (one sp.) Fam. Psechridae – Tjurunga Lehtinen, 1967 (one sp.) Fam. Miturgidae (Zorinae) – Odomasta Simon, 1909 (one sp.) Fam. Araneidae – Collina Urquhart, 1891 (one sp.) Fam. Amphinectidae Tanganoides Davies, 2005 (six spp., incl. one in Victoria); Tasmabrochus Davies, 2002 (five spp.); Tasmarubrius Davies, 1998 (five spp.); Teeatta Davies, 2005 (three spp.) Fam. Amaurobiidae – Only endemic species Fam. Desidae Cicirra Simon, 1886 (one sp.); Namandia Lehtinen, 1967 (one sp.); Ommatauxesis Simon, 1903 (one sp.); Toxops Hickman, 1940 (one sp.) Fam. Hahniidae – Neoaviola Butler, 1929 (one sp.) Fam. Stiphidiidae – Tjurunga Lehtinen, 1967 (one sp.) Notes on several families: Actinopodidae Three genera, two in South America and one (Missulena Walckenaer, 1805) sheared between Australia (10 spp.) and Chile (Missulena tussulena Goloboff, 1994). Greater Nothogea? Austrochilidae Small family of nine spp., illustrating well the concept of Notogea including the southern end of South America. Two subfamilies: Austrochilinae with 8.21 Parasitiformes (Some Ixodida and Mesostigmata): Australia and New Zealand 767 two genera from Chile and Argentina and Hickmaniinae with one species from a cave in Tasmania. The same could be said concerning the fam. Malkaridae: one genus in Tasmania, two in mainland Australia, and one (Chilenodes Platnick et Forster, 1987) in Chile and Argentina. Hexathelidae Atracinae (three genera with three spp., endemic to Australia), Plesiothelinae is endemic to Tasmania and many others in Europe, Asia, and Africa, Macrothelinae has an isolated end. genus in New Zealand (Porrhothele Simon, 1892), and Hexathelinae is also a good example for a bigger Notogea – three genera in Australia, two in Chile and Argentina, and one in New Zealand. Nicodamidae Nine genera with 29 spp. Four genera endemic to Australian continent, one end. to Tasmania, one (Dimidamus Harvey, 1995) distributed in New Guinea and Australia, and one (Ambicodamus Harvey, 1995) shared between Australia and Tasmania. Malkaridae Four genera, two (Malkara and Perissopmeros) endemic to mainland Australia, one (Carathea) endemic to Tasmania, and one (Chilenodes Platnick and Forster, 1987) known from Chile and Argentina. Again “Greater Nothogea”! To this family was attached as junior synonym Pararchaeidae from Australia, Tasmania, and New Zealand. Subfam. Micropholcommatinae The distribution of the subfamily of Anapidae comprises mainland Australia, Tasmania (22 spp.), Auckland Islands, New Guinea, Campbell Islands, New Zealand, New Caledonia, Lord Howe Island, Chile, and Brazil. Thanks to the work of Rix and Harvey (2010), now we have several examples of “traditional” Gondwanan distribution: Gigiella Rix et Harvey, 2010 (one sp. in mainland Australia and Tasmania, one sp. in Chile); Eperiella Rix et Harvey, 2010 (one sp. in Tasmania, one sp. in Chile); and Normplatnicka Rix et Harvey, 2010 (two spp. in m. Australia, one sp. in Chile). They are also good examples for a bigger Notogea. Synotaxidae Three subfamilies: Pahorinae (endemic to New Zealand), Physogleninae (Australia, Tasmania, New Zealand, and Chile), and Synotaxinae (Australia, Chile, South America). 8.21 Parasitiformes (Some Ixodida and Mesostigmata): Australia and New Zealand Ixodida New Zealand has 11 named species of ticks, four of which are endemic: Aponomma sphenodonti (Dumbleton, 1953) (on tuatara); Ixodes anatis Chilton, 1904 (on kiwi and Anatidae); I. jacksoni Hoogstraal, 1967 (from nest of Stictocarbo- 768 8 Some Peculiarities of the Distribution of Arachnida Phalacrocorax punctatus); and Carios quadridentatus Heath, 2012 (from the endemic bat Mystacina tuberculata). Six are known also from Australia (Dumbleton 1953, 1963; Heath 1977, 2012; Heath et al. 2011; Hoogstraal 1967; Spain and Luxton 1971). Acariformes (Some Examples) Sarcoptiformes Suborder Oribatida Ref.: Australia. Colloff and Haliday (1998); New Zealand. Hammer (1966, 1967, 1968), Colloff and Cameron (2014). According to the catalogue of Australian Oribatida (Colloff and Halliday 1998), in Australia are registered 340 named species (256 endemic) and 237 unnamed species or a total of 577 species. There are no superfamilies indigenous to Australia. Thanks to the research of M. Hammer, from New Zealand have been listed at least 50 families of Oribatida (Spain and Luxton 1971). Trombidiformes (Some examples) Ref.: Beron (2008a), Domrow (1978), Domrow and Lester (1985), Haliday (1998), Southcott (1961), Womersley (1936), Southcott (1966), Hirst (1926), Southcott (1988), Zhang (2000) Prostigmata Erythraeoidea Endemic Genera in Australia Fam. Smarididae – Sphaerotarsus Womersley, 1936 Fam. Erythraeidae Erythrellus Southcott, 1946; Erythrites Southcott, 1946; Erythroides Southcott, 1946; Rainbowia Southcott, 1961; Pussardia Southcott, 1961; Mypongia Southcott, 1961; Wartookia Southcott, 1961 End. species in New Caledonia: Charletonia rageaui Southcott, 1966 Endemic genera in New Zealand: Neosmaris Hirst, 1926; Taranakia Southcott, 1988; Ramsayella Zhang, 2000 8.22 8.22.1 New Zealand Geography, General Zoogeography, and Paleogeography The archipelago, called by the Europeans in 1645 New Zealand, is one of the amazing places on Earth, when discussing zoogeography. It is composed of two bigger islands (North and South Islands), separated by the Cook Straight (22 km. wide), one less big (Stewart), and many smaller islands with a total surface of 268 thousand km2. New Zealand is 1500 km. far from Australia. The relief is very varied with highest point Mount Cook, or Aoraki (3754 m.), and 17 other summits higher than 3000 m. The biota is very altered by humans, mostly by the white settlers and the 8.22 New Zealand 769 animals they introduced, one way or another. For the invertebrates, most important was the change in the environment because of the land use, deforestation, etc. To understand the history of the biota of New Caledonia, New Zealand, and Lord Howe Island, we have to follow up the hypothetical development of Zealandia, the New Zealand microcontinent, having existed until some 23 million years ago as a result of the breakaway from Australia about 60–85 million years ago and from Antarctic 85–130 Ma ago (Craw 1988, 1989; Fleming 1949, 1962, 1963a, 1963b, 1967, 1970, 1975; Grehan 1989; Griffiths 1975; Page 1989; Sharma and Wheeler 2013; Udvardy 1975). According to Fleming (1975), the isolation of New Zealand becomes effective for the land vertebrates (and maybe invertebrates) before the end of Cretaceous. Cracraft (1973) thinks that “Australia separated from Antarctica in the Eocene and with spreading rates approximately twice as fast as those for New Zealand and this differential movement between Australia-New Zealand and Antarctica formed the Tasman Sea” (see also Griffiths and Varne, 1972). In any case, New Zealand was detached from Antarctida more than 80 Ma ago (Cracraft, 1973), together with the ancestors not only of the moa, kiwi, tuatara, and other well-known cases also with the ancestors of many insects and other invertebrates. In Cracraft (1974), we read that “The next portion of Gondwanaland to separate was New Zealand, which was adjacent to West Antarctica and which began drifting in the late Cretaceous at about 40–45 Ma ago.” Arachnogeography It would be hardly possible to assess the entire endemism and richness of New Zealand arachnofauna some 40 years ago. Forster (1949c, 1973, 1975) analyzed the particularities of this amazing fauna, but since (mostly with the efforts of Forster himself and his collaborators), new data have been accumulated. The orders Scorpiones, Ricinulei, Thelyphonida (Uropygi), Amblypygi, Schizomida, and Opilioacarida are completely missing from New Zealand and the surrounding islands (naturally). Represented are Pseudoscorpiones, with six endemic genera, Opiliones (amazingly rich in endemics and with elements in common with faraway countries, one endemic family), and Araneae (huge number of spiders in an island country). Among spiders there are 93 endemic genera and 93% endemic species. Pseudoscorpiones The pseudoscorpion fauna of New Zealand and the Subantarctic islands in the area consists now of 70 species of 28 genera and 9 families: Chthoniidae, Syarinidae, Garypidae, Garypinidae, Olpiidae, Cheiridiidae, Cheliferidae, Chernetidae, Withiidae, including some species and 6 genera endemic. One (Philomaoria) is shared with Lord Howe Island, another (Synsphyronus) with Tasmania, and 11 with Australia (Beier 1966, 1967, 1969, 1973, 1976; Chamberlin 1925; Harvey 2009, 2013f). Endemic genera: Fam. Chthoniidae Maorichthonius Chamberlin, 1925 – New Zealand (one sp.) Sathrochthoniella Beier, 1967 – New Zealand (one sp.) 770 8 Some Peculiarities of the Distribution of Arachnida Tyrannochthoniella Beier, 1966 – New Zealand (one sp.) Fam. Olpiidae Nelsoninus Beier, 1967 – New Zealand (South Island) (one sp.) Fam. Chernetidae Apatochernes