bishop museum bulletins in cultural and environmental studies

BISHOP MUSEUM 

BULLETINS IN CULTURAL AND 

ENVIRONMENTAL STUDIES 

Marine Bioinvasions of Hawai‘i 

The Introduced and Cryptogenic 

Marine and Estuarine 

Animals and Plants of the Hawaiian Archipelago 

JAMEs T. CArlTon And luCIus G. EldrEdGE 

Bishop Museum Bulletin in Cultural and Environmental Studies 4 

Bishop Museum Press 

Honolulu

Marine Bioinvasions of Hawai‘i

Marine Bioinvasions of Hawai‘i 

The Introduced and Cryptogenic 

Marine and Estuarine 

Animals and Plants of the Hawaiian Archipelago 

James T. Carlton & Lucius G. Eldredge 

Bishop Museum Bulletin in Cultural and Environmental Studies 4 

Bernice P. Bishop Museum, Honolulu 

2009

Cover: Map of the 2000–2002 Pacific shipping routes. It is 24 months of data from the voluntary WMO/NOAA weather 

reports generated automatically from vessels fitted with met stations; taken usually every 6 hours but sometimes less 

frequently. Data from fixed buoys, floating buoys, research vessels were extracted. The data show approximately 

825,000 records for that 24-month period. [courtesy Maritime Safety and Envir o nmental Strategy, Australian Maritime 

Safety Authority] 

Published by 

Bishop Museum Press 

1525 Bernice Street 

Honolulu, Hawai‘i 96817-2704, USA 

Copyright ©2009 Bishop Museum 

All Rights Reserved 

Printed in the United States of America 

ISBN-10: 1-58178-096-6 

ISBN-13: 978-1-58178-096-3 

ISSN 1548-9620

TABLE OF CONTENTS 

INTRODUCTION ......................................................................................................................... 7 

METHODS ..................................................................................................................................... 8 

ACKNOWLEDGMENTS ............................................................................................................. 9 

TABLES 

1. Marine and estuarine introduced and cryptogenic species of the Hawaiian Islands ......... 10 

2. Number of species treated in this monograph and their status .......................................... 20 

3. Vectors transporting nonindigenous marine and estuarine species to the Hawaiian 

Islands (and Acronyms Used in Table 1) ........................................................................... 21 

SYSTEMATIC ACCOUNT OF INTRODUCED AND CRYPTOGENIC SPECIES 

Viruses, Bacteria, and Fungi .......................................................................................... 21 

Dinoflagellata ................................................................................................................. 22 

Ciliophora ....................................................................................................................... 23 

Porifera ........................................................................................................................... 24 

Cnidaria .......................................................................................................................... 32 

Ctenophora ..................................................................................................................... 47 

Nematoda ........................................................................................................................ 47 

Rotifera ........................................................................................................................... 48 

Platyhelminthes .............................................................................................................. 48 

Annelida ......................................................................................................................... 51 

Mollusca ........................................................................................................................ 68 

Crustacea ........................................................................................................................ 90 

Insecta ............................................................................................................................ 116 

Pycnogonida ................................................................................................................... 124 

Phoronida ...................................................................................................................... 127 

Kamptozoa ..................................................................................................................... 128 

Bryozoa ......................................................................................................................... 128 

Echinodermata ............................................................................................................... 134 

Ascidiacea ...................................................................................................................... 135 

Pisces ............................................................................................................................. 142 

Algae .............................................................................................................................. 151 

Anthophyta ..................................................................................................................... 162 

LITERATURE CITED ............................................................................................................... 166 

INDEX ......................................................................................................................................... 196

INTRODUCTION 

We present here the first comprehensive monograph of the introduced and cryptogenic marine and 

estuarine organisms of the Hawaiian Archipelago, from the island of Hawai‘i to Kure and Midway 

Atolls, between the latitudes of 19° N and 29° N. Covered here are protoctists, fungi, invertebrates, 

fish, algae, and flowering plants, inhabiting coastal environments influenced by the sea. The habitats 

we cover are thus the supralittoral zone (supporting “strand” or “maritime” species), the intertidal 

zone, and the sublittoral (subtidal), including non-native parasites, commensals, or other symbionts. 

