R evue
suisse de
Zoologie 117 (4): 557-598; décembre 2010
Global checklist of pseudoscorpions (Arachnida)
found in birds’ nests
Paola TURIENZO1, Osvaldo DI IORIO1 & Volker MAHNERT2
Departamento de Biodiversidad y Biología Experimental, Facultad de
Ciencias Exactas y Naturales, 4º Piso, Pabellón II, Ciudad Universitaria
C1428EHA, Buenos Aires, Argentina. E-mail: paolaturienzo@yahoo.com
2 Volker Mahnert, Muséum d’histoire naturelle, 1, route de Malagnou,
CH-1208 Geneva, Switzerland. E-mail: volker.mahnert@wanadoo.fr
1 Entomología.
Global checklist of pseudoscorpions (Arachnida) found in birds’ nests.
- A compilation of the information on pseudoscorpions recorded from birds’
nests is presented. Species and families of birds, and specimens, species and
families of pseudoscorpions involved worldwide are given. 14 families (63
genera with 85 species) of Pseudoscorpiones were found in birds’ nests, of
which the Chernetidae is ranked first, with 22 genera and 35 species. Only
45 families of birds have been recorded as hosts worldwide, represented by
a total of 98 species and a certain number of unidentified taxa. Geographical
origin of these records reflects collecting efforts of individuals and gives no
precise indication on the biological association between these two groups.
Keywords: Ecology - behaviour - phoresy - host association.
INTRODUCTION
The arachnid order Pseudoscorpiones, commonly known as pseudoscorpions,
false scorpions, or book scorpions, comprise about 3.3% of the described arachnids,
with 3 385 known species (Harvey, 2009). Pseudoscorpions are small (typically less
than 4 mm in length) predacious arachnids, feeding upon small invertebrates (e.g.,
mites, springtails, various flies and gnats, ants, insect larvae, and occasionally small
caterpillars). The general ecology of the group is poorly known; the most intensive
studies on tropical pseudoscorpions have been carried out in the Manaus Amazon
region, under the guidance of the late Professor J. Adis (Plön, Germany) (e.g. Adis,
1979, 1992; Adis & Mahnert, 1985, 1990). They live in leaf litter, under rocks, in
compost piles, under bark and in decaying wood, in caves, and in various invertebrate
and vertebrate nests (Beier, 1948, 1963b; Weygoldt, 1969).
No answer can be given to the question whether pseudoscorpions are arriving
accidentally in the nests, if they are regular visitors, or if nests represent their favoured
habitat. Beier (1948) had in his compilation enumerated 12 species found in birds’
nests or on birds, citing two species as living exclusively in this biotope: Diplotemnus
insularis Chamberlin and Hesperochernes montanus Chamberlin, but all these
conclusions were based on isolated findings. An exhaustive ecological study had been
Manuscript accepted 20.07.2010
p. tuRienZo et al.
558
carried out by Nordberg (1936) on the nidicolous arthropods of bird’s nests in Finland
(mainly from Aland, a few nests also from Tvärminne and Helsingfors), but unfortunately only three pseudoscorpion species had been identified. Krumpal & Cyprich
(1988) tried to define ecological preferences of the 19 pseudoscorpion species found
during their survey in the Czech Republic in 162 nests (representing 54 bird nest types)
in relation to nest type (free nests, nests in nest boxes, nests in synanthropic habitats,
nests in hollow trees) and seasonal changes. Ressl (1963) and Simon (1966), but also
Jones (1978) discussed the eventual dispersal of pseudoscorpions by birds and the
possible colonization of birds’ nests. Recently, Christophoryova & Krumpalova (2010)
distinguish within the pseudoscorpions nidixenous species (which occur accidentally
in bird nests, the nymphal stages are represented sporadically) and nidiphilous or
nidicolous species. They occur regularly in the bird nests, the nymphal stages are
represented numerously, and prefer nests in certain habitats with specific building and
microclimagtic conditions.
An ongoing study on nidicolous arthropods in birds’ nests from Argentina
(Turienzo & Di Iorio, 2008; Turienzo et al., 2008, Di Iorio et al., 2010; Turienzo & Di
Iorio, unpublished data) has yielded 246643 insects, 5378 soft ticks (Acarina:
Argasidae) and 6406 pseudoscorpions. Consequently, a summarization of the
knowledge concerning the pseudoscorpions known from this biotope worldwide is
needed for further comparison. The main purpose of the present work is to: 1) provide
a compilation of the information on pseudoscorpion species and families recorded
from birds’ nests, and 2) state which bird species and families are involved worldwide.
MATERIAL AND METHODS
The taxonomic and biological publications on pseudoscorpions were reviewed,
as well as papers treating inhabitants of birds’ nests; also included are unpublished
identifications collated by one of us (V. M.). The bird families and species, followed
by those of the associated pseudoscorpions, are listed in alphabetical order. All
references were verified, except those indicated as “cited by”, Museum collections and
indications not given in original sources are cited between square brackets [ ]. Bird
species lists of the American Ornithologists’ Union (1998), Remsen et al. (2008) and
the Comitê Brasileiro de Registros Ornitológicos (2007) were used for current nomenclature; subspecific taxa not treated in these lists were updated from Peterson (2002).
The nomenclature of pseudoscorpions is adopted from Harvey (2009). Old combinations, synonyms, misidentifications, original localities, number of specimens, and
depositories when they were stated, are provided in each case. Pertinent comments in
the references and controversial data are included in notes below the respective items.
Collections mentioned in the text
AM
BISHOP
CNCI
EMB
Australian Museum, Sydney, Australia.
Bishop Museum, Honolulu, Hawaii.
Canadian National Collection of Insects, Ottawa, Canada.
Ellen M. Benedict collection, Pacific University, Forest Grove, Oregon,
USA
pseudoscoRpions in biRds’ nests
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INPA
Instituto Nacional de Pesquisas da Amazônia, Núcleo de Pesquisas em
Ciências Humanas e Sociais, Manaus, Brazil.
LZPb
Laboratorio de Zoología, Departamento de Biología, Universidade Federal
da Paraíba, João Pessoa, Paraíba, Brazil.
MNRJ Museu Nacional, Rio de Janeiro, Brazil.
NHMW Naturhistorisches Museum, Wien, Austria.
WAM
Western Australian Museum, Perth, Australia.
PSEUDOSCORPION TAXA ASSOCIATED WITH BIRDS, NESTS AND GUANO
ATEMNIDAE
Anatemnus orites major Beier: Pycnonotidae.
Anatemnus orites orites (Thorell): Apodidae.
Diplotemnus insularis Chamberlin: “in bird nests and on birds“ (probably
Sulidae and Laridae).
Oratemnus navigator (With): Apodidae.
Paratemnoides nidificator (Balzan): Passeriformes.
CHEIRIDIIDAE
Apocheiridium (A.) indicum Murthy & Ananthakrishnan: Passeridae.
Apocheiridium (A.) minutissimum Beier: Accipitridae, Halcyonidae, Phasianidae.
Apocheiridium (A.) rossicum Redikorzev: Anatidae.
Apocheiridium (A.) stannardi Hoff: Anatidae.
Cheiridium museorum (Leach): Apodidae, Columbidae, Corvidae, Fringillidae,
Hirundinidae, Motacillidae, Paridae, Passeridae, Phasianidae, Remizidae,
Sturniidae, Turdidae; birds’ nests.
CHELIFERIDAE
Chelifer cancroides cancroides (Linnaeus): Apodidae, Columbidae, Corvidae, Fringillidae, Hirundinidae, Laridae, Phasianidae, Picidae, Strigidae, Sturniidae,
Sylviidae, Turdidae, Tyrannidae; birds’ nests.
Chelifer cancroides orientalis Morikawa: Hirundinidae.
Chelifer sp.: Passeridae.
Dactylochelifer latreillei latreillei (Leach): Columbidae, Emberizidae, Hirundinidae, Laniidae, Paridae, Passeriformes, Remizidae, Sylviidae, Turdidae.
Hansenius torulosus (Tullgren): Phasianidae.
Paisochelifer callus (Hoff): birds’ nests.
Rhacochelifer maculatus (L. Koch): Phalacrocoracidae.
CHERNETIDAE
Acuminochernes crassipalpus (Hoff): Strigidae.
Acuminochernes tacitus Hoff: Picidae.
Allochernes powelli (Kew): Hirundinidae, Passeriformes.
Allochernes wideri (C. L. Koch): Coraciidae, Corvidae, Muscicapidae, Paridae,
Passeridae, Phasianidae, Picidae, Sittidae, Strigidae, Turdidae; birds’ nests.
Anthrenochernes stellae Lohmander: Corvidae.
Americhernes oblongus (Say): Anatidae.
Apatochernes antarcticus pterodromae Beier: Procellariidae.
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Apatochernes nestoris Beier: Procellariidae, Psittacidae.
Calidiochernes musculi Beier: Furnariidae, Troglodytidae.
Chelanops skottsbergi (Beier): Procellariidae.
Chernes cimicoides (Fabricius): Ardeidae, Hirundinidae, Laniidae, Paridae,
Strigidae, Sylviidae.
Chernes hahnii (L. Koch): Meropidae, Paridae, Passeriformes; birds’ nests.
Chernes vicinus (Beier): birds’ nests.
Chernes sp.: Passeridae.
Dendrochernes cyrneus (L. Koch): Passeridae; birds’ nests.
Dinocheirus panzeri (C. L. Koch): Columbidae, Corvidae, Hirundinidae, Laniidae,
Paridae, Passeriformes, Phasianidae, Strigidae, Sturnidae, Sylviidae, Turdidae.
Dinocheirus sicarius Chamberlin: Phasianidae.
Hesperochernes montanus Chamberlin: Corvidae; birds’ nests.
Lamprochernes chyzeri (Tömösvary): Phasianidae.
Lamprochernes nodosus (Schrank): Meropidae, Panuridae, Paridae, Remizidae;
birds’ nests.
Lamprochernes savignyi (Simon): Phasianidae; birds’ nests.
Lamprochernes sp.: Hirundinidae.
Neochelanops patagonicus (Tullgren): birds’ nests.
Opsochernes carbophilus Beier: birds’ nests.
Parachernes argentinus Beier: Furnariidae; birds’ nests.
Parachernes cf. pulcher Mahnert: Furnariidae.
Parachernes squarrosus Hoff: Turdidae.
Parazaona bucheri Beier: Furnariidae, Psittacidae.
Parazaona morenensis (Tullgren): Psittacidae.
Pselaphochernes anachoreta (Simon): Corvidae.
Pselaphochernes scorpioides (Hermann): Corvidae, Passeridae, Upupidae; birds’
nests.
Pselaphochernes lacertosus (L. Koch): Procellariidae.
Reischekia exigua exigua Beier: Ardeidae.
Rhopalochernes sp.: Icteridae.
Sundochernes malayanus Beier: Pycnonotidae.
Tejachernes stercoreus (Turk): Hirundinidae.
Troglochernes dewae (Beier): Cacatuidae, Climacteridae.
CHTHONIIDAE
Chthonius (Chthonius) ischnocheles (Hermann): Hirundinidae, Turdidae.
Chthonius (Chthonius) rhodochelatus Hadzi: Passeridae.
Chthonius (Ephippiochthonius) fuscimanus Simon: Emberizidae, Passeriformes,
Turdidae.