Beier, 1948 – New Zealand (Campbell Islands, Snares Islands, Auckland Islands, Chatham Islands, Norfolk Island (17 spp.) Heterochernes Beier, 1966 – New Zealand (one sp.) Opiliones Ref.: Boyer and Giribet (2007, 2009), Crawford (1992), Forster (1947, 1948, 1952, 1954, 1975, 16 papers from 1943 to 1975), Groh and Giribet (2015), Hirst (1925), Roewer (1931), Taylor (2004, 2009, 2011, 2013). In New Zealand (s. str.) live 27 genera of Opiliones, out of them 24 endemic (Forster, 1975, actualized), from the families Pettalidae, Acropsopilionidae, Neopilionidae (incl. Monoscutidae), Synthetonychiidae, and Triaenonychidae. Cyphophthalmi In New Zealand have been recorded 23 spp. of the fam. Pettalidae: Rakaia Hirst, 1925 – Twelve spp. (endemic genus) Aoraki Boyer et Giribet, 2007 – Eight spp. (endemic genus) Neopurcellia Forster, 1948 – South Island (three spp.) (endemic genus) Eupnoi Fam. Neopilionidae – Megalopsalidinae: Megalopsalis Forster, Forsteropsalis Taylor (nine spp.), Mangatangi Taylor (one sp.), Pantopsalis Simon (nine sp.) Monoscutinae (endemic subfamily): Acihasta Forster, Monoscutum Forster, Templar Taylor (New Zealand, Auckland, Snares, Campbell Islands) Dyspnoi: Fam. Acropsopilionidae – Acropsopilio Silvestri (one sp.) Laniatores In New Zealand are registered 68 spp. of Laniatores of 15 genera and 2 families. Endemic supraspecific taxa: Fam. Synthetonychiidae – Endemic family Synthetonychia Forster, 1954 – Fourteen spp. Fam. Triaenonychidae Hedwiga Roewer, 1931 (1 sp.); Hendea Roewer, 1931 (14 spp.); Hendeola Forster, 1954 (2 spp.); Metanuncia Roewer, 1914 (Stewart Island) (1 sp.); Neonuncia Roewer, 1914 (5 spp.); Prasma Roewer, 1931 (3 spp.); Prasmiola Forster, 1954 (one sp.); Psalenoba Roewer, 1931 (one sp.); Triregia Forster, 1948 (3 spp.); Algidia Hogg, 1920 (8 spp.); Cenefia Roewer, 1931 (4 spp.); Muscicola Forster, 1954 (1 sp.); Pristobunus Roewer, 1931 (2 spp.) Subfam. Soerensenellinae – Endemic subfamily (eight spp.) 8.22 New Zealand 771 Karamea Forster, 1954 (four spp.); Soerensenella Pocock, 1903 – four species Araneae Recently an overview of New Zealand spiders has been published by Paquin, Vink, and Dupérré (2010). Best characterized is the spider fauna of New Zealand by the book review of Duffey (2010, Newsl. Br. Arachnol. Soc., 119): “Britain and New Zealand are comparable in area but the former has 658 species while the latter has 1126 described and another 536 awaiting description, making a present total of about 1662 species in 236 genera and 57 families. As new species are still being found the authors think the true total could be about 2000. One can’t help feeling this is a conservative estimate because even the well-worked fauna in Britain, which recorded a total of 584 in 1958, has since added another 74 species. The best known New Zealand arachnologist, R.R. Forster, though the New Zealand total could be as much as 2500 species. Whether 2000 or 2500, the fascinating question is why New Zealand has such a rich fauna when no country in Europe reaches even the lower figure. France, one of the largest, is two and half times the area [of NZ], but has a total of 1569 sp. (Le Peru 2007). The family Linyphiidae is the largest in New Zealand but only 12% of the total and they are all in the subfamily Linyphiinae. Endemic Erigoninae apparently do not exist as all known species are introduced. In Britain about 40% of the spider fauna are Linyphiidae, of which over 70% are Erigoninae.” The endemism of New Zealand spiders is amazing: 93% of the fauna consists of endemic species. From New Zealand (including the nearby islands Auckland, Campbell, Snares, etc.) have been recorded 2 endemic families (Huttoniidae and Megadictynidae), 1 subfamily (Pahorinae), and 93 endemic genera, and at least 93% of the species of spiders are endemic (Simon 1892; Blest 1979; Forster, 1955, 1959, 1964, 1970, 1975; Forster and Blest 1979; Forster and Platnick 1977, 1984; Forster and Wilton 1968, 1973; Parrott 1942; Platnick and Forster 1989; Rix 2006; Vink and Dupérré 2010; Zabka et al. 2002): Fam. Hexathelidae Hexathele Ausserer, 1871 (20 spp.); Porrhothele Simon, 1892 (5 spp.) Fam. Anapidae Novanapis Platnick et Forster, 1989 (one sp.); Paranapis Platnick et Forster, 1989 (two spp.); Zealanapis Platnick et Forster, 1989 (ten sp.) Fam. Pararchaeidae – Forstrarchaea Rix, 2006 (one sp.); Pararchaea Forster, 1955 (one sp.) Fam. Idiopidae – Cantuaria Hogg, 1902 (42 spp.) Fam. Gradungulidae