We treat 490 species, of which 301 are introduced and 117 are cryptogenic (Table 1). These 

and other categories are defined below. The rest of the species (Table 2) are either of unknown establishment, 

species that escaped or were intentionally released but did not establish, species that were 

unintentionally introduced but failed to establish, intercepted species, species that represent doubtful 

or erroneous records, native species previously treated as introduced, waifs, or species that are of 

uncertain presence in marine waters. Brock (1952, 1960) reviewed introduced marine and freshwater 

species, covering 61 taxa in the latter paper. Eldredge (1994) reviewed the history of introductions 

of commercially significant species to the Pacific Islands, including Hawai‘i. 

The marine and estuarine biota of the Hawaiian Islands are derived from a combination of nonhuman 

mediated and human-mediated dispersal processes. “Natural” processes include ocean currents 

transporting larval, juvenile, or adult stages (as plankton or by rafting), and migratory birds on 

long-established flyways (Zimmerman, 1948a; Newman, 1986; Jokiel, 1990; Mueller-Dombois & 

Fosberg, 1998). Human-mediated processes involve a long list of vectors (sensu Carlton & Ruiz, 

2005), the major ones of which for the Archipelago include ships and intentional importations (and 

the hitchhikers associated with such transplantations) (Table 3). 

Drifting organisms began colonizing the aboriginal islands soon after they were formed; this 

process has been on-going for tens of millions of years, leading to the evolution of many endemic 

marine, freshwater, and terrestrial organisms in the Archipelago. As the most isolated islands in the 

world, however, a great many species failed to naturally colonize, and thus assisted passage was 

required to bridge vast expanses of ocean. 

We presume that Polynesian (Kirch, 2002) vessels of the 11th century (if not earlier human colonization 

waves) brought the first hull-fouling (if not bilge and solid ballast) organisms to Hawai‘i. 

We have not systematically attempted here to sift through the paleontological, archeological, or historical 

sedimentological record to identify Polynesian-mediated introductions of marine organisms. 

If and when such species are identified, we would still regard them as introduced species, as part of 

the human-influenced construction of the biota, as do our terrestrial colleagues who discern the 

translocation of distant plants by aboriginal settlers to the Hawaiian flora. Speculation about the role 

of Polynesians in introducing marine or brackish species has not been extensive; we note, however, 

that the freshwater and brackish water snails Tarebia granifera and Melanoides tuberculata, transportable 

in taro roots, are two of a number of candidate species (Cowie, 1998). 

Captain James Cook’s arrival in 1778 marked the beginning of the modern era of vessel-mediated 

invasions in Hawai‘i (Beechert, 1991). By the early 1800s, global vessel traffic visiting the 

Islands had become common. Between 1820 and 1840 alone, more than 1,500 visits to the Port of 

Honolulu were made by whaleships (hailing from all over the world) and more than 400 visits were 

made by Trans-Pacific traders from around the Pacific Rim (Richards, 2000). Rapid interisland 

exchange was facilitated by over 1,100 interisland transits in the same period (Richards, 2000). By 

the 1900s many tens of thousands of vessel visits (for example, Worden, 1980; Beechert, 1991) with 

7

8 

BISHOP MUSEUM BULLETIN IN CULTURAL AND ENVIRONMENTAL STUDIES 4 (2009) 

thousands of species on and in their hulls, and in their solid and water ballast, from the world over, 

had come to the port and harbor systems of the islands that had evolved from the original coastal 

rivers and estuaries. 

In a separate work (Carlton & Eldredge, in prep.) we analyze the temporal history of accidental 

and intentional introductions, the proportional role of vectors, biogeographic tracks, the systematic 

and taxonomy history, and the ecological and societal impacts of the introduced fauna and flora. 

Most species have arrived through international shipping (as fouling and in ballast), and most species 

hail from the Indo-West Pacific, Eastern Pacific, or North Atlantic. The absence of historical surveys 

in the Islands prevents the timely detection of virtually all early invasions: many species first collected 

in the first half of the 20th century were almost certainly introduced in the 1700s and 1800s. 

Thus by 1900, only nine ship-dispersed species had been collected in Hawai‘i, representing fouling, 

boring, and solid ballast introductions (Carlton & Eldredge, in prep.), a number that soberingly 

reflects the nearly complete lack of knowledge of what was living in Pearl Harbor, Honolulu Harbor, 

Kāne‘ohe Bay, or other archipelago estuaries, only a century ago. 