Chthonius (Ephippiochthonius) tetrachelatus (Preyssler): Anatidae, Passeriformes,
Turdidae.
Chthonius sp.: Muscicapidae, birds’ nests.
Mundochthonius styriacus Beier: Paridae.
Pseudochthonius homodentatus Chamberlin: Passeriformes, Turdidae.
Tyrannochthonius heterodentatus Beier: Passeridae.
pseudoscoRpions in biRds’ nests
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Tyrannochthonius kermadecensis (Beier): Turdidae.
Tyrannochthonius migrans Mahnert: Troglodytidae.
GARyPIDAE
Garypus titanius Beier: in guano of birds’ colonies.
GEOGARyPIDAE
Geogarypus albus Beier: Pycnonotidae; birds’ nests.
Geogarypus longidigitatus (Rainbow): Cisticolidae, Estrilidae, Nectarinidae,
Pycnonotidae.
Geogarypus minor (Tullgren): Cisticolidae.
Geogarypus minutus (Tullkgren): Cisticolidae.
IDEORONCIDAE
Xorilbia cf. gracilis (Mahnert): Troglodytidae.
LARCIDAE
Larca lata (Hansen): Phasianidae, Turdidae, Tytoniidae; birds’ nests.
Larca notha Hoff: Hirundinidae.
NEOBISIIDAE
Neobisium (Neobisium) carcinoides (Hermann): Cinclidae, Emberizidae, Troglodytidae; birds’ nests; birds’ nests type Motacilla; nests Cinclidae+Motacilla.
Neobisium (Neobisium) crassifemoratum Beier: Tetraonidae; birds’ nests.
Neobisium (Neobisium) inaequale Chamberlin: Sylviidae; birds’ nests type
Motacilla.
Neobisium (Neobisium) sylvaticum (C. L. Koch): Corvidae, Laniidae, Sylviidae,
Turdidae.
Neobisium sp.: Emberizidae, Hirundinidae, Motacillidae, Turdidae.
OLPIIDAE
Calocheiridius olivieri (Simon): Phalacrocoracidae.
Indolpium loyolae (Murthy): Passeridae.
Pachyolpium irmgardae Mahnert: Formicariidae, Troglodytidae.
Xenolpium pacificum pacificum (With): birds’ nests.
PSEUDOGARyPINIDAE
Pseudogarypinus frontalis (Banks): birds’ nests.
SyARINIDAE
Ideobisium sp.: Passeriformes.
WITHIIDAE
Dolichowithius argentinus Beier: birds’ nests.
Stenowithius bayoni (Ellingsen): Anatidae, Laniidae.
Stenowithius duffeyi Beier: birds’ guano in nesting colonies.
Victorwithius proximus (Beier): Furnariidae.
Withius indicus Murthy & Ananthakrishnan: Phasianidae.
Withius kaestneri (Vachon): Phasianidae.
Withius piger (Simon): Phasianidae.
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BIRD TAXA AND ASSOCIATED PSEUDOSCORPIONS
ACCIPITRIDAE
Ictinaetus malayensis Temminck, 1892
CHEIRIDIIDAE
Apocheiridium minutissimum Beier, 1964
Malaysia: Pahang, Fraser’s Hill, 15-I-1956, McClure leg., 1 exuvia, Ser. No.
M-03118, Band No. H. 87082, in a nest of Black Eagle (Beier, 1964).
ANATIDAE
Aix sponsa (Linnaeus, 1758)
CHEIRIDIIDAE
Apocheiridium stannardi Hoff, 1952
USA: Michigan, Shiawassee County, in nest box (Nelson, 1971: “accidental
occurrence?”).
CHERNETIDAE
Americhernes oblongus (Say, 1821) (= Lamprochernes oblongus: Nelson, 1971).
USA: Michigan, Shiawassee County, in nest box (Nelson, 1971: “accidental occurrence”).
Anas platyrhynchos (Linnaeus, 1758)
CHTHONIIDAE
Chthonius (Ephippiochthonius) tetrachelatus (Preyssler, 1790)
Slovakia: (Fenda et al., 1998).
Anas undulata (Dubois, 1839)
WITHIIDAE
Stenowithius bayoni (Ellingsen, 1910)
Uganda: Chagwe, Seziwa Swamps (Ellingsen, 1913; Beier, 1948).
Undetermined species (“wild duck”: Redikorzev, 1938; Beier, 1948).
CHEIRIDIIDAE
Apocheiridium rossicum Redikorzev, 1935
Russia: Siberia, Ienissei, on head of wild duck (Redikorzev, 1938; Beier,
1948).
APODIDAE
Apus affinis (Gray, 1830) (= Cypselus affinis: Ellingsen, 1914, Beier, 1948).
ATEMNIDAE
Anatemnus orites orites (Thorell, 1889)
India: Calcutta, from a nest in the Museum buildings (Ellingsen, 1914;
Beier, 1948).
Oratemnus navigator (With, 1906)
India: Calcutta, from a nest in the Museum buildings (Ellingsen, 1914;
Beier, 1948).
CHEIRIDIIDAE
Cheiridium museorum (Leach, 1817)
pseudoscoRpions in biRds’ nests
563
India: Calcutta, from a nest in the Museum buildings (Ellingsen, 1914;
Beier, 1948).
Apus apus (Linnaeus, 1758)
CHEIRIDIIDAE
Cheiridium museorum (Leach, 1817)
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1983).
CHELIFERIDAE
Chelifer cancroides cancroides (Linnaeus, 1758)
Austria: Lower Austria, Scheibbs Distr. (Ressl & Beier, 1958; Ressl, 1963,
1983).
ARDEIDAE
Ardea cinerea Linnaeus, 1758
CHERNETIDAE
Chernes cimicoides (Fabricius, 1793)
England: Kent, High Halstow (Jones, 1975, 1978).
Egretta alba modesta (Gray, 1831)
CHERNETIDAE
Reischekia exigua exigua Beier, 1976
New Zealand: Westland, Okarito, in old broken-down white heron nests,
sample 68/5 (Beier, 1976).
CACATUIDAE
Cacatua galerita galerita (Latham, 1790)
CHERNETIDAE
Troglochernes dewae (Beier, 1967)
Australia: Queensland, Fringe Dwellers, Iron Range, circa 12º38’S,
143º05’E, 9-X-1998, S. Legge & R. Heinsohn leg., 1� [WAM T66299],
nest of Sulphur-Crested Cockatoo (Harvey & Volschenk, 2007).
Calyptorhynchus latirostris Carnaby, 1948 (= Cacatua latirostris: Harvey &
Volschenk, 2007).
CHERNETIDAE
Troglochernes dewae (Beier, 1967)
Australia: Western Australia, Shire of Moora, 30º35’S, 116º01’E, 20-XI1998, P. Mawson leg., 1� [WAM T66300], ex nest in healthy hollow of
Eucalyptus salmonophloia (Harvey & Volschenk, 2007); Gingin Shire,
30º59’S, 115º45’E, 20-XI-1998, P. Mawson leg., 1 tritonymph, 1�
[WAM T48341], ex nest # 84 (Harvey & Volschenk, 2007).
Eulophus roseicapilla (Vieillot, 1817) (= Cacatua roseicapilla: Beier, 1967; Harvey &
Volschenk, 2007).
CHERNETIDAE
Troglochernes dewae (Beier, 1967) (= Sundochernes dewae: Beier, 1967).
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Australia: New South Wales, Brewarrina, 29º58’S, 146º52’E, � holotype
[AM KS5867], 1 protonymph, 2 deutonymphs, 2 tritonymphs, 1�
(paratypes) [AM KS5868], 1 nymph, 1�, 2� (paratypes) [NHMW],
V-1964, B. Dew leg., from nest of Galah in hollow tree (Beier, 1967,
Harvey & Volschenk, 2007).
CINCLIDAE
Cinclus cinclus (Linnaeus, 1758)
NEOBISIIDAE
Neobisium (Neobisium) carcinoides (Hermann, 1804)
Slovakia: (Fenda et al., 1998).
Cinclus cinclus (Linnaeus, 1758) + Motacilla sp.
Neobsium (Neobisium) carcinoides (Hermann, 1804)
Slovakia: (Fenda et al., 1998).
CISTICOLIDAE
Cisticola lais Hartlaub & Finsch, 1870
GEOGARyPIDAE
Geogarypus minutus (Tullgren, 1907) (= Garypus minutus: Elingsen, 1912).
South Africa: Cape Province, King William’s Town Div.: Pirie, a very
immature specimen rather doubtfully referable to G. minutus, from a
nest of a grass warbler (Cisticola lais Sharpe) (Ellingsen, 1912).
Prinia flaviventris (Delessert, 1840)
GEOGARyPIDAE
Geogarypus longidigitatus (Rainbow, 1897) (= Geogarypus elegans audyi
Beier, 1952: Beier, 1963a).
Malaysia: Selangor, Rantau Panjang, 8 km N of Klang, 12-VII-1961, 1 tritonymph, nest 234, B.53395 (Beier, 1963a).
CLIMACTERIDAE
Climacteris rufa Gould, 1841
CHERNETIDAE
Troglochernes dewae (Beier, 1967)
Australia: Western Australia, yilliminning Agricultural Region, 32º56’S,
117º25’E, 6-III-1999, G. Luck leg., 1 deutonymph, 2 tritonymphs, 1�,
1� [WAM T66301], ex nest # 206 (Harvey & Volschenk, 2007).
COLUMBIDAE
Columba livia domestica Gmelin, 1789 (= Columba sp., city pigeon: Woodroffe,
1953).
CHEIRIDIDAE
Cheiridium museorum (Leach, 1817)
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1963).
pseudoscoRpions in biRds’ nests
565
England: Leicester, The Newarke, from nest of feral pigeon (Jones, 1975).
Finland: Aland (? Tvärminne or Helsingfors) (Nordberg, 1936).
Germany: Rheinland-Westfalen, Peppenhoven am Vorgebirge; Morenhoven
am Vorgebirge, in einem Taubenschlag (Ellingsen, 1911; Beier, 1948).
[Undetermined species]
Brazil: Amazonas, Manaus, VII/XI-2005, 3 ex. [INPA] in nest # 5, 4 ex.
[INPA] in nest # 8 from a total of 14 examined nests, 0.28% of total
arthropods (Ogawa, 2008).
CHELIFERIDAE
Chelifer cancroides cancroides (Linnaeus, 1758)
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1963, 1983).
England: abundant in some pigeons’ nests (Woodroffe, 1953).
Finland: Aland (? Tvärminne or Helsingfors) (Nordberg, 1936).
Germany: Rheinland-Westfalen, Peppenhoven am Vorgebirge, in einem
Taubenschlag (Ellingsen, 1911; Beier, 1948).
Sweden: (Lohmander, 1939; Beier, 1948).
CHERNETIDAE
Dinocheirus panzeri (C. L. Koch, 1837)
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1983).
Ireland: Portaferry, County Down, pigeon’s nest in an old tower (Legg &
O’Connor, 1997; Jones, 1975).
Columba oenas Linnaeus, 1758
CHELIFERIDAE
Chelifer cancroides cancroides (Linnaeus, 1758)
Finland: Aland (? Tvärminne or Helsingfors) (Nordberg, 1936).
Dactylochelifer latreillei (Leach, 1817)
Finland: Aland (? Tvärminne or Helsingfors) (Nordberg, 1936; Krištofik
et al., 1993, 2002) (mis-identified, see Chernes cimicoides).