Gradungula Forster, 1955; Pianoa Forster, 1987; Spelungula Forster, 1987 (one sp.) Fam. Huttoniidae: Huttonia Pickard-Cambridge, 1880 772 8 Some Peculiarities of the Distribution of Arachnida Fam. Orsolobidae: Anopsolobus Forster et Platnick, 1985; Ascuta Forster, 1956; Bealeyia Forster et Platnick, 1985; Dugdalea Forster et Platnick, 1985; Duripelta Forster, 1956; Maoriata Forster et Platnick, 1985; Orongia Forster et Platnick, 1985; Paralobus Forster et Platnick, 1985; Pounamuella Forster et Platnick, 1985; Subantarctia Forster, 1955; Tangata Forster et Platnick, 1985; Tautukua Forster et Platnick, 1985; Turretia Forster et Platnick, 1985; Waiporia Forster et Platnick, 1985; Waipoua Forster et Platnick, 1985; Wiltonia Forster et Platnick, 1985 Fam. Mecysmauchenidae (Aotearoa Forster et Platnick, 1974; Zearchaea Wilton, 1946 Fam. Anapidae (Micropholcommatinae): Pua Forster, 1959; Tinytrella Rix et Harvey, 2010; Taliniella Rix et Harvey, 2010 Fam. Cyatholippidae: Hanea Forster, 1988; Tekella Urquhart, 1894; Tekelloides Forster, 1988 Fam. Synotaxidae: Nomaua Forster, 1990; Pahora Forster, 1990; Pahoroides Forster, 1990; Runga Forster, 1990; Meringa Forster, 1990; Mangua Forster, 1990; Zeatupua Fitzgerald et Sirvid, 2009 Fam. Agelenidae: Ahua Forster et Wilton, 1973; Huka Forster et Wilton, 1973; Mahura Forster et Wilton, 1973; Neoramia Forster et Wilton, 1973; Neorepukia Forster et Wilton, 1973; Oramia Forster et Wilton, 1973 (also in Lord Howe Island); Oramiella Forster et Wilton, 1973; Orepukia Forster et Wilton, 1973; Paramyro Forster et Wilton, 1973; Porotaka Forster et Wilton, 1973; Tararua Forster et Wilton, 1973; Tuapoka Forster et Wilton, 1973 Fam. Tetragnathidae: Eryciniolia Strand, 1912 (one sp.); Nanometa Simon, 1908 (one sp.); Pinkfloydia Dimitrov et Hormiga, 2011 (one sp.) Fam. Lycosidae: Notocosa Vink, 2002 (one sp.) Fam. Psechridae – [Borrala Gray et Smith, 2004 (31 spp., 1 in New Caledonia)]; Nanocambridgea Forster et Wilton, 1973 (1 sp.); Pillara Gray et Smith, 2004 (4 spp.) Fam Amaurobiidae – Auhunga Forster et Wilton, 1973 (1 sp.); Maloides Forster et Wilton, 1989 (1 sp.); Muritaia Forster et Wilton, 1973 (5 spp.); Otira Forster et Wilton, 1973 (6 spp.); Pakeha Forster et Wilton, 1973 (18 spp.); Paravoca Forster et Wilton, 1973 (2 spp.); Poaka Forster et Wilton, 1973 (1 sp.); Waitetola Forster et Wilton, 1973 (1 sp.) Fam. Amphinectidae: Akatorea Forster et Wilton, 1973 (2 spp.); Amphinecta Simon, 1898 (11 sp.); Aorangia Forster et Wilton, 1973 (16 sp.); Dunstanoides Forster et Wilton, 1989 (9 spp.); Holomamoea Forster et Wilton, 1973 (1 sp.); Huara Forster, 1964 (12 spp.); Makora Forster et Wilton, 1973 (5 spp.); Mamoea Forster et Wilton, 1973 (19 spp.); Maniho Marples, 1959 (10 spp.); Marplesia Lehtinen, 1967 (2 spp.); Neolana Forster et Wilton, 1973 (3 spp.); Neororea Forster et Wilton, 1973 (2 spp.); Oparara Forster et Wilton, 1973 (2 spp.); Paramamoea Forster et Wilton, 1973 (10 spp.); Rangitata Forster et Wilton, 1973 (1 sp.); Reinga Forster et Wilton, 1973 (5 spp.); Rorea Forster et Wilton, 1973 (2 spp.); Waterea Forster et Wilton, 1973 (1 sp.) 8.22 New Zealand 773 Fam. Desidae: Gasparia Marples, 1956; Gohia Dalmas, 1917; Goyenia Forster et Wilton, 1970; Hapona Forster, 1970; Helsonia Forster, 1970; Hulua Forster et Wilton, 1973; Lamina Forster, 1970; Mangareia Forster, 1970; Matachia Dalmas, 1917; Mesudus Özdikmen, 2007; Neomyro Forster et Wilton, 1973; Notomatachia Forster, 1970; Nuisiana Forster et Wilton, 1973; Otagoa Forster, 1970; Panoa Forster, 1970; Rapua Forster, 1970; Toxopsoides Forster et Wilton, 1973; Tuakana Forster, 1970 Fam. Dictynidae: Arangina Lehtinen, 1967; Paradictyna Forster, 1970; Viridictyna Forster, 1970 Fam. Hahniidae: Kapanga Forster, 1970; Porioides Forster, 1989; Rinawa Forster, 1970 Fam. Megadictynidae: Forstertyna Harvey, 1995; Megadictyna Dahl, 1906 Fam. Zodariidae: Forsterella Jocqué, 1991 Fam. Zoropsidae: Haurokoa Forster et Wilson, 1973 Fam. Cycloctenidae: Plectophanes Bryant, 1935 (5 spp.); Toxopsiella Forster, 1964 (12 spp.); Uzakia Koçak et Kemal, 2008 (1 sp.) Fam. Miturgidae: Pacificana Hogg, 1904; Zealoctenus Forster et Wilton, 1973 Fam. Gnaphosidae: Kaitawa Forster, 1979; Matua Forster, 1979; Nauhea Forster, 1979; Notiodrassus Bruyant, 1935; Zelanda Özdikmen, 2009 Fam. Thomisidae: Cymbachina Bryant, 1933 (one sp.) Fam. Salticidae: Hinewaia Zabka et Pollard, 2002 (one sp.) Acariformes Ref.: Beron (2008a), Clark (2014), Hammer (1966, 1968), Spain and Luxton (1971) Trombidiformes Prostigmata Fam. Erythraeidae Endemic genera: Neosmaris Hirst, 1926; Ramsayella Zhang, 2000; Taranakia Southcott, 1988; Pukakia Clark, 2014 Parasitiformes Ixodida New Zealand has nine named species of ticks, three of which are endemic: Aponomma sphenodonti (Dumbleton, 1953) (on tuatara); Ixodes anatis Chilton, 1904 (on kiwi and Anatidae); and I. jacksoni Hoogstraal, 1967 (from nest of Stictocarbo-Phalacrocorax punctatus). Six are known also from Australia (Dumbleton 1953, 1963; Heath 1977; Hoogstraal 1967; Spain and Luxton 1971). One unnamed Argasidae is known from the endemic bat Mystacina tuberculata. Holothyrida Recorded (loc.?) is Allothyrus (?) australasiae (Womersley, 1935) known from Australia. 774 8.23 8.23.1 8 Some Peculiarities of the Distribution of Arachnida New Caledonia Geography, General Zoogeography, and Paleogeography New Caledonia is located in the subregion of Melanesia in the South West Pacific. It comprises the main island (Grande Terre), the Loyalty Islands, and several smaller islands. It has a land area of 18,575.5 square kilometers. The Grande Terre is by far the largest of the islands, with an area of 16,372 square kilometers, and is elongated northwest-southeast, 350 km in length and 50–70 km wide. A mountain range runs the length of the island, with five peaks over 1500 m. The highest point is Mont Panié at 1628 m elevation. New Caledonia is one of the northernmost parts of a (93%) submerged continent called Zealandia. It sank after rifting away from Australia 60–85 million years ago (mya) and from Antarctica between 130 and 85 mya. New Caledonia itself is separated from Australia in the Late Cretaceous (65–66 Ma ago), and subsequently drifted in a north-eastern direction, reaching its present position about 50 Ma ago. According to Neall and Trewick (2008), New Caledonia separated from New Zealand 83 Ma ago. There is also a hypothesis that the islands emerged 37 Ma ago (Oligocene) and that its biota started developing by this time. Some geologists insist that New Caledonia has been entirely submerged several times and repopulated after that. However, biologists accept that parts of the land remained above water as refugia for the archaic animals and plants. Many species from the Gondwanan flora in the Late Cretaceous and early Tertiary probably survived in the temperate climate of New Caledonia and died out in Australia during its strong aridisation. This opinion is contested too. The detailed analysis of Grandcolas et al. (2008) concerning the formation of the New Caledonian biota comes to the conclusion that it is not a continental, but rather an oceanic island, having its biota formed not earlier than 37 Ma ago during the Oligocene (confirmed by molecular research). New Caledonia is extremely interesting from biogeographical point of view, and different problems are raised. As a matter of fact, it is an archipelago of one big and very varied island, the Loyalty Islands, the Pine Island, and many other islets and reefs. The polulation is 258 000. The isolation of New Caledonia dates at least from the Miocene (perhaps from the Oligocene) and thus have been preserved many relict animal and plant, and also neoendemics have been formed. After the arrival of the first settlers (the Canaques) ca. 3200–3300 years ago, many endemic animals disappeared, and the Europeans (since 1853 New Caledonia is a French territory) contributed to it mostly by mining and agrarian activities, forest destruction, planting foreign trees, and others. New Caledonia is 1300 km. far from Australia, 1500 km. from New Zealand, and 1200 km. from Fiji (Darlington 1957; Heads 2008a, 2008b, 2010; Lillie and Brothers 1970; Neall and Trewick 2008; Paramonov 1958, 1960; Sarasin 1925). Arachnogeography There are several hypotheses about the origin of the arachnofauna (and the remaining fauna) of the Big Island, far away from any mainland. Besides the animals, brought on the “Three Ws” (wind, waves, and wings), the fauna is either quite recent (after the total submersion of the island) or contains 8.23 New Caledonia 775 remains from much older fauna (Grandcolas et al., 2008). The presence, at least, of the amazing relict family of (cave dwelling) opilions Troglosironidae speaks rather for at least some elements surviving from older fauna. Besides, on the islands, there are some other orders of Arachnida: Scorpiones, Amblypygi, Schizomida, and