Thus the history of marine invasions in Hawai‘i, perhaps more so than any other region in the 

world, is intimately tied to the history of interested and dedicated biologists, and their visits or residency 

in the Islands. Premier among these (Carlton & Eldredge, in prep.) was Charles Howard Ed - 

mond son, to whom we owe the early 20th century documentation of many presumably 19th century 

invaders (Eldredge, 2007). 

METHODS 

We became interested in the marine bioinvasions of the Hawaiian Islands in the 1970s, and after 

accumulating numerous incidental records, began a systematic study in the 1990s. We studied most 

of the published systematic and biogeographic literature on marine and brackish-water invertebrates, 

algae, flowering plants, and fish of the Hawaiian Islands, corresponded with many colleagues (see 

Acknowledgments), and examined museum collections (referenced at individual species), particularly 

the Bishop Museum (Honolulu) and the National Museum of Natural History (Washington, 

D.C.). Along with numerous colleagues over the years, we conducted field work in Pearl Harbor and 

Kāne‘ohe Bay (species and pertinent dates of collection are noted throughout the text). Commencing 

in 1996, Lu Eldredge, Steven Coles, Scott Godwin, Ralph DeFelice, and their associates (Bishop 

Museum), began a series of port, harbor, estuary, and reef studies through the entire Archipelago, 

revealing a plethora of invasions that had arrived over prior decades. 

We assigned all species (Tables 1 and 2) with which we were concerned to one of the categories 

listed below. We assessed historical, systematic, biogeographic, and other criteria (see Chapman & 

Carlton, 1991, 1994) relative to the potential history of a given species, as detailed in many of the 

individual treatments. Many species that had never been previously considered relative to their biogeographic 

status in the Hawaiian Islands are newly treated in this monograph as introductions or 

cryptogens. In addition to the categories listed below, we relegated 19 species (Table 2) to doubtful 

or erroneous records, natural waifs, or uncertain presence in marine waters. These are shown in the 

“Other” category on Table 2, which also include several species that we concluded were native, 

although previously suggested as introduced. 

INTRODUCED: 

Species that have been transported by human agency, whether intentionally or accidentally, and are 

established in the wild. The following terms are in cluded here: nonindigenous, invasive, adventive, 

acclimatized, exotic, nonnative, immigrant, alien, foreign, naturalized, transplant, transfer, and import. 

Subcategories: 

Establishment Unknown: continued presence and reproduction are unknown, and for which 

subsequent species-specific searches may not have been undertaken. Examples include the 

snails Amphithalamus inclusus, Boonea cincta, Ividella navisa, and Peristichia pedroana. 

Failed: known to have reproduced or grown in the wild but then subsequently died out. 

Examples include the soft coral Dendronephthya sp., the snail Bulla adamsi, and the mussel 

Mytilus galloprovincialis.

Carlton & Eldredge — Marine Bioinvasions of Hawai‘i 

9 

Intercept: associated with fisheries products or fouled vessels brought to Hawai‘i, with the species 

concerned having subsequent access to open ocean waters. Examples include the polychaete 

Boccardia proboscidea, the crab Charybdis helleri, and the alga Sargassum muticum. 

Deleted: one or a few individuals collected more than 50 years ago, and for which subsequent 

searches have not revealed the species. Examples include the isopod Sphaeroma terebrans, 

the mudshrimp Upogebia pugettensis, the crab Carcinus maenas and the sea spider 

Anoplodactylus eroticus. 

Released; Not Established: intentional release but with no subsequent evidence of having 

established a naturally reproducing population in the wild. Examples include the snails 

Haliotis spp. and Trochus niloticus, the shrimp Macrobrachium rosenbergii, and numerous 

marine fish species. 

Escaped; Not Established: species that escaped from culture operations, but with no subsequent 

evidence of having established a naturally reproducing population in the wild. Examples 

include penaeid shrimp, the alga Mazzaella volans, and the alga Macrocystis pyrifera. 

CRYPTOGENIC: 

Species that are not clearly native or introduced (Carlton, 1996a). 

NATIvE: 

Species present prior to human contact. endemic species are found only in Hawai‘i; indigenous 

species are native but not unique to the Islands. 