CHERNETIDAE
Chernes cimicoides (Fabricius, 1793) and Dinocheirus panzeri (C. L. Koch,
1837)
Finland: Aland (? Tvärminne or Helsingfors) (Nordberg, 1936). The
samples detemined as D. latreillei consist of individuals of two other
species, Chernes cimicoides and Toxochernes (= Dinocheirus) panzeri
(Kaisila, 1949).
CORACIIDAE
Coracias garrulus Linnaeus, 1758
CHERNETIDAE
Allochernes wideri (C. L. Koch, 1843)
Slovakia (and/or Austria-Czech
Krumpalova, 2010).
Republic)
(Christophoryova
&
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CORVIDAE
Corvus corone corone Linnaeus, 1758
CHERNETIDAE
Chernes hahnii L. Koch, 1873
France: Haute-Savoie, Bossy-sur-Frangy, nid, 6-V-1972, leg. J. Steffen,
1�, det.V. Mahnert.
Pselaphochernes anachoreta (Simon, 1878)
France: Haute-Savoie, Bossy-sur-Frangy, nid, 6-V-1972, leg. J. Steffen,
1�, det.V. Mahnert.
NEOBISIIDAE
Neobisium (Neobisium) sylvaticum (C. L. Koch, 1835)
Austria: Lower Austria, Scheibbs Distr. (Ressl & Beier, 1958; Ressl 1983).
Note: Records of Chelifer cancroides, Allochernes wideri and Dinocheirus panzeri
from nests of Corvus corone by Ressl (1963) were apparently due to a typographical error, and concern the occurrence in nests of Sturnus vulgaris. They
were implicitly corrected by Ressl (1983).
Corvus monedula Linnaeus, 1758 (= Colaeus monedula: Nordberg 1936).
CHEIRIDIIDAE
Cheiridium museorum (Lach, 1817)
United Kingdom: Wales, Ceredigion, in a jackdaw’s nest in the lofts (Chater,
1989).
CHELIFERIDAE
Chelifer cancroides cancroides (Linnaeus, 1758)
Finland: Aland (? Tvärminne or Helsingfors) (Nordberg, 1936).
CHERNETIDAE
Allochernes wideri (C. L. Koch, 1843)
Germany: Hessen, Wiesbaden, Biebricher Schlosspark, nest in broken
Populus alba (Helversen, 1966).
Anthrenochernes stellae Lohmander, 1939
Sweden: Blekinge, probably nest of C. monedula (Gärdenfors & Wilander,
1995).
Dinocheirus panzeri (C. L. Koch, 1837) (= Toxochernes panzeri (C. L. Koch);
= Chernes rufeolus (Simon): Helversen, 1966).
Germany: Hessen, Wiesbaden, Biebricher Schlosspark, nest in broken
Populus alba (Helversen, 1966).
Sweden: Skane, Torna Hällestad; Östergötland, Alvastra (Lohmander, 1939;
Beier, 1948); probably nest of C. monedula (Gärdenfors & Wilander,
1995).
Pselaphochernes scorpioides (Hermann, 1804)
England: Berkshire, Wytham Wood, Jackdaw’s nest at 15 ft in an oak tree
(Jones, 1975, 1978).
Pica hudsonia Sabine, 1823 (= Pica pica hudsonicus Sabine: Chamberlin, 1935).
CHERNETIDAE
Hesperochernes montanus Chamberlin, 1935
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567
USA: Montana, Bitterroot Valley, 1931, “one pseudoscorpion that has been
determined as Hesperochernes montanus new species by J. C.
Chamberlin who is describing this arachnid elsewhere” (Jellison &
Philip, 1933), Ravally County, Girds Creek, 30-V-1931, W. L. Jellison
leg., 1�, # JC-796.01001, in a bird’s nest (Chamberlin, 1935; Beier,
1948).
EMBERIZIDAE
Emberiza citrinella Linnaeus, 1758
CHTHONIIDAE
Chthonius (Ephippiochthonius) fuscimanus Simon, 1900 (= Chthonius austriacus: Ressl, 1963).
Austria: Lower Austria, Scheibbs Distr., Purgstall (Ressl, 1963, 1983).
CHELIFERIDAE
Dactylochelifer latreillei latreillei (Leach, 1817)
Austria: Lower Austria, Scheibbs Distr. (Ressl & Beier, 1958; Ressl, 1963,
1983).
NEOBISIIDAE
Neobisium (Neobisium) carcinoides (Hermann, 1804)
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1983).
Neobisium sp.
Slovakia: (Fenda et al., 1998).
ESTRILDIDAE
Lonchura maja (Linnaeus, 1766)
GEOGARyPIDAE
Geogarypus longidigitatus (Rainbow, 1897) (= Geogarypus elegans audyi
Beier, 1952: Beier, 1963a).
Malaysia: Selangor, Rantau Panjang, 8 km N of Klang, 15-III-1961, 1 tritonymph, nest 54, B.53085 (Beier, 1963a).
Lonchura malacca (Sykes, 1832)
GEOGARyPIDAE
Geogarypus longidigitatus (Rainbow, 1897) (= Geogarypus elegans audyi
Beier, 1952: Beier, 1963a).
Malaysia: Selangor, Rantau Panjang, 8 km N of Klang, 18-I-1961, 1 D, 1 T,
3�, 1�, nest 13, B.52897; 31-V-1961, 1�, nest 166, B.53259; 14-VI1961, 2�, nest 192, B.53341 (Beier, 1963a).
Lonchura punctulata (Linnaeus, 1758)
GEOGARyPIDAE
Geogarypus longidigitatus (Rainbow, 1897) (= Geogarypus elegans audyi
Beier, 1952: Beier, 1963a).
Malaysia: Selangor, Rantau Panjang, 8 km N of Klang, 18-X-1961, 1 gravid
�, nest 250, B.53576 (Beier, 1963a).
568
p. tuRienZo et al.
FORMICARIIDAE
Formicariidae sp. 2
OLPIIDAE
Pachyolpium irmgardae Mahnert, 1979
Brazil: Amazonas, Municipios Silves & Itacotiara, Mil Madereira Itacotiara
Ltda., 3º00’S, 58º40’ W, VII/XII-2000, 1 nymph (Torres, 2001).
FRINGILLIDAE
Fringilla coelebs Linnaeus, 1758
CHEIRIDIIDAE
Cheiridium museorum (Leach, 1817)
Europe: (Roewer, 1937; Beier, 1948).
CHELIFERIDAE
Chelifer cancroides cancroides (Linnaeus, 1758)
Finland: Arland (? Tvärminne or Helsingfors) (Nordberg, 1936; Kaisila,
1949).
FURNARIIDAE
Anumbius annumbi (Vieillot, 1817)
CHERNETIDAE
Parachernes cf. pulcher Mahnert, 1979
Argentina: Córdoba, Dto Unión, Bell Ville (Grimaldi et al., 2006).
Parazaona bucheri Beier, 1967
Argentina: Santa Fe, near Santa Fe City, in a nest superimposed to a nest of
Phacellodomus ruber (Vieillot, 1817) [Aves: Furnariidae] (Turienzo et
al., 2008).
WITHIIDAE
Victorwithius proximus (Ellingsen, 1905)
Argentina: Buenos Aires [Campo de Mayo] (Turienzo et al., 2008).
Phacellodomus ruber (Vieillot, 1817)
CHERNETIDAE
Parazaona bucheri Beier, 1967
Argentina: Santa Fe, near Santa Fe City (Turienzo et al., 2008).
Phacellodomus rufifrons inornatus Ridgway, 1887
CHERNETIDAE
Calidiochernes musculi Beier, 1957
Venezuela: “Alta Gracia de Oviteno”, det. M. Beier, in nest, hand-written
note by M. Beier in his private copy of Beier, 1957a: 138.
MONOSPHyRONIDA
undetermined species
Colombia: Department of Meta, El Porvenir, 4º45’N, 71º25’W, in 22 examined nests (Barreto et al., 1984).
pseudoscoRpions in biRds’ nests
569
UNDETERMINED FAMILy
Undetermined species (= “Chelifer sp.”: Pifano, 1938).
Venezuela: Estado yaracuy, Distrito San Felipe, Carretera San Javier Marín, II-1938, Anduze & Pifano leg., in company of Coleoptera and
Psammolestes arthuri (Pinto) [Hemiptera: Reduviidae] (Pifano, 1938;
Carcavallo et al., 1975).
Phacellodomus rufifrons rufifrons (Wied-Neuwied, 1821)
PSEUDOSCORPIONES, undetermined species
Brazil: Goias (Silva & Lustosa, 1993).
Phacellodomus rufifrons sincipitalis Cabanis, 1883
PSEUDOSCORPIONES, undetermined species
Brazil: (Lent, 1939); Pernambuco, March to June 1970, 344 ex. [LZPb],
from a total of 28 nests (Lins Duarte, 1978).
Phacellodomus sp.
CHERNETIDAE
Parachernes argentinus Beier, 1967
Argentina: Tucumán, La Soledad, near Cañete, 1965-1966, identified by
Beier (Bucher, 1974).
Parazaona bucheri Beier, 1967
Argentina: Tucumán, Tapia-Raco, identified by Beier (Bucher, 1974).
Note: P. argentinus and P. bucheri were described by Beier (1967) on specimens
collected by Bucher in 1966 and sent to Beier for identification. Bucher (1974)
specified that the specimens had been collected from nests of Phacellodomus,
information probably unknown to Max Beier.
HALCyONIDAE
Halcyon pileata (Boddaert, 1783)
CHEIRIDIIDAE
Apocheiridium minutissimum Beier, 1964
Malaysia: Subang, 12-XII-1962, H. E. McClure leg., � holotype [BISHOP
3603], Ser. No. M-02066, Band No. H. 87082, in nest (Beier, 1964).
HIRUNDINIDAE
Delichon urbica (Linnaeus, 1758)
CHEIRIDIIDAE
Cheiridium museorum (Leach, 1817)
Austria: Lower Austria, Scheibbs Distr., Purgstall (Ressl, 1963, 1983).
Scotland: near Rannoch, Perthshire, in nest of house martin (Jones, 1975,
1978).
CHELIFERIDAE
Chelifer cancroides cancroides (Linnaeus, 1758)
Austria: Lower Austria, 1862, Ferrari leg, “aus einem Schwalbennest”
(Beier, 1929a); under Hirundo rustica and Delichon urbica (Beier,
1948); Scheibbs Distr. (Ressl, 1963 1983).
570
p. tuRienZo et al.
Finland: Aland (? Tvärminne or Helsingfors) (Nordberg, 1936).
Sweden: Östergötland, Lake Täkern, leg. V. A. Engholm (Lohmander, 1939;
Beier, 1948).
Switzerland: (Lessert, 1911; Beier, 1948); canton of Vaud, Commugny,
1964/1966, in nests, leg. J. Steffen, 5�, 10�, 1 trito-, 3 deutonymphs, det.
V. Mahnert.
Chelifer cancroides orientalis Morikawa, 1954
Japan: Nagano Prefecture, Agematsu Town, 17-XI-1967, in 8 examined
nests (Uchikawa, 1970).
Dactylochelifer latreillei latreillei (Leach, 1817)
Finland: Aland (? Tvärminne or Helsingfors) (Nordberg, 1936; Krištofik et
al., 1993) (mis-identified, see Chernes cimicoides).