Pseudoscorpiones, endemic genera of Laniatores, many endemic spiders, and most interesting Holothyrida – one genus endemic and one shared with Lord Howe Island. Scorpiones Three species (two endemic and one widespread) are recorded (Kraepelin 1914; Simon 1877; Vachon 1976). Fam. Buthidae Isometrus (Reddyanus) heimi Vachon Fam. Hormuridae Liocheles australasiae (Fabricius) L. neocaledonicus Simon Amblypygi Charinus neocaledonicus Simon, 1895 (Charinidae), endemic species of a widespread genus, is recorded from New Caledonia. Other members of the same genus (all of them known from caves) are three subspecies of Ch. australianus and Charinus pecki Weigoldt, 2006. Schizomida Only Hubbardiidae indet. are mentioned from New Caledonia (Reddell and Cokendolpher 1995). Pseudoscorpiones In New Caledonia and the Loyalty Islands are known 14 species of Pseudoscorpiones from 10 genera of 7 families: Chthoniidae, Tridenchthoniidae, Parahyidae, Syarinidae, Garypinidae, Atemnidae, and Chernetidae (Beier (1940, 1964, 1966, 1966d, 1968, 1976, 1979; Simon 1880). Among them are the endemic species Hebridochernes caledonicus, H. gressitti, H. maximus, Nesidiochernes caledonicus, Paraldabrinus (end. genus) novaecaledoniae, Amblyolpium ruficeps, Ideobisium antipodum, Anaulacodithella novacaledonica, A. reticulata, Sathrochthonius kaltenbachi, Tyrannochthonius troglophilus, and T. zonatus – 12 spp. or 86% endemism. Opiliones The most interesting is the endemic family of Cyphophthalmi Troglosironidae with 1 genus Troglosiro Juberthie, 1979, and 13 spp. (Shear 1993; Sharma and Giribet 2009). Dyspnoi - Missing Laniatores From New Caledonia and the Loyalty Islands are known Laniatores from the families Triaenonychidae and Zalmoxidae (eight endemic species of genus 776 8 Some Peculiarities of the Distribution of Arachnida Zalmoxis) (Simon 1881; Roewer 1912, 1914, 1949; Goodnight and Goodnight 1948; Sharma and Giribet 2012). Endemic Laniatores in New Caledonia are the following genera: Fam. Triaenonychidae Diaenobunus Roewer, 1914; Triconobunus Roewer, 1914 Santobius Roewer, 1949 (Mesoceras Soerensen, 1886; preoccup., Mesoceratula Roewer, 1949) – Vanuatu, Fiji The presence of fam. Zalmoxidae has been explained by Sharma and Giribet (2012): “The family Zalmoxidae, similar to the Pacific iguanas, constitutes the unusual case of a lineage of Neotropical origin that colonized the Indo-Pacific likely by ancient transoceanic dispersal during the Late Cretaceous.” Araneae In New Caledonia have been recorded spiders of al least 22 families: Anapidae, Barychelidae, Dipluridae, Desidae, Pararchaeidae, Lamponidae, Theridiidae, Pisauridae, Miturgidae (Zorinae), Salticidae, Scytodidae, Tetrablemmidae, Telemidae, Segestriidae, Oonopidae, Mimetidae, Deinopidae, Uloboridae, Mysmenidae, Linyphiidae, Tetragnathidae, and Araneidae (Berland 1924, 1929; Platnick and Forster 1989, 1993; Platnick 1993; Rainbow 1920; Raven 1994; Rix and Harvey 2010; Simon 1889; 1892). Until 1993 have been recorded spiders of 112 genera and 194 spp. (Platnick 1993). Platnick and Forster (1993) synonymized the “endemic family” Bradystichidae Simon, 1884, with Pisauridae. Rix and Harvey (2010b) recorded from the island the first representative of Pararchaeidae outside Australia and New Zealand. Endemic genera: Fam. Dipluridae: Stenygrocercus Simon, 1892 (six spp.) Fam. Anapidae: Caledanapis Platnick et Forster, 1989 (six spp.); Mandanapis Platnick et Forster, 1989 (one sp.); Montanapis Platnick et Forster, 1989 (one sp.); Caledothele Raven, 1991 Fam. Barychelidae, Barychelinae: Barycheloides Raven, 1994 (5 spp.); Barychelus Simon, 1889 (2 spp.); Encyocrypta Simon, 1889 (32 spp.); Natgeogia Raven, 1994 (1 sp.); Orstom Raven, 1994 (6 spp.); Questocrypta Raven, 1994 (1 sp.) Fam. Desidae: Canala Gray, 1992 (three spp.) Fam. Lamponidae: Centrocalia Platnick, 2000 (three spp.) Fam. Theridiidae: Anatea Berland, 1927 (one sp.) Fam. Pisauridae: Bradystichus Simon, 1884 (five sp.), Pseudohostus Rainbow, 1915 (five spp.) Fam. Miturgidae (incl. Zoridae): Zoroides Berland, 1924 (one sp.) Fam. Salticidae: Corambis Simon, 1901 (two spp.); Lystrocteisa Simon, 1884 (one sp.); Rhondes Simon, 1901 (six spp.) Acariformes - Ref.: Beron (2008) Trombidiformes – Prostigmata Fam. Erythraeidae: Charletonia rageaui Southcott, 1966 8.24 Lord Howe Island 777 Parasitiformes (Ixodida and Mesostigmata) Holothyrida First recorded by Berlese (1923). Endemic genus is Haplothyrus Lehtinen (two spp.). One genus (Lindothyrus Lehtinen) is shared with Lord Howe Island (on New Caledonia endemic sp. Lindothyrus rubellus Lehtinen, 1995). Both genera belong to the family Holothyridae and not to the Allothyridae, known from Australia and New Zealand. The New Guinean genera also are not represented in New Caledonia. The same observation is valid for Lord Howe Island. 