ACKNOWLEDGMENTS 

We are grateful to a large number of colleagues who have provided advice, assistance, clarification 

of records, reprints, and excavation of obscure literature. These include Isabella Abbott, Shane 

Ahyong, James Allen, Julie Bailey-Brock, Rolando Bastida-Zavala, Michael Berrill, Rudiger Bieler, 

Chip Biernbaum, Barbara Butler, David Ronald Bybee, Stephen Cairns, Dale Calder, Roy Caldwell, 

Debby Carlton, John Chapman, Donald Chandler, C. Alan Child, Eugene Coan, Andrew Cohen, 

Linda Cole, Steven Coles, Rachel Collin, William Cooke, Robert Cowie, Gerald L. Crow, Peter 

Davie, Michael Dawson, Amanda Demopoulos, Matthew Dick, Bert Draper, Ralph DeFelice, 

Ronald Englund, Christer Erséus, Neal Evenhuis, Daphne Fautin, Darryl Felder, Paul Fofonoff, 

William Font, Charles Galt, Jonathan Geller, Lisa-Ann Gershwin, Christopher Glasby, Scott 

Godwin, D. Goodwin, Terry Gosliner, David Greenfield, Danièle Guinot, David Gulko, Eduardo 

Hajdu, Michael Hadfield, S.N. Harkantra, Leslie Harris, Richard Heard, Robert Hershler, Brenden 

Holland, Charlotte Holmquist, Bridget Holohan, John Hooper, John Hoover, Frank Howarth, Samuel 

Ensung Kahng, Regina Kawamoto, Alison Kay, Robert Kinzie, Phyllis Knight-Jones, Alan Kohn, 

Gretchen Lambert, Janet Lamberson, James Lowry, Josh Mackie, Antonio Frias Martins, Karen 

Maruska, George Matsumoto, Linda McCann, Claudia Mills, Anthony Montgomery, Bruce Mundy, 

Todd Newberry, William Newman, Peter Ng, Gordon Nishida, Arne Nygren, Daniel Otte, Brian 

Paavo, Gustav Paulay, Sara Pelleteri, Mary Petersen, Andrew Pitman, Cory Pittman, Richard Pyle, 

John E. Randall, Geoff Read, David Reid, Raul Aguilar Rosas, Gary Rosenberg, Michael Roy, 

Dennis Russell, Katsushi Sakai, Al Samuelson, Marilyn Schotte, Michelle Kelly-Borges Shanks, 

Alison Sherwood, Karla McDermid Smith, Michael Stat, Isabel Stirling, Stefano Taiti, C. Takahashi, 

Harry ten Hove, Thomas Theeger, Wesley Thorsson, Rob van Soest, Roy Tsuda, Catherine Unabia, 

Daniel Wagner, Kerstin Wasson, Les Watling, Judith Winston, Robert Woollacott, Brian Wysor, 

Chela Zabin, and Helmut Zibrowius. Funding for our work was provided by The Charles H. and 

Margaret B. Edmondson Trust to which we are deeply thankful.

10 


Table 1. Marine and estuarine introduced and cryptogenic species of the Hawaiian Islands 

Species Date Vector* Native to Status** 

FUNGI 

Cytospora rhizophorae 1902 R Western Atlantic I 

Etheirophora blepharospora 1902 R Western Atlantic I 

PROTOCTISTA 

Dinoflagellata: Dinophyceae 

Perkinsus marinus 1972 R Atlantic Ocean I 

Ciliophora (ciliates) 

Zoothamnium sp./spp. 1935 SF, BW, R C 

Halofolliculina annulata — SF, R C 

Parafolliculina violaceae — SF, R C 

Metafolliculina nordgardi — SF, R C 

Metafolliculina andrewsi — SF, R C 

Lagotia viridis — SF, R C 

Ascobius simplex — SF, R C 

Eufolliculina lignicola 1961 SB Unknown I 

Mirofolliculina limnoriae 1961 SB Indo-Pacific I 

Cephaloidophora communis 1949 SF Unknown I 

PORIFERA (sponges) 

Cliona sp./Pione sp. 1947 SF, R C 

Tethya ornata 1985 SF C 

Suberites aurantiacus 1902 SF Unknown I 

Halichondria melanadocia 1960s [


11 

Table 1. (continued) 

Species Date Vector Native to Status 

CNIDARIA 

Hydrozoa (hydroids) 