CHERNETIDAE
Chernes cimicoides (Fabricius, 1793) and Dinocheirus panzeri (C. L. Koch,
1837)
Finland: Aland (? Tvärminne or Helsingfors) (Nordberg, 1936). The
samples detemined as D. latreillei consist of individuals of two other
species, Chernes cimicoides and Toxochernes (=Dinocheirus) panzeri
(Kaisila, 1949).
CHTHONIIDAE
Chthonius (Chthonius) ischnocheles (Hermann, 1804)
England: Churcham, Gloucestershire (Jones, 1975).
Delichon urbica (Linnaeus, 1758) + Passer domesticus (Linnaeus, 1758)
CHEIRIDIIDAE
Cheiridium museorum (Leach, 1817)
England: Ashleworth, Gloucestershire, in nest of house martin, later
occupied by sparrow (Jones, 1975, 1978).
Hirundo rustica (Linnaeus, 1758)
CHEIRIDIIDAE
Cheiridium museorum (Leach, 1817)
Europe (?): (Roewer, 1937; Beier, 1948).
CHELIFERIDAE
Chelifer cancroides cancroides (Linnaeus, 1758)
Austria: Lower Austria, Dornbach bei Wien, 1862, Ferrari leg., “aus einem
Schwalbennest” (Beier, 1929a); “Hirundo rustica” (Beier, 1948);
Scheibbs Distr. (Ressl & Beier, 1958; Ressl 1963, 1983).
Finland: Aland (? Tvärminne or Helsingfors) (Nordberg, 1936).
France: Haute-Savoie, Bossy-sur-Frangy, 450m, in nests in stable and
kitchen of farmstead, 1969-1980, leg. J. Steffen, det. V. Mahnert.
Germany: Hessen, Naturpark Hoher Vogelsberg, Ilbeshausen (Jost, 1982).
Sweden: (Lohmander, 1939).
Switzerland: (Lessert, 1911; Beier, 1948); canton of Vaud, Commugny, in
nest, 8-VII-1962, leg. J. Steffen, 3�, 1�, det. Mahnert.
pseudoscoRpions in biRds’ nests
571
USA: Oregon, Lane Co., 2.5 mi N of Cheshire, 4-XII-1971, leg. E. M.
Benedict, 5 nymphs, 5�, 4� [EMB], hay, mouse and barn swallow
nests in old sheep shed (Benedict & Malcolm, 1979).
Dactylochelifer latreillei latreillei (Leach, 1817)
Finland: Aland (? Tvärminne or Helsingfors) (Nordberg, 1936; Krištofik et
al., 1993, 2002) (mis-identified, see Chernes cimicoides).
CHERNETIDAE
Allochernes powelli (Kew, 1916)
Germany: Hessen, Naturpark Hoher Vogelsberg, Ilbeshausen (Jost, 1982).
Chernes cimicoides (Fabricius, 1793) and Dinocheirus panzeri (C. L. Koch,
1837).
Finland: Aland (? Tvärminne or Helsingfors) (Nordberg 1936). Samples
identified by Nordberg as D. latreillei consist of individuals of two other
species, Chernes cimicoides and Toxochernes (=Dinocheirus) panzeri
(Kaisila, 1949).
Dinocheirus panzeri (C. L. Koch, 1837) (= Toxochernes panzeri (C. L. Koch,
1837).
Austria: Dornbach bei Wien, in einem Schwalbennest (Beier, 1929a); nest
of Hirundo rustica (Beier, 1948).
Progne subis (Linnaeus, 1758) + Passer domesticus (Linnaeus, 1758)
CHELIFERIDAE
Chelifer sp.
USA: Wisconsin, near Madison, 26-X-1962, 15 adults in a Purple Martin
house occupied by House Sparrows (Thompson, 1966).
Riparia riparia Linnaeus, 1758 (= bank swallows: Muchmore, 1981).
CHERNETIDAE
Lamprochernes sp.
Slovakia: Chotín, 5 February 1990, 1 deutonymph (Krištofik et al., 1994).
LARCIDAE
Larca notha Hoff, 1961
Canada: Saskatchewan, Val Marie, 10-VI-1955, J. R. Vockeroth leg.,
1 deutonymph, 3 tritonymphs, 3�, 2� [CNCI], from the nest
(Muchmore, 1981).
NEOBISIIDAE
Neobisium sp.
Slovakia: (Fenda et al., 1998).
Undetermined sp. (= “swallow's nest”: Hoff, 1957).
CHEIRIDIIDAE
Cheiridium museorum (Leach, 1817)
United Kingdom: Wales, Ceredigion, in two swallow nests (Chater, 1989).
572
p. tuRienZo et al.
CHERNETIDAE
Tejachernes stercoreus (Turk, 1949)
USA: Texas, Uvalde County, Frio Cave, from swallow nest constructed
from mud at cave entrance (Hoff, 1957).
ICTERIDAE
Psarocolius decumanus (Pallas, 1769)
CHERNETIDAE
Rhopalochernes sp.
Brazil: Amazonas, Municipios Silves & Itacotiara, Mil Madereira Itacotiara
Ltda., 3º00’S, 58º40’W, VII/XII-2000, 4 nymphs, 9 adults in nest # 59,
21 nymphs, 21 adults in nest # 60, 84 nymphs, 19 adults in nest # 61,
17 nymphs, 23 adults in nest # 62, 13 nymphs, 18 adults in nest # 63,
35 nymphs, 13 adults in nest # 64, 7 nymphs, 5 adults in nest # 65 of a
total of 7 examined nests [Colony 2] (Torres, 2001).
LANIIDAE
Lanius collaris Linnaeus, 1766
WITHIIDAE
Stenowithius bayoni (Ellingsen, 1910)
South Africa: Cape Province, King William’s Town Div., Pirie, in the nest
of the Fiscal Shrike (Ellingsen, 1912; Beier, 1948).
Lanius collurio collurio Linnaeus, 1758
CHELIFERIDAE
Dactylochelifer latreillei latreillei (Leach, 1817)
Slovakia: Vysoká pri Morave, 2-VIII-1993, 1�; Lehnice, 15-VII-1993,
1�; Gabčíkovo, 19-VII-1994, 1 tritonymph, 3�, 1�; Ipelské
Predmostie, 21-VII-1994, 1�, 1� (Krištofik et al., 2002).
NEOBISIIDAE
Neobisium (Neobisium) sylvaticum (C. L. Koch, 1835)
Slovakia: Očová, 6-VII-1993, 1 tritonymph, 1� (Krištofik et al., 2002).
Lanius minor minor Gmelin, 1788
CHERNETIDAE
Chernes cimicoides (Fabricius, 1793)
Slovakia: Detva, settlement of Latok, 19-VII-1996, 1� (Krištofik et al.,
2002).
Dinocheirus panzeri (C. L. Koch, 1837)
Slovakia: Holice, 5-VII-1994, 3 tritonymphs, 1�, 2� (Krištofik et al.,
2002).
LARIDAE
Larus delawarensis Ord, 1815
PSEUDOSCORPIONES, undetermined species
pseudoscoRpions in biRds’ nests
573
Canada: Lake Superior, Black Bay, Granite Island, 48°43’N, 88°29’W, 9-V1983 through 29-VI-1983, 5 specimens out of a total of 40 collected gull
nests (Ryder & Freitag, 1974).
Larus ridibundus Linnaeus, 1766
CHELIFERIDAE
Chelifer cancroides cancroides (Linnaeus, 1758)
Sweden: Lake Takern, Östergötland (Lohmander, 1939; Beier, 1948).
MEROPIDAE
Merops apiaster Linnaeus, 1758
CHERNETIDAE
Chernes hahnii (C. L. Koch, 1839)
Slovakia: Chotín, 48º11’N, 18º14’E, 22-VIII-1991, 1� in one of 9 examined nests (Krištofik et al., 1996).
Lamprochernes nodosus (Schrank, 1761)
Slovakia: Jarovce, 47º14’N, 17º06’W, 16-VIII-1993, 1� in one of 21 examined nests (Krištofik et al., 1996).
MOTACILLIDAE
Anthus trivialis (Linnaeus, 1758)
NEOBISIIDAE
Neobisium sp.
Slovakia: (Fenda et al., 1998).
Motacilla alba Linnaeus, 1758
CHEIRIDIIDAE
Cheiridium museorum (Leach, 1817)
Slovakia (and/or Austria-Czech
Krumpalova, 2010).
Republic):
(Christophoryova
&
Republic):
(Christophoryova
&
MUSCICAPIDAE
Ficedula albicollis (Temminck, 1815)
CHTHONIIDAE
Chthonius sp.
Slovakia: (Fenda et al., 1998).
NEOBISIIDAE
Neobisium sp.
Slovakia: (Fenda et al., 1998).
PSEUDOSCORPIONES
Slovakia (and/or Austria-Czech
Krumpalova, 2010).
Ficedula hypoleuca (Pallas, 1764)
CHERNETIDAE
Allochernes wideri (C. L. Koch, 1843)
574
p. tuRienZo et al.
Finland: Korppoo (V), in a nesting box, where Muscicapa hypoleuca had
been nesting previously, but where bats were living when the nest was
investigated (Kaisila, 1949).
NECTARINIIDAE
Anthreptes malacensis (Scopoli, 1786)
GEOGARyPIDAE
Geogarypus longidigitatus (Rainbow, 1897) (= Geogarypus elegans audyi
Beier, 1952: Beier, 1963a).
Malaysia: Selangor, Rantau Panjang, 8 km N of Klang, 7-VI-1961, 1 deutonymph, in nest 178 (Beier, 1963a).
PANURIDAE
Panurus biarmicus (Linnaeus, 1758)
CHERNETIDAE
Lamprochernes nodosus (Schrank, 1803)
Italy: Po River delta, Argenta, 44°41’N, 11°50’E, 1�, 1�; Campotto,
44°42’N, 11°50’E, 1� (Krištofik et al., 2007).
PARIDAE
Parus caeruleus (Linnaeus, 1758)
CHERNETIDAE
Allochernes wideri (C. L. Koch, 1843)
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1963, 1983).
Chernes hahnii (C. L. Koch, 1839)
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1983).
Parus major Linnaeus, 1758
CHERNETIDAE
Allochernes wideri (C. L. Koch, 1843)
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1983).
Dinocheirus panzeri (C. L. Koch, 1837)
England: Churchdown, Gloucestershire (George, 1961; Jones, 1975, 1978).
CHTHONIIDAE
Mundochthonius styriacus Beier, 1971
Austria: Styria, Pöls bei Zwaring, 20 km S of Graz, in nest 3.5 m above
ground (Beier, 1971; Jost, 1982).
Parus montanus (Conrad von Baldenstein, 1827)
CHELIFERIDAE
Dactylochelifer latreillei latreillei (Leach, 1817)
Finland: Aland (? Tvärminne or Helsingfors) (Nordberg, 1936; Krištofik
et al., 1993) (mis-identified, see Chernes cimicoides).
CHERNETIDAE
Chernes cimicoides (Fabricius, 1793) and Dinocheirus panzeri (C. L. Koch,
1837)
pseudoscoRpions in biRds’ nests
575
Finland: Aland (? Tvärminne or Helsingfors) (Nordberg, 1936). Samples
identified by Nordberg as D. latreillei consist of individuals of two other
species, Chernes cimicoides and Toxochernes (= Dinocheirus) panzeri
(Kaisila, 1949).