8.24 Lord Howe Island The small Lord Howe Island is called “The riddle of Pacific” (Paramonov 1958). On a surface of 14.55 km2 (10 km long and up to 2 km wide) live many endemic species, and the island is 600 km. far from Australia and 900 km. far from Norfolk Island. Its inhabitants (387 permanent and up to 400 tourists) are in the “settled area,” in the lowland. It is because since 1981 70% of the island is a reserve and, fortunately, there are some forests left on the 875 meter high Mount Gower. The archipelago includes also 28 uninhabited islands. It is considered that Lord Howe is part of the island chain, having existed along the Western edge of the shelf called Lord Howe Rise, 3000 km. long and 300 wide. This Rise extended from New Zealand to west of New Caledonia and consists of continental rocks, detached from Australian plate 60–80 Ma ago. The shelf is part of Zealandia microcontinent. Pseudoscorpiones From Lord Howe Island, Beier (1976) has published eight spp. of pseudoscorpions (four endemics in the island), seven genera, and six families (Chthoniidae, Dithidae, Olpiidae, Atemnidae, Chernetidae, Cheliferidae). One endemic subgenus Pholeochthonius – troglobite. Notogean genera are Anaulacodithella and Philomaoria. Araneae Rix and Harvey (2010a) described the endemic genus and species of Micropholcommatidae Patelliella adusta and another endemic micropholcommatid species – Taphiassa magna. Endemic genera Fam. Micropholcommatidae Patelliellini (end. tribe) Patelliella Rix et Harvey, 2010 Fam. Cyatholepidae Lordhowea Griswold, 2001 Fam. Zodariidae Basasteron Baehr, 2003 778 8 Some Peculiarities of the Distribution of Arachnida Fam. Phrurolithidae Dorymetaecus Rainbow, 1920 (one sp.) 8.25 Conclusion The level of representation of Arachnida in the classical Notogea (with Papuan area but without Patagonia) is much lower than the level in the vertebrates, with their endemic subclasses, orders, and suborders. Even in the most isolated area (New Zealand), there are no endemics of very high rank. Here are the endemisms above genus: Australia (cont.): One endemic family of scorpions (Urodacidae) Tasmania: Only endemic subfamilies of spiders (Plesiothelinae and Hickmanniinae) New Guinean area: No endemics above genus New Caledonia: One endemic family of Opiliones (Troglosironidae) New Zealand: One endemic family of spiders (Huttoniidae) and one of Opiliones (Synthetonychiidae) Characteristics of the Arachnida in Australia, Tasmania, Papuan Area, New Zealand, New Caledonia, and Lord Howe Island Kingdom Notogaea – No endemic orders or suborders among Arachnida Region of Australia Palpigradi – Two spp. brought from Europe, one local (?) Solifugae - Missing Amblypygi – Fam. Charinidae (Charinus Simon) Thelyphonida (Uropygi) – Not found in Australia Schizomida – Fam. Hubbardiidae (Apozomus Harvey, Attenuizomus Harvey, Bamazomus Harvey, Brignolizomus Harvey, Draculoides Harvey, Julattenius Harvey, Notozomus Harvey, Ovozomus Harvey, Paradraculoides Harvey et al.) Scorpiones – Endemic family is Urodacidae, end. genera Urodacus Peters with 19 species, Isometroides Keyserling and Cercophonius Peters (Australia and Tasmania), and Aops Volschenk et Prendini. Pseudoscorpiones – No endemic families; 150 spp., 17 fam. Endemic genera of pseudoscorpions in Australia (without Tasmania) are fam. Olpiidae, Austrohorus Beier (one sp.), Linnaeolpium Harvey et Leng (one sp.); fam. Cheliferidae, Australochelifer Beier (one sp.); fam. Chernetidae, Conicochernes Beier (four spp.), Marachernes Harvey (three spp.); in Tasmania Neopseudogarypus Morris Opiliones – No endemic families Cyphophthalmi – Fam. Pettalidae (two genera, endemic in Queensland) (Austropurcellia Juberthie) and Western Australia (Karripurcellia Giribet) 8.25 Conclusion 779 Laniatores – Fam. Triaenonychidae (endemic: Breviacantha Kauri, Callihamina Roewer, Callihamus Roewer, Cluniella Forster, Heteronuncia Roewer, Holonuncia Forster, Perthacantha Roewer, Yatala Roewer, Conoculus Forster, Dingupa Forster, Dipristes Roewer), Assamiidae (Dampetrus Karsch, Metamermerus Roewer), Samoidae, Zalmoxidae (Stygnoleptinae) Eupnoi – Fam. Neopilionidae (incl. Monoscutidae), Ballarriinae (Arrallaba Hunt et Cokendolpher, Ballarra Hunt et Cokendolpher, Plesioballarra crinis Hunt et Cokendolpher, Vibone Kauri). Hesperopilio Shear – Unclear family place Dyspnoi – Fam. Acropsopilionidae (Acropsopilio Silvestri) Araneae – Seventy-three families with at least 237 endemic or subendemic genera (without Tasmania). In Tasmania – Two endemic subfamilies and 29 endemic genera Opilioacarida – One undeterm. recorded Holothyrida – Fam. Allothyridae (Allothyrus van der Hammen, Australothyrus van der Hammen) Region New Zealand Very unbalance fauna. Seven orders of Arachnida are missing; the remaining are Pseudoscorpiones (67 spp.), Opiliones (111 spp.), Araneae (1662 spp.), Holothyrida (1 sp.), Ixodida (11 spp.), Mesostigmata, Sarcoptiformes, and Trombidiformes. Particularly well is represented the order Araneae. Palpigradi – Missing Solifugae – Missing Amblypygi – Missing Thelyphonida (Uropygi) – Missing Schizomida – Missing Scorpiones – Missing Pseudoscorpiones – No endemic families; 67 spp. of 27 genera and 8 families. Endemic genera: Maorichthonius Chamberlin, Sathrochthoniella Beier, Tyrannochthoniella Beier, Nelsoninus Beier, Apatochernes Beier (together with Campbell Islands, Snares Islands, Auckland Islands, Chatham Islands, Norfolk Island), Heterochernes Beier Opiliones – One endemic family Cyphophthalmi – Three endemic genera (Rakaia Hirst, Aoraki Boyer et Giribet, Neopurcellia Foster) from the family Pettalidae (in total 21 spp.) Laniatores – Endemic family: Synthetonychiidae (Synthetonychia Forster), Triaenonychidae: Hedwiga Roewer, Hendea Roewer, Hendeola Forster, Metanuncia Roewer (Stewart Island), Neonuncia Roewer, Prasma Roewer, Prasmiola Forster, Psalenoba Roewer, Triregia Forster, Algidia Hogg, Cenefia Roewer, Muscicola Forster, Pristobunus Roewer, subfam. Sørensenellinae (Karamea Forster, Sorensenella Pocock) 780 8 Some Peculiarities of the Distribution of Arachnida Eupnoi – Fam. Caddidae (Acropsopilio Silvestri), Neopilionidae (incl. Monoscutidae) (Forsteropsalis Taylor (nine spp.), Mangatangi Taylor, Pantopsalis Simon; Monoscutinae (end. subfamily): Acihasta Forster, Monoscutum Forster, Templar Taylor) (New Zealand, Auckland, Snares, Campbell Islands) Dyspnoi – Missing Araneae: One endemic family (Huttoniidae), 1 subfamily (Pahorinae), and at least 93 endemic genera of spiders: Fam. Huttoniidae (Huttonia Pickard-Cambridge), Hexathelidae (Hexathele Ausserer, Porrhothele Simon), Anapidae (Novanapis Platnick et Forster, Paranapis Platnick et Forster, Zealanapis Platnick et Forster), Pararcheidae (Forstrarchaea Rix, Pararchaea Forster), Cycloctenidae (Plectophanes Bryant, Toxopsiella Forster, Uzakia Koçak et Kemal, Cantuarea Hogg), Gradungulidae (Gradungula Forster, Pianoa Forster, Spelungula Forster), Orsolobidae (Anopsolobus Forster et Platnick, Ascuta Forster, Bealeyia Forster et Platnick, Dugdalea Forster et Platnick, Duripelta Forster, Maoriata Forster et Platnick, Orongia Forster et Platnick, Paralobus Forster et Platnick, Pounamuella Forster et Platnick, Subantarctia Forster, Tangata Forster et Platnick, Tautukua Forster et Platnick, Turretia Forster et Platnick, Waiporia Forster et Platnick, Wiltonia Forster et Platnick), Mecysmauchenidae (Aotearoa Forster et Platnick, Zearchaea Wilton, Parapua Forster, Pua Forster, Forstarchaea Rix, Tekelloides Forster, Waitkerra Opell, Nomana Forster, Pahora Forster, Pahoroides Forster, Runga Forster, Wairua Forster, Meringa Forster, Mangua Forster, Ahua Forster et Wilton, Huka Forster et Wilton, Mahura Forster et Wilton, Neoramia Forster et Wilton, Oramia Forster et Wilton, Oramiella Forster et Wilton, Orepukia Forster et Wilton, Paramyro Forster et Wilton, Porotaka Forster et Wilton, Tararua Forster et Wilton, Tuapoka Forster et Wilton, Anhunga Forster et Wilton, Maloides Forster et Wilton, Muritaia Forster et Wilton, Pakeha Forster et Wilton, Paravoka Forster et Wilton, Poaka Forster et Wilton, Gasparia Forster et Wilton, Gohia Forster et Wilton, Goyenia Forster et Wilton, Hapona Forster, Helsinia Forster, Hulua Forster et Wilton, Lamina Forster, Mangareia Forster, Matachia Dalmas, Mesudus Özdikmen, Neomyro Forster et Wilton, Notomatachia Forster, Nuisiana Forster et Wilton, Otagoa Forster, Panoa Forster, Rapua Forster, Toxopsoides Forster et Wi

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