Bougainvillia muscus 1967 SF, BW Atlantic I 

Eudendrium sp. 1972 SF, BW C 

Cladonema radiatum 1972 SF, BW North Atlantic I 

Pennaria disticha 1928 SF, BW Unknown I 

Halecium beani 1943 SF, BW C 

Ventromma halecioides 1999 SF, BW C 

Plumularia floridana 1999 SF, BW C 

Plumularia strictocarpa 1999 SF, BW C 

Plumularia setacea 1972 (1946?) SF, BW C 

Antennella secundaria 1999 SF, BW C 

Halopteris diaphana 1970s SF, BW C 

Halopteris polymorpha 1999 SF, BW C 

Lytocarpia phyteuma 2001 SF, BW C 

Anthohebella parasitica 1999 SF, BW C 

Obelia dichotoma 1972 SF, BW Unknown I 

Obelia bidentata 1946 SF, BW Unknown I 

Clytia latitheca 1999 SF, BW C 

Clytia hemisphaerica 1972 SF, BW C 

Thyroscyphus fruticosus 1973 SF, BW C 

Synthecium megathecum 1972 SF, BW C 

Dynamena crisioides 1972 SF, BW C 

Dynamena cornicina 1972 SF, BW C 

Dynamena quadridentata 2001 SF, BW C 

Sertularella areyi 1999 SF, BW C 

Sertularella tongensis 1999 SF, BW C 

Sertularella diaphana 1972 SF, BW C 

Tridentata loculosa 1972 SF, BW C 

Tridentata hupferi 1972 SF, BW C 

Tridentata turbinata 2001 SF, BW C 

Tridentata distans 2001 SF, BW C 

Tridentata marginata 2001 SF, BW C 

Anthozoa (corals and sea anemones) 

Culicia rachelfitzhardingeae 1983 SF Indo-Pacific I 

Sarcothelia sp. 2000 SF, BW C 

Carijoa riisei 1972 SF, BW Indo-Pacific I 

Diadumene leucolena 1950s SF, BW, R NW Atlantic I 

Diadumene lineata 1999 SF, BW, R Japan I 

Diadumene franciscana 1998 SF, BW Unknown I 

Scyphozoa (jellyfish) 

Aurelia sp. 1953 SF, BW Indo-Pacific I 

Cassiopea andromeda 1950 SF, BW Indo-Pacific I 

Phyllorhiza punctata 1933 SF, BW Indo-Pacific I 

Anomalorhiza shawi 1983 SF, BW Philippines I 

Cubozoa (cubomedusae) 

Carybdea sivickisi 1996 BW Indo-Pacific I 

CTENOPHORA (comb jellyfish) 

Vallicula multiformis 1992 BW Caribbean I

12 




NEMATODA (roundworms) 

Camallanus cotti 1993 R Asia I 

Spirocamallanus istiblenni 1962 R? C 

ROTIFERA (rotifers) 

Brachionus plicatilis 2001 R, BW C 

Colurella adriatica 2001 R, BW C 

Hexarthra oxyuris


13 



ANNELIDA (true worms) 

Polychaeta (polychaetes) 

Malacoceros sp. 1978 SF, BW C 

Minuspio sp. 1970s BW C 

Armandia intermedia 1979 SF, BW C 

Chaetopterus sp. 1960 SF, BW Unknown I 

Branchiomma japonica 1946 SF, BW Japan I 

Sabellastarte spectabilis 1946 SF, BW Philippines I 

Ficopomatus enigmaticus 1937 SF, BW Australia I 

Hydroides brachyacanthus 1939 SF, BW Eastern Pacific I 

Hydroides cruciger 1936 SF, BW Eastern Pacific I 

Hydroides diramphus 1900 SF, BW Western Atlantic? I 

Hydroides elegans 1929 SF, BW Indo-Pacific? I 

Pomatoleios kraussii 1967 SF, BW Indo-Pacific I 

Salmacina tribranchiata 1935 SF, BW Eastern Pacific I 

Serpula watsoni 1936 SF, BW Indo-West Pacific I 

Eulaeospira orientalis 1960s SF, BW C 

Janua pagenstecheri 1960s SF, BW Northeast Atlantic I 

Leodora knightjonesi 1960s SF, BW C 

Neodexiospira foraminosa 1960s SF, BW C 

Neodexiospira nipponica 1960s SF, BW C 

Neodexiospira pseudocorrugata 1960s SF, BW C 

Pileolaria militaris 1960s SF, BW Northeast Atlantic? I 

Pileolaria pseudoclavus 1960s SF, BW C 

Simplicaria pseudomilitaris 1960s SF, BW C 

Spirorbis marioni 1960s SF, BW C 

Vinearia koehleri 1960s SF, BW C 

MOLLUSCA 

Gastropoda (snails) 

Diodora ruppelli 1962 SF, BW Indo-Pacific I 

Tarebia granifera 1856 R Indo-Pacific I 

Melanoides tuberculata 1994 R? Asia-Africa I 

Pyrgophorus coronatus 1998 BW? Caribbean I 

Vermetus alii

14 




MOLLUSCA 

Bivalvia (bivalves) 

Chama pacifica 1996 SF Indo-West Pacific I 

Sphenia coreanica 1968 SF Japan I 

Abra sp. 1996 SF, BW Unknown I 

Venerupis philippinarum 1918 R Japan I 

Lioconcha fastigiata 1985 BW Northwest Pacific I 

Hiatella arctica


15 



CRUSTACEA 

Tanaidacea (tanaids) 

Parapseudes pedispinis 1996 SF, BW Northeast Pacific I 

Apseudes sp. 1996 SF, BW Japan? I 

Leptochelia “dubia” 1932 SF, BW C 

Isopoda (isopods) 

Gnorimosphaeroma rayi 1972 SF, BW Japan I 

Sphaeroma walkeri 1943 SF, BW Indian Ocean I 

Paradella dianae 2002 SF, BW eastern Pacific I 

Pistorius bidens 2002 SF, BW Australia I 

Paracerceis sculpta 1943 SF, BW Northeast Pacific I 

Exosphaeroma sp. 1996 SF, BW Unknown I 

Mesanthura sp. 1996 SF, BW Unknown I 

Limnoria tripunctata 1922 SB Southern Hemisphere? I 

Paralimnoria andrewsi 1922 SB Indo-West Pacific I 

Caecijaera horvathi 1975 SB Southern Hemisphere? I 

Littorophiloscia culebrae 1984 SBA North Atlantic? I 

Halophiloscia couchii 1997 SBA Northeast Atlantic I 

Armadilloniscus ellipticus 1985 SBA Unknown I 

Alloniscus oahuensis 1879 SBA Indo-Pacific I 

Porcellio lamellatus 1973 SBA Northeast Atlantic I 

Buchnerillo sp. 1985 SBA Atlantic Ocean? I 

Olibrinus truncatus 1985 SBA Southern Hemisphere? I 

Ligia exotica 1996 SBA, SF Unknown I 

Cumacea (cumaceans) 

Nannastacus sp. 1996 BW Unknown I 

Scherocumella sp.

16 




INSECTA 

Odonata (damselflies and dragonflies) 

Enallagma civile 1936 R, O Western North America I 

Ischnura ramburii 1973 R, O North/South America I 

Crocothemis servilia 1994 R, O Middle East, Asia, Australia I 

Tramea lacerata 1873 R, O Eastern North America I 

Hemiptera (water bugs) 

Trichocorixa reticulata 1877 R, O North America I 

Mesovelia amoena 1971 R, O North/South America I 

Mesovelia mulsanti 1933 R, O North/South America I 

Micracanthia humilis 1988 R, O North America I 

Hymenoptera (wasps) 

Kleidotoma bryani 1923 R, O Guam, Palmyra I 

Coleoptera (beetles) 

Enochrus sayi 1931 O Western North Atlantic I 

Tropisternus salsamentus 1968 BW California I 

Cercyon fimbriatus 2001 R, O Eastern Pacific I 

Parathroscinus murphyi 1996 BW Southeast Asia I 

Diptera (flies) 

Canaceoides angulatus 1922 R, O Tropical E Pacific I 

Procanace williamsi 1944 R, O Asia (Japan?) I 

Tethina willistoni 1919 R, O North America I 

Atrichopogon jacobsoni 1958 R, O Western Pacific I 

Atrichopogon sp. 1998 R, O Unknown I 

Cricotopus bincinctus 1955 R, O Unknown I 

Goeldichironomus holoprasinus 1969 R, O North/South America I 

Dolichopus exsul 1930 R, O W Indian (Caribbean) I 

Medetera grisescens 1914 R, O Indo-Pacific I 

Syntormon flexible 1917 R, O Indo-West Pacific I 

Psychoda salicornia 1945 R, O California I 

Brachydeutera ibari 1980 R, O Asia I 

Ceropsilopa coquilletti 1946 R, O North America I 

Clasiopella uncinata 1946 R, O Taiwan, Australasia I 

Discocerina mera 1948 R, O Western/South Pacific I 

Donaceus nigronotatus 1958 R, O Asia I 

Ephydra gracilis 1946 R, O Eastern Pacific I 

Ephydra milbrae 1950 R, O Eastern Pacific I 

Hecamede granifera 1923 R, O Western/South Pacific I 

Mosillus tibialis 1944 R, O North America I 

Paratissa pollinosa 1945 R, O Caribbean, S America I 

Placopsidella marquesana 1951 R, O Indo-Pacific I 

Psilopa girschneri 1952 R, O Unknown I 

Scatella stagnalis 1967 R, O Unknown I 

Hostis guamensis 1946 R, O Australia I 

Collembola 

Oudemansia esakii 1939 R, O C 

Dermaptera (earwigs) 

Anisolabis maritima 1953 SBA C


17 



INSECTA 


Thinophilus hardyi 1996 R, O C 

Orthoptera (crickets) 

Thetella tarnis

18 




CHORDATA 

Urochordata: Ascidiacea (sea squirts) 

Didemnum candidum 1930 SF Northeast Atlantic I 

Didemnum perlucidum 1999 SF Unknown I 

Didemnum psammathodes 1998 SF Indo-West Pacific I 

Diplosoma listerianum 1900 SF Unknown I 

Lissoclinum fragile 1962 SF Unknown I 

Ciona intestinalis 1933 SF North Atlantic I 

Ecteinascidia imperfecta 1979 SF Western Pacific I 

Corella minuta


19 



RHODOPHYTA (red algae) 

Acanthophora spicifera 1952 SF, R Indo-Pacific I 

Eucheuma denticulatum 1970 R Philippines I 

Kappaphycus alvarezii 1974 R Philippines I 

Kappaphycus striatum 1970 R Phliippines,Pohnpei I 

Asparagopsis “taxiformis” - Lineage 4 1991 SF? Indo-Pacific I 

Gracilaria salicornia

20 


Table 2. Number of species treated in this monograph and their status 

Taxon 

Introduced 

Cryptogenic 

Establishment 

Unknown 

Released/ 

Escaped 

Not Established 

Failed 

Intercept 

Other* 

Fungi 2 – – – – – – 

Dinoflagellata 1 – – – – – – 

Ciliophora 3 7 – – – – – 

Porifera 9 18 – – – – – 

Hydrozoa 8 27 – – – – 1 

Anthozoa 5 1 1 – 1 – 1 

Scyphozoa 4 – – – – – – 

Cubozoa 1 – – – – – – 

Ctenophora 1 – – – – – – 

Nematoda 1 1 – – – – – 

Rotifera – 5 – – – – – 

Turbellaria 1 1 – – – – – 

Trematoda 1 – – – – – 1 

Monogenea 2 – – – – – – 

Cestoidea 1 – – – – – – 

Hirudinea – – – – – – 1 

Oligochaeta – 7 – – – – – 

Polychaeta 21 21 – – – 1 2 

Gastropoda 11 4 7 2 1 – 3 

Bivalvia 21 – 1 9 – – – 

Copepoda 3 1 – – – – – 

Amphipoda 20 5 – – – – – 

Tanaidacea 2 1 – – – – – 

Isopoda 18 – – – – – 1 

Cumacea 2 – – – – – – 

Mysidacea 1 – – – – – – 

Cirripedia 4 – – – – – – 

Decapoda 10 – – 6 – 3 7 

Stomatopoda 1 – – – – – – 

Insecta 38 4 1 – – – – 

Pycnogonida 12 – – – – – 1 

Phoronida 1 – – – – – – 

Kamptozoa 1 – – – – – – 

Bryozoa 21 – – – – – – 

Echinodermata 1 1 – – – – – 

Ascidiacea 30 1 – – – – – 

Pisces 18 – 3 10 – – 1 

Rhodophyta 9 – – 1 1 – – 

Chlorophyta 1 12 – – – – – 

Phaeophyta 3 – – – – 1 – 

Anthophyta 12 – – 4 – – – 

TOTALS 301 117 13 32 3 5 19 

* Other: Doubtful or erroneous records, native species