PASSERIDAE
Passer domesticus (Linnaeus, 1758)
CHEIRIDIIDAE
Apocheiridium indicum Murthy & Ananthakrishnan, 1977
India: West Bengal, 1984-1985, from a total of 192 nests (Bhattacharyya,
1990a, b, 1999).
Cheiridium museorum (Leach, 1817)
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1963, 1983).
England: Kirton-in-Kindsey, Lincolnshire, in deserted sparrow nest
(Pickard-Cambridge, 1892); Churchdown, Gloucestershire; Woodwalton
Fen N.N.R., Hunts. (Jones, 1975, 1978).
Finland: Aland (? Tvärminne or Helsingfors) (Nordberg, 1936).
India: West Bengal, 1984-1985, from a total of 192 nests (Bhattacharyya,
1990a, 1999).
Switzerland: (Lessert, 1911; Beier, 1948).
CHELIFERIDAE
Chelifer cancroides cancroides (Linnaeus, 1758)
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1963, 1983).
France: Haute-Savoie, Bossy-sur-Frangy, 450m, nest in farm stable, VI1969, leg. J. Steffen, 2�, 7�, 6 tritonymphs, 1 deutonymph, det. V.
Mahnert.
Sweden: (Lohmander, 1939; Beier, 1948).
Switzerland: (Lessert, 1911; Beier, 1948); Geneva City, Natural History
Museum, 2�, 2�, 1983/1993, from nests, det. V. Mahnert.
Chelifer sp.
India: West Bengal, 1984-1985, from a total of 192 nests (Bhattacharyya,
1990a, 1999).
CHERNETIDAE
Chernes sp.
India: West Bengal, 1984-1985, from a total of 192 nests (Bhattacharyya,
1990a, 1999).
Pselaphochernes scorpioides (Hrmann, 1804)
Germany: “Ostdeutschland”, in nest of house sparrow (Drogla & Lippold,
2004).
CHTHONIIDAE
Chthonius (Chthonius) rhodochelatus Hadzi, 1933
France: Haute-Savoie, Bossy-sur-Frangy, 450 m, in nest fallen from a tree,
22-X-1972, leg. J. Steffen, 1�, det. V. Mahnert.
Tyrannochthonius heterodentatus Beier, 1930 (= Tyrannochthonius madrasensis
Murthy, 1961: Bhattacharyya, 1990a, 1999).
576
p. tuRienZo et al.
India: West Bengal, 1984-1985, from a total of 192 nests (Bhattacharyya,
1990a, 1999).
OLPIIDAE
Indolpium loyolae (Murthy, 1961) (= Minniza loyolae: Bhattacharyya, 1990a,
1999).
India: West Bengal, 1984-1985, from a total of 192 nests (Bhattacharyya,
1990a, 1999).
Passer montanus (Linnaeus, 1758)
CHERNETIDAE
Allochernes wideri (C. L. Koch, 1843)
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1963, 1983).
Dendrochernes cyrneus (L. Koch, 1873)
Slovakia (and/or Austria-Czech Republic): (Christophoryova
Krunmpalova, 2010).
&
Prunella modularis (Linnaeus, 1758)
CHEIRIDIIDAE
Cheiridium museorum (Leach, 1817)
Austria: Tirol, Ötztaler Alpen, Obergurgl, 2000 m (Thaler, 1979).
PASSERIFORMES, undetermined genera and species
ATEMNIDAE
Paratemnoides nidificator (Balzan, 1888) (= Paratemnoides minor (Balzan):
Torres, 2001)
Brazil: Amazonas: Municipios Silves & Itacotiara, Mil Madereira Itacotiara
Ltda., 3º00’S, 58º40’W, VII/XII-2000, 1 nymph in nest # 4 (Torres,
2001).
CHERNETIDAE
Allochernes wideri (C. L. Koch, 1843)
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1983).
Allochernes powelli (Kew, 1916)
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1983).
Dinocheirus panzeri (C. L. Koch, 1837)
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1983).
Chernes hahnii (C. L. Koch, 1839)
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1983).
CHELIFERIDAE
Dactylochelifer latreillei latreillei (Leach, 1817)
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1983).
CHTHONIIDAE
Chthonius (Ephippiochthonius) tetrachelatus (Preyssler, 1790)
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1983).
Chthonius (Ephippochthonius) fuscimanus Simon, 1900 (= Chthonius (E.) austriacus: Ressl, 1983).
pseudoscoRpions in biRds’ nests
577
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1983).
Pseudochthonius homodentatus Chamberlin, 1929
Brazil: Amazonas, Municipios Silves & Itacotiara, Mil Madereira Itacotiara
Ltda., 3º00’S, 58º40’W, VII/XII-2000, 1� in nest # 20 (Torres, 2001).
SyARINIDAE
Ideobisium sp.
Brazil: Amazonas, Municipios Silves & Itacotiara, Mil Madereira Itacotiara
Ltda., 3º00’S, 58º40’W, VII/XII-2000, 1 nymph, 1� in nest # 20
(Torres, 2001).
PHALACROCORACIDAE
Phalacrocorax aristotelis (Linnaeus, 1761)
CHELIFERIDAE
Rhacochelifer maculatus (L. Koch, 1873)
France: Corsica, Îles Lavezzi, Rattino, VI.1983, leg. C. Guiguen, 2�, det.
V. Mahnert.
OLPIIDAE
Calocheiridius olivieri (Simon, 1879)
France: Corsica, Îles Lavezzi, Rattino, VI.1983, leg. C. Guiguen, 2�, 2�,
det. V. Mahnert.
PHASIANIDAE
Gallus gallus (Linnaeus, 1758) (= Gallus domesticus: Ressl, 1963).
CHEIRIDIIDAE
Cheiridium museorum (Leach, 1817)
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1963, 1983).
United Kingdom: Wales, Ceredigion, in hen’s nests of straw in an outbuilding (Chater, 1989).
CHELIFERIDAE
Chelifer cancroides cancroides (Linnaeus, 1758)
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1963, 1983).
USA: Illinois (Hoff 1944); Colorado: Weld County, 2�, east of Latham
Reservoir, SE of Greeley, in chicken house (Hoff, 1961).
Hansenius torulosus (Tullgren, 1907) (= Chelifer kewi: Ellingsen 1912).
South Africa: Cape Province: in a hen-house (Ellingsen, 1912).
CHERNETIDAE
Allochernes wideri (C. L. Koch, 1843)
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1963, 1983).
Dinocheirus panzeri (C. L. Koch, 1837)
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1963; 1983).
Dinocheirus sicarius Chamberlin, 1952
USA: Oregon, Lane Co., 5 mi N of Elmira (130 m), litter-dung of chicken
house (Benedict & Malcolm, 1982).
578
p. tuRienZo et al.
Lamprochernes chyzeri (Tömösvary, 1882)
Switzerland: canton of Geneva, Avully, in chicken dung, 6-IX-1966, leg. Cl.
Besuchet, 1�, 1 trito-, 1 deutonymph, det. V. Mahnert.
Lamprochernes savignyi (Simon, 1881)
New Zealand: chicken coops (Beier, 1976).
South Africa: Medunsa, 30 km NW Pretoria, VI.1999, leg. Green, 1�, 1�,
in chicken litter, det. V. Mahnert.
LARCIDAE
Larca lata (Hansen, 1884)
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1983).
WITHIIDAE
Withius indicus Murthy & Ananthakrishnan, 1977
India: Tamil Nadu, in chicken house (Murthy & Ananthakrishnan, 1977).
Withius kaestneri (Vachon, 1937)
South Africa: Cape Province, Albany Distr., 33°32’S, 26°34’W, 10.V.1979
D. Gautier leg., 12�, 6�, in chicken dung, det. V. Mahnert (published by
Hulley, 1983); in manure samples from poultry farms in Albany Distr.
(33°32’S, 26°34’E), Bathurst Distr. (33°34’S, 26°43’S), Ciskei
(33°24’E, 27°08’E), East London (32°53’S, 27°53’E), and Port
Elizabeth (33°54’S, 25°09’E) (Hulley, 1983).
Withius piger (Simon, 1878) (= Withius subruber: Beier, 1976; = Whitius piger:
Pinto et al., 2005).
Brazil: Rio Grande do Sul: Pelotas, in poultry farm (Pinto et al., 2005).
New Zealand: Auckland, Kumeu, in chicken manure, sample # 75/144
(Beier, 1976).
PSEUDOSCORPIONES
USA: California, Sonomo County, in poultry manure (Peck & Anderson,
1969); California, Riverside County, in caged layer poultry manure
(Wills et al., 1990); Florida, Putnam Co., in poultry manure (Propp &
Morgan, 1985).
Polyprectrom inopinatum (Rothschild, 1903)
CHEIRIDIIDAE
Apocheiridium minutissimum Beier, 1964
Malaysia: Pahang, Fraser’s Hill, 26-III-1957, 1� (torso) paratype, Ser. No.
M-02117, Band No. H. 87082, in nest of Rothchild’s Peacock Pheasant
(Beier, 1964).
PICIDAE
Colaptes auratus cafer (Gmelin, 1788) (= Colaptes cafer: Hoff, 1961).
CHERNETIDAE
Acuminochernes tacitus Hoff, 1961
USA: Colorado, Larimer County, 2.5 miles east of Fort Collins, at about
5000 feet in elevation (Hoff, 1961).
pseudoscoRpions in biRds’ nests
579
Dendrocopus medius (Linnaeus, 1758)
CHERNETIDAE
Allochernes wideri (C. L. Koch, 1843)
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1963, 1983).
Dryocopus martius (Linnaeus, 1758)
CHELIFERIDAE
Chelifer cancroides cancroides (Linnaeus, 1758)
Finland: Aland (? Tvärminne or Helsingfors) (Nordberg, 1936).
Picoides borealis Vieillot, 1809
PSEUDOSCORPIONES, unidentified species
USA: Georgia, Fort Stewart Military Reservation, 1997-1998, 7% of all
cavities or 15% of natural cavities of a total of 42 examined cavities
[20 natural cavities, 22 insert cavities] (Pung et al., 2000).
PROCELLARIIDAE
Calonectris diomedea (Scopoli, 1796)
CHERNETIDAE
Pselaphochernes lacertosus (L. Koch, 1873)
Portugal: Islas Selvagem, Selvagem Grande, 1984, P. D. Gabbutt leg., 7�,
7� 5 tritonymphs, in nests, det. V. Mahnert.
Pachyptila turtur Kuhl, 1820
CHERNETIDAE
Apatochernes antarcticus pterodromae Beier, 1964
New Zealand: Auckland Is. (Ocean I.), Stewart I. (Motunau I.) (Beier,
1976).
Procellaria westlandica (Falla, 1946)
CHERNETIDAE
Apatochernes nestoris Beier, 1962
New Zealand: North I., Bay of Plenty (Whale I.); South I., Westland (Nelson
Creek, Barrytown, Murrays Mistake; Open Bay Is.); Stewart I.
(Bench I.) (Beier, 1976).
Pterodroma externa externa (Salvin, 1875)
CHERNETIDAE
Chelanops skottsbergi (Beier, 1957) (= Stigmachernes skottsbergi: Beier,
1957b).
Chile: Juan Fernandez Is., Masafuera, probably in phagophily within the
birds’ colonies (Beier, 1957b).
Pterodroma macroptera (A. Smith, 1840)
CHERNETIDAE
Apatochernes nestoris Beier, 1962
New Zealand: North I., Bay of Plenty (Whale I.); South I., Westland (Nelson
Creek, Barrytown, Murrays Mistake; Open Bay Is.); Stewart I.
(Bench I.) (Beier, 1976).
580
p. tuRienZo et al.
Pterodroma lessoni (Garnot, 1826)
CHERNETIDAE
Apatochernes antarcticus pterodromae Beier, 1964
New Zealand: Auckland Is (Ocean I.), Stewart I. (Motunau I.) (Beier, 1976).
Puffinus griseus (Gmelin, 1789)
CHERNETIDAE
Apatochernes nestoris Beier, 1962
New Zealand: North I., Bay of Plenty (Whale I.); South I., Westland (Nelson
Creek, Barrytown, Murrays Mistake; Open Bay Is.); Stewart I.
(Bench I.) (Beier, 1976).
PSITTACIDAE
Anodorhynchus hyacinthinus (Latham, 1790)
PSEUDOSCORPIONES, unidentified species
Brazil: Mato Grosso do Sul, Pantanal de Miranda, 1 ex. in a natural nest
(Dutra de Carvalho et al., 2005).
Myiopsitta monachus monachus (Boddaert, 1783)
CHERNETIDAE
Parazaona bucheri Beier, 1967
Argentina: Santa Fe, near Santa Fe City (Turienzo et al., 2008).
Parazaona morenensis (Tullgren, 1908)
Argentina: Santa Fe, near Santa Fe City (Turienzo et al., 2008).
PSEUDOSCORPIONES, undetermined species
Argentina: Buenos Aires, without detailed localities, from a total of 43 beds
of fresh vegetal matter and one communal nest (Aramburú et al., 2009).
Nestor meridionalis (Gmelin, 1788)
CHERNETIDAE
Apatochernes nestoris Beier, 1962
New Zealand: North I., Bay of Plenty (Whale I.); South I., Westland (Nelson
Creek, Barrytown, Murrays Mistake; Open Bay Is.); Stewart I.
(Bench I.) (Beier, 1976).
PyCNONOTIDAE
Iole olivacea Blyth, 1844 (= Microscelis olivacea: Beier, 1963).
CHERNETIDAE
Sundochernes malayanus Beier, 1963
Malaysia: Selangor, Rantau Panjang, 5 mi N of Klang, 28-VI-1961, 1�
holotype [BISHOP 3465], nest 191, B.53365 (Beier, 1963).
GEOGARyPIDAE
Geogarypus albus Beier, 1963
Malaysia: Selangor, Rantau Panjang, 5 mi N of Klang: 22-II-1961, 1�
paratype, nest 69, B.53013; 14-VI-1961, 1 deutonymph, paratype, nest
193, B.53338 (Beier, 1963).
pseudoscoRpions in biRds’ nests
581
Pycnonotus goiavier (Scopoli, 1786)
ATEMNIDAE
Anatemnus orites major Beier, 1963
Malaysia: Selangor: Rantau Panjang, 5 mi N of Klang: 1960, 1�, nest 26;
29-III-1961, 2�, nest 110, B.53105; 19-IV-1961, 1 deutonymph, 1 tritonymph, 1�, nest 146, B.53150 (Beier, 1963).
CHERNETIDAE
Sundochernes malayanus Beier, 1963
Malaysia: Selangor, Rantau Panjang, 5 mi N of Klang, 7-VI-1961, 1 tritonymph paratype, nest 182 (Beier, 1963).
GEOGARyPIDAE
Geogarypus longidigitatus (Rainbow, 1897) (= Geogarypus elegans audyi
Beier, 1952: Beier, 1963).
Malaysia: Selangor, Rantau Panjang, 8 km N of Klang: 11-I-1961, 1�, nest
8; 2-II-1961, 1�, nest 50, B.52939; 5-II-1961, 1 deutonymph, nest 44,
1961-249; 9-II-1961, 2�, nest 64; 22-II-1961, 1 protonymph, nest 56,
B.53011; 22-II-1961, 1� (juvenile), nest 58, B-53014; 1-III-1971, 1
deutonymph, 1 tritonymph, nest 70, B-53046; 15-III-1961, 1 tritonymph,
1�, nest 104, B.53080; 29-III-1961, 1� (gravid), nest 112, B.53104; 13IV-1961, 1 tritonymph, 1�, 1� (gravid), nest 142, B.53136; 7-VI-1961,
1 tritonymph, nest 197; 14-VI-1961, 1�, 3�, nest 190, B.53334; 14-VI1961, 1�, nest 200, B.53342; 14-VI-1961, 1 deutonymph, nest 204,
B.53339; 21-VI-1961, 1�, nest 212, B.53347; 21-VI-1961, 1�, 1�,
nest 199, B.53348; 21-VI-1961, 1�, nest 161, B.53349; 5-VII-1961,
2�, 1�, nest 225, B.53373; 26-VII-1961, 1�, nest 235, B.53417 (Beier,
1963).
REMIZIDAE
Remiz pendulinus (Linnaeus, 1758)
CHEIRIDIIDAE
Cheiridium museorum (Leach, 1817) (= Cheridium museorum: Krištofik et al.,
1995)
Slovakia: Dolny Stal, in nest with eggs (Krištofik et al., 1995).
CHELIFERIDAE
Dactylochelifer latreillei latreillei (Leach, 1817)
Slovakia: Velké Blahovo, 1� in 19 examined nests (Krištofik et al., 1993,
2002).
CHERNETIDAE
Lamprochernes nodosus (Schrank, 1803) (= Laprochermes nodosus: Krištofik
et al., 1995)
Slovakia: Jakubor, in nest with chicks (Krištofik et al., 1995).
SITTIDAE
Sitta europeae Linnaeus, 1758
582
p. tuRienZo et al.
CHERNETIDAE
Allochernes wideri (C. L. Koch, 1843)
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1983).
STRIGIDAE
Aegolius funereus (Linnaeus, 1758)
CHELIFERIDAE
Chelifer cancroides cancroides (Linnaeus, 1758)
Slovakia: Chernovrší, area of Pec, 27-VII-1993, 1�; Škamranka, area of
Postřekov, 27-VII-1993, 1 tritonymph (Krištofik et al., 2003).
CHERNETIDAE
Chernes cimicoides (Fabricius, 1793)
Slovakia: Velká Skála, area of Postřekov, 27-VII-1993, 1� 1� (Krištofik et
al., 2003).
Dinocheirus panzeri (C. L. Koch, 1837)
Slovakia: 2� (Krištofik et al., 2003).
Otus asio (Linnaeus, 1758)
CHERNETIDAE
Acuminochernes crassopalpus (Hoff, 1945)
USA: New york, Siracuse, Oakwood Cemetery, 119 ex. in the material
extracted from one nest (Philips & Dindal, 1990).
Dinocheirus panzeri (C. L. Koch, 1837)
Netherlands: Zuid-Holland, Meyendel, uit nest ransuil (Tooren, 2005).
Otus scops (Linnaeus, 1758)
CHERNETIDAE
Allochernes wideri (C. L. Koch, 1843)
Slovakia (and/or Austria-Czech
Krumpalova, 2010).
Strix aluco Linnaeus, 1758
CHERNETIDAE
Allochernes wideri (C. L. Koch, 1843)
Slovakia (and/or Austria-Czech
Krumpalova, 2010).
Republic):
(Christophoryova
&
Republic):
(Christophoryova
&
Strigidae sp.
CHERNETIDAE
Dinocheirus panzeri (C. L. Koch, 1837) (= Chelifer panzeri: Kew, 1911; =
Toxochernes panzeri: Jones, 1978).
England: (Kew, 1911; Beier, 1948); Wickham, Kent, in an owl’s nest (Jones,
1975, 1978).
STURNIDAE
Sturnus vulgaris Linnaeus, 1758
pseudoscoRpions in biRds’ nests
583
CHEIRIDIIDAE
Cheiridium museorum (Leach, 1817)
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1983).
Switzerland: (Lessert, 1911; Beier, 1932, 1948).
CHELIFERIDAE
Chelifer cancroides cancroides (Linnaeus, 1758)
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1983).
Finland: Aland (? Tvärminne or Helsingfors) (Nordberg, 1936).
Italy: S. Vito (Modena), Emilia, nei nidi de storni (Beier, 1929b, 1948).
New Zealand: Kaikoura, Wharekiri, Donegals Flat (Beier, 1976).
Sweden: (Lohmander, 1939; Beier, 1948).
CHERNETIDAE
Allochernes wideri (C. L. Koch, 1843)
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1983).
Chernes hahnii (C. L. Koch, 1839)
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1983).
Dinocheirus panzeri (C. L. Koch, 1837) (= Chelifer panzeri: Kew, 1911; =
Toxochernes panzeri: Beier, 1948; Jones, 1975).
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1983).
England: (Kew, 1911; Beier 1948; Jones, 1975).
Sweden: (Lohmander, 1939; Beier, 1948).
NEOBISIIDAE
Neobisium (Neobsium) sylvaticum (C. L. Koch, 1835)
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1963).
Note: The record of this species is apparently due to a typographical error and
has been implicitly corrected by Ressl (1983); it concerns nests of Corvus
corone.
SyLVIIDAE
Acrocephalus palustris (Bechstein, 1798)
NEOBISIIDAE
Neobisium (Neobisium) inaequale Chamberlin, 1930
Slovakia: Malacky-Vinohrádok, 12-VI-2001, 1�, in nest; 17-VI-2001, 1�,
in nest (Krištofik et al., 2005).
Phylloscopus collybita (Vieillot, 1817)
NEOBISIIDAE
Neobisium (Neobisium) sylvaticum (C. L. Koch, 1835)
Austria: Tirol, Mutters near Innsbruck, 3-VI-1970, W. Gstader leg., 1�, 3�,
V. Mahnert det.
Sylvia atricapilla (Linnaeus, 1758) (= Parus atricapillus: Nordberg, 1936).
CHELIFERIDAE
Chelifer cancroides cancroides (Linnaeus, 1758)
Finland: Aland (? Tvärminne or Helsingfors) (Nordberg, 1936).
584
p. tuRienZo et al.
Dactylochelifer latreillei latreillei (Leach, 1817)
Finland: Aland (? Tvärminne or Helsingfors) (Nordberg, 1936) (mis-identified, see Chernes cimicoides). Chernetidae
CHERNETIDAE
Chernes cimicoides (Fabricius, 1793) and Dinocheirus panzeri (C. L. Koch,
1837)
Finland: Aland (? Tvärminne or Helsingfors) (Nordberg, 1936). The
samples determined as D. latreillei consist of individuals of two other
species, Chernes cimicoides and Toxochernes (= Dinocheirus) panzeri
(Kaisila, 1949).
Sylvia borin (Boddaert, 1783)
CHELIFERIDAE
Chelifer cancroides cancroides (Linnaeus, 1758)
Finland: Aland (? Tvärminne or Helsingfors) (Nordberg, 1936)
TETRAONIDAE
Tetrao urogallus Linnaeus, 1758
NEOBISIIDAE
Neobsium (Neobisium) crassifemoratum (Beier, 1928)
Slovakia: (Fenda et al., 1998).
TROGLODyTIDAE
Troglodytes musculus Neumann, 1823
CHERNETIDAE
Calidiochernes musculi Beier, 1957
Venezuela: Llanos, nest of Troglodytes musculus (“cucaracharo”) (Beier,
1957a).
Troglodytes troglodytes (Linnaeus, 1758)
NEOBISIIDAE
Neobisium (Neobisium) carcinoides (Hermann, 1804)
Slovakia: (Fenda et al., 1998).
Thryothorus leucotis Lafresnaye, 1845
CHTHONIIDAE
Pseudochthonius homodentatus Chamberlin, 1929
Brazil: Amazonas, Municipios Silves & Itacotiara, Mil Madereira Itacotiara
Ltda., 3º00’S, 58º 40’W, VII/XII-2000, 1 nymph in nest # 1 of a total of
8 examined nests (Torres, 2001).
Tyrannochthonius migrans Mahnert, 1979
Brazil: Amazonas, Municipios Silves & Itacotiara, Mil Madereira Itacotiara
Ltda., 3º00’S, 58º40’W, VII/XII-2000, 1 � in nest # 1 of a total of 8 examined nests (Torres, 2001).
IDEORONCIDAE
Xorilbia cf. gracilis (Mahnert, 1985) (= Albiorix sp. cf. gracilis: Torres, 2001).
pseudoscoRpions in biRds’ nests
585
Brazil: Amazonas: Municipios Silves & Itacotiara, Mil Madereira Itacotiara
Ltda., 3º00’S, 58º40’W, VII/XII-2000, 1 nymph in nest # 5 of a total of
8 examined nests (Torres, 2001).
OLPIIDAE
Pachyolpium irmgardae Mahnert, 1979
Brazil: Amazonas, Municipios Silves & Itacotiara, Mil Madereira Itacotiara
Ltda., 3º00’S, 58º40’W, VII/XII-2000, 2 nymphs in one nest of a total of
8 examined nests (Torres, 2001).
TURDIDAE
Phoenicurus ochruros gibraltariensis (Gmelin, 1789)
CHEIRIDIIDAE
Cheiridium museorum (Leach, 1817)
Europe: (Roewer, 1937; Beier, 1948).
Phoenicurus ochruros (Gmelin, 1789) (= black redstart: Ranius & Wilander, 2000).
CHEIRIDIIDAE
Cheiridium museorum (Leach, 1817)
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1963, 1983).
CHELIFERIDAE
Chelifer cancroides cancroides (Linnaeus, 1758)
Austria: Lower Austria, Scheibbs Distr. (Ressl & Beier, 1958; Ressl, 1963,
1983).
France: Haute-Savoie, Bossy-sur-Frangy, VIII-1983, leg. J. Steffen, 2�, det.
V. Mahnert.
Switzerland: canton of Vaud, Commugny, 29-IX-1963, leg. J. Steffen, 4�,
5�, det. V. Mahnert.
LARCIDAE
Larca lata (Hansen, 1884)
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1963, 1965, 1983).
Phoenicurus phoenicurus (Linnaeus, 1758)
CHEIRIDIIDAE
Cheiridium museorum (Leach, 1817)
Finland: Aland (? Tvärminne or Helsingfors) (Nordberg, 1936).
CHERNETIDAE
Allochernes wideri (C. L. Koch, 1843)
Austria: Lower Austria, Scheibbs Distr. (Ressl & Beier, 1958; Ressl, 1963,
1983).
Dinocheirus panzeri (C. L. Koch, 1837) (= Dinocheirus panzeri panzeri: Ressl,
1983; = Dinocheirus panzeri rufeolus: Ressl, 1983).
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1963, 1983).
Sialia sp.
CHERNETIDAE
p. tuRienZo et al.
586
Parachernes squarrosus Hoff, 1949
USA: Illinois, Quincy, in bluebird box (Hoff, 1949).
Turdus merula Linnaeus, 1758
CHTHONIIDAE
Chthonius (Ephippiochthonius) fuscimanus Simon, 1900
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1983).
Chthonius (Chthonius) ischnocheles (Hermann, 1804)
England: Newcastle-upon-Tyne (Jones, 1975).
Tyrannochthonius kermadecensis (Beier, 1976) (= Paraliochthonius kermadecensis: Beier, 1976).
New Zealand: Kermadec Is., in deserted nest (Beier, 1976).
CHELIFERIDAE
Chelifer cancroides cancroides (Linnaeus, 1758)
Finland: Aland (? Tvärminne or Helsingfors) (Nordberg, 1936).
Dactylochelifer latreillei latreillei (Leach, 1817)
Slovakia (and/or Austria-Czech Republic): (Christophoryova
Krumpalova, 2010).
&
CHERNETIDAE
Dinocheirus panzeri (C. L. Koch, 1837) (= Dinocheirus panzeri rufeolus: Ressl,
1983).
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1983).
NEOBISIIDAE
Neobisium (Neobisium) sylvaticum (C. L .Koch, 1835)
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1983).
Turdus philomelas Brehm, 1831
CHELIFERIDAE
Dactylochelifer latreillei latreillei (Leach, 1817)
Slovakia (and/or Austria-Czech Republic):
Krumpalova, 2010).
(Christophoryova
&
Turdus pilaris Linnaeus, 1758
CHEIRIDIIDAE
Cheiridium museorum (Leach, 1817)
Finland: Aland (? Tvärminne or Helsingfors) (Nordberg, 1936).
Turdus sp.
CHTHONIIDAE
Chthonius (Ephippiochthonius) tetrachelatus (Preyssler, 1790)
Czech Republic: Prague, Zavadilova ul., nest of Turdus sp. (Stahlavsky,
2001).
TyRANNIDAE
Sayornis phoebe (Latham, 1790)
CHELIFERIDAE
pseudoscoRpions in biRds’ nests
587
Chelifer cancroides cancroides (Linnaeus, 1758)
North America: (Beier, 1948).
TyTONIDAE
Tyto alba (Scopoli, 1769)
CHELIFERIDAE
Chelifer cancroides cancroides (Linnaeus, 1758)
France: Haute-Savoie, Bossy-sur-Frangy, 450m, among pellets, 1983-1984,
leg. J. Steffen, 5�, 15� 1 trito-, 1 deutonymph, det. V. Mahnert.
Switzerland: Genève, Jussy-Monniaz, “Les Gressevaux”, among barn owl
pellets, 1976, leg. M. Jaussi, 1�, det. V. Mahnert.
LARCIDAE
Larca lata (Hansen, 1884)
Netherlands: Gelderland, Vorchten (Heerde), loft of a church, among barn
owl pellets (Tooren, 2001).
UPUPIDAE
Upupa epops Linnaeus, 1758
CHERNETIDAE
Pselaphochernes scorpioides (Hermann, 1804)
Slovakia: (Christophoryova & Krumpalova, 2010).
UNIDENTIFIED FAMILIES: Nests and guano in colonies of unidentified birds
ATEMNIDAE
Diplotemnus insularis Chamberlin, 1933
Brazil: St. Paul’s Rocks (Arquipélago de São Pedro e São Paulo),
0°55’10’’N/29°20’33’’W, (Atlantic Ocean), in bird nests and on birds
(Chamberlin, 1933; Beier, 1940, 1948).
Note: Three species of sea birds are breeding on the islets of this archipelago: Sula
leucogaster (Pelcaniformes, Sulidae), Anous stolidus and A. minutus
(Charadriiformes, Laridae) (Edwards, 1979).
CHEIRIDIIDAE
Apocheiridium rossicum Redikorzev, 1935
Russia: Ural Mts, Perm, on unidentified bird (Redikorzev, 1938).
Cheiridium museorum (Leach, 1817)
England: (Kew, 1911, Beier, 1948); Belton, Leicestershire, in old bird’s
nest (Robin/house sparrow?) in dry stone wall (Jones, 1975).
Germany: “Ostdeutschland”, in birds’ nests (Drogla & Lippold, 2004)
Slovakia: (Krumpal & Cyprich, 1988; Christophoryova & Krumpalova,
2010).
CHELIFERIDAE
Chelifer cancroides cancroides (Linnaeus, 1758)
Slovakia: (Krumpal & Cyprich, 1988; Christophoryova & Krumpalova,
2010).
588
p. tuRienZo et al.
Dactylochelifer latreillei latreillei (Leach, 1817)
Slovakia: (Krumpal & Cyprich, 1988).
Paisochelifer callus (Hoff, 1945)
USA: eastern and central USA, in birds’ nests (Muchmore, 1990).
CHERNETIDAE
Allochernes wideri (C. L. Koch, 1843)
Austria: (Beier, 1932; Beier, 1948); Lower Austria, Scheibbs Distr. (Ressel,
1963).
Germany: in birds’ nests (Schenkel, 1928); Bayern, Aschaffenburg, “in
Nestern von Höhlenbrütern” (Helversen, 1966); “Ostdeutschland”, in
birds’ nests (Drogla & Lippold, 2004).
Slovakia: (Krumpal & Cyprich, 1988; Christophoryova & Krumpalova,
2010).
Sweden: Östergötland: in 40.9% of hollow oaks with bird nests (Ranius &
Wilander, 2000).
Chernes cimicoides (Fabricius, 1793)
Slovakia: (Krumpal & Cyprich, 1988).
Chernes hahnii (L. Koch, 1873)
Slovakia: (Krumpal & Cyprich, 1988; Christophoryova & Krumpalova,
2010).
Chernes vicinus (Beier, 1932) (= Allochernes vicinus: Krumpal & Cyprich,
1988).
Slovakia: (Krumpal & Cyprich, 1988).
Chernes sp.
England: frequently in nests, often in very large numbers (Woodroffe,
1953).
Slovakia: (Krumpal & Cyprich, 1988).
Dendrochernes cyrneus (L. Koch, 1878)
Slovakia: (Krumpal & Cyprich, 1988; Christophoryova & Krumpalova,
2010).
Dinocheirus panzeri (C. L. Koch, 1937)
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1963).
Netherlands: Zuid-Holland, Meyendel, uit divers nest-materiaal (Tooren,
2005).
Slovakia: (Krumpal & Cyprich, 1988).
Hesperochernes montanus Chamberlin, 1935
USA: Montana, Girds Creek, Ravalli County, in a bird’s nest (Chamberlin,
1935).
Lamprochernes chyzeri (Tömösvary, 1882)
Slovakia: (Krumpal & Cyprich, 1988; Christophoryova & Krumpalova,
2010).
Lamprochernes nodosus (Schrank, 1761)
Slovakia: (Krumpal & Cyprich, 1988; Christophoryova & Krumpalova,
2010).
pseudoscoRpions in biRds’ nests
589
Neochelanops patagonicus (Tullgren, 1900)
Argentina: Santiago del Estero, Añ atuya, in birds’ nests, identified by Feio
[MNRJ] (Mazza, 1943).
Note: This locality is mentioned in Feio (1945) as “Añatinga”, an error
repeated by Ceballos & Rosso de Ferradás (2008).
Opsochernes carbophilus Beier, 1966
New Zealand: Occasionally in bird nests (Beier, 1976).
Parachernes cf. argentinus Beier, 1967
Argentina: [locality not specified], in birds’ nests [not specified] (Turienzo
et al., 2008).
Pselaphochernes scorpioides (Hermann, 1804)
Austria: (Beier, 1948).
CHTHONIIDAE
Chthonius (Ephippiochthonius) tetrachelatus (Preyssler, 1790)
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1963).
Slovakia: (Krumpal & Cyprich, 1988).
Chthonius sp.
Slovakia: (Krumpal & Cyprich, 1988).
GARyPIDAE
Garypus titanius Beier, 1961
Ascension Island: Bos’nbird Isle, “im Vogelguano der Nistkolonien” [in
birds’s guano of the nesting colony], 13 species of sea birds (Beier,
1961).
GEOGARyPIDAE
Geogarypus albus Beier, 1963
Malaysia: Selangor, Rantau Panjang, 5 mi N of Klang, 1960, � holotype,
nest # 65, from same locality, 1960, � allotype [BISHOP], nest # 62,
1960, 1� paratype, nest # 26, 1960, 1� paratype, nest # 111 (Beier,
1963).
Geogarypus longidigitatus (Rainbow, 1897) (= Geogarypus elegans audyi
Beier, 1952: Beier 1963).
Malaysia: Selangor, Rantau Panjang, 8 km N of Klang, 1960, 1 deutonymph, 1 tritonymph, nest # 231; 1960, 1 tritonymph, 3��, nest # 255;
1960, 1 deutonymph, nest # 257; 1960, 1 deutonymph, nest # 256; 1960,
1� (gravid), nest # 270; 1960, 1 tritonymph, nest # 274; 13-VII-1960,
1�, nest # 282; 1960, 1 tritonymph, nest # 290; 5-X-1960, 1 protonymph, nest # 317 (Beier, 1963).
LARCIDAE
Larca lata (Hansen, 1884)
Slovakia (and/or Austria-Czech Republic): (Christophoryova &
Krumpalova, 2010).
Sweden: province of Östergötland, in hollow oaks with bird nests (Ranius &
Wilander, 2000).
590
p. tuRienZo et al.
NEOBISIIDAE
Neobisium (Neobisium) carcinoides (Hermann, 1804) (= Neobisium muscorum:
Chater, 1989; Krumpal & Cyprich, 1988).
Slovakia: (Krumpal & Cyprich, 1988; Fenda et al., 1998: nest type
Motacilla; Christophoryova & Krumpalova, 2010).
United Kingdom: Wales, Ceredigion, found in birds' nests on the ground
(Chater, 1989).
Neobisium (Neobisium) crassifemoratum Beier, 1928
Slovakia: (Krumpal & Cyprich, 1988).
Neobisium (Neobisium) inaequale Chamberlin, 1930
Slovakia: (Krumpal & Cyprich, 1988; Fenda et al., 1998: nest type
Motacilla).
Neobisium (Neobisium) sylvaticum (C. L. Koch, 1843)
Austria: Lower Austria, Scheibbs Distr. (Ressl, 1963).
OLPIIDAE
Xenolpium pacificum pacificum (With, 1907)
New Zealand: (Beier, 1976).
PSEUDOGARyPINIDAE
Pseudogarypinus frontalis (Banks, 1909)
USA: western United States, in birds’ nests (Muchmore, 1990).
WITHIIDAE
Dolichowithius argentinus Beier, 1959
Argentina: [province, locality not specified] in birds’ nests [not specified]
(Turienzo et al., 2008).
Dolichowithius cf. argentinus Beier, 1959
Argentina: [province, locality not specified] in birds’ nests [not specified]
(Turienzo et al., 2008).
Stenowithius duffeyi Beier, 1961
Ascension Island: Bos’nbird Isle, “auf dem Vogelguano der Nistkolonien”
[on the birds’ guano in the nesting colony”] (13 species of sea birds)
(Beier, 1961).
PSEUDOSCORPIONES, undetermined species
Argentina: Salta, Dto Santa Bárbara, in the proximities of the river Lavayen,
together with Psammolestes coreodes (Bergroth, 1911) [Hemiptera:
Reduviidae: Triatominae] (Mazza, 1943).
India: West Bengal, 788 adult pseudoscorpions in 1536 nests of 8 species of
birds [Streptopelia chinensis (Scopoli); Acridotheres tristis (Linnaeus);
Sturnus contra Linnaeus; Corvus splendens Vieillot; Prinia socialis
Sykes; Ploceus philippinus (Linnaeus); Passer domesticus (Linnaeus)]
(Bhattacharyya, 1990).
USA: Florida, in poultry manure (Propp & Morgan, 1985).
pseudoscoRpions in biRds’ nests
591
DISCUSSION
14 families of pseudoscorpions with 63 genera and 85 species are recorded from
birds’ nests. Chernetidae are dominant with 22 genera and 35 species, these numbers
represent about 19% of the described genera and 5% of the described species in this
family (Harvey, 2009). It is followed by Chthoniidae (5 genera/subgenera and 9
species) (= about 1.3% of the described species), Cheliferidae (5 genera and 6
species/subspecies) (=about 8% of the decribed genera and about 2% of the described
species!), Olpiidae (4 genera and 4 species), Withiidae (4 genera and 7 species)
(= about 11% of the described genera and 4.5% of the described species!), Atemnidae
(4 genera and 5 species/subspecies), Cheiridiidae (2 genera and 5 species), Neobisiidae
and Geogarypidae (1 genus and 4 species each), Larcidae (1 genus and 2 species),
Garypidae, Ideoroncidae, Pseudogarypinidae (1 species each), Syarinidae (1 species).
The predominance of Chernetidae might be explained partly by the corticolous biology
of many of its species.
45 families of birds have been recorded, represented by a total of 98 identified
species and a certain number of unidentified taxa, a very small part compared to the
number of species and families recognized world-wide (about 142 families and 9700
species). The ranking is headed by Procellariidae, Turdidae (7 spp.) each, Furnariidae
(6 or 5 spp.), Hirundinidae (5 spp.), Picidae, Sylviidae (4 spp. each), Strigidae (4 spp.),
Anatidae, Cacatuidae, Corvidae, Estrildidae, Laniidae, Paridae, Psittacidae,
Troglodytidae (3 spp. each), Anatidae, Passeridae (3 spp. each), Apodidae, Ardeidae,
Cisticolidae, Columbidae, Laridae, Motacillidae, Muscicapidae, Phasianidae,
Pycnonotidae (2 spp. each), Accipitridae, Cinclidae, Climacteridae, Coraciidae,
Emberizidae, Formicariidae, Fringillidae, Halcyonidae, Icteridae, Meropidae,
Nectariidae, Panuridae, Phalacrocoracidae, Remizidae, Sittidae, Sturniidae, Tetraonidae, Tyrannidae, Tytonidae, and Upupidae (1 sp. each). Easy accessibility of nests
for human observers explains probably the predominance of certain families.
Little information can be gathered from listing pseudoscorpions and birds’nests
per country. That reflects only the interest of collectors in the mentioned countries, or
particular interest in other arthropod groups present in birds’ nests. This fact is emphasized by the leading position of Austria (17 bird species studied, but compare this to
the over 200 bird species breeding in Austria!) which is almost exclusevily due to the
activity of one single person, Franz Ressl from the Scheibbs District, Lower Austria.
The prominence of Slovakia is due to a research program initiated mainly by J.
Kristofik, and the many data from Malaysia are the results of the collecting by H. E.
McClure and Lim Boo-Liat (Beier, 1963, 1964). From some European countries we do
not have any record (e.g. Spain, Belgium, Greece, etc.). Several countries in tropical
areas (i.e. Brazil, Venezuela, India, Uganda) have very few records of pseudoscorpions
from birds’ nests. A similar situation was presented for the insect fauna in birds’ nests
from the Neotropical Region (Di Iorio & Turienzo, 2009). Furthermore, it can be
pointed out that indications of origin in older collections were sometimes incomplete,
without any information on habitats (see Parazaona bucheri and Parachernes argentinus under Phacellodomus sp.).
The presence of pseudoscorpions in birds’ nests is certainly favoured by good
or optimal living conditions offered by this particular niche: abundance of potential
592
p. tuRienZo et al.
prey (mites, larvae of various insects, e.g. fleas and dipteras, in the nests, biting lice
(Mallophaga) directly on the birds), and also microclimatic conditions without large
variation in humidity, temperature and light (nests in hollow trees) (see Krumpal &
Cyprich, 1988 and Christophoryova & Krumpalova, 2010).
It is evident that the building site of nests highly influences the presence of one
or another pseudoscorpion family. Nests built on the ground will be visited mainly by
soil-dwelling genera and species, e.g. Chthoniidae (genera Chthonius, Tyrannochthonius), Neobisiidae (Neobisium), Olpiidae, Geogarypidae and, to a lesser extent,
Garypidae and Syarinidae or even Withiidae (animals of this family are found in litter
or under stones, but also under the bark of living or fallen trees). Mundochthonius
styriacus (Chthoniidae) and Anthrenochernes stellae (Chernetidae) seem to prefer the
constant conditions of hollow trees and are therefore frequently found in birds’ nests
constructed in this habitat. On the other hand, many species of usually bark-inhabiting
families (Cheiridiidae, Chernetidae, Cheliferidae) are regularly colonizing nests in
trees, but the presence of several genera, e.g. Dactylochelifer (Cheliferidae), Apatochernes, Chelanops, Chernes or Sundochernes (Chernetidae) is common, though probably not very frequent. Biology is explaining the sporadic presence of the tritonymph
of Neobisium sylvaticum in nests in trees: this stage frequently climbs on shrubs and
trees (Beier, 1963b) and arrives accidentally in nests. Some species of Cheiridiidae
(Apocheiridium spp., Cheiridium museorum), Cheliferidae (Chelifer cancroides) and
Chernetidae (Dinocheirus panzeri) are regularly colonizing nests of synanthropic birds
(e.g. Columba livia, Passer domesticus, Gallus gallus) or may be passively dispersed
by other birds (e.g. Larca lata by Phoenicurus ochrurus: Ressl, 1963b, 1965).
The abundance of pseudoscorpions is usually low (particularly in most ground
nests), but may be very high in old nests in the nature, and even more so in nests of
synanthropic bird species.
Only a few species can be considered as (more or less) exclusive colonizers of
tree nests of birds, as far as we can judge from the scarce surveys and collecting data.
These are Diplotemnus insularis Chamberlin (St Paul’s Islands), Hesperochernes montanus (USA) (Beier, 1948), Troglochernes dewae (Australia) (Harvey & Volschenk,
2007), Parazaona bucheri, Parazaona morenensis (Turienzo, Di Iorio & Mahnert, unpublished data) and Calidiochernes musculi (South America) (Beier, 1957). On the
other hand, some common bark-inhabiting species of the genera Victorwithius and
Dolichowithius (Withiidae from South America) have never or very rarely been found
in birds’ nests. Even more surprising is the quasi total absence in nests of some
Atemnidae, e.g. Paratemnoides nidificator (Balzan), P. pallidus (Balzan), and P. elongatus (Banks). These species show evidence of social behaviour and can be found in
huge numbers under the bark of standing or fallen trees.
ACKNOWLEDGEMENTS
The authors express their deep thanks to Dr Mark S. Harvey (Western
Australian Museum, Perth) and to a second, anonymous, reviewer for their linguistic
ameliorations, comments and bibliographic help, they improved considerably the first
version of this list. Sincere thanks are also due to Dr Peter Schwendinger (Muséum
pseudoscoRpions in biRds’ nests
593
d’histoire naturelle, Genève) for his intensive help during the revisional process of this
paper. We are also thankful to Mrs Christelle Mougin (Library, Muséum d’Histoire
naturelle, Geneva) for her efficient help in bibliographic research.
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Paola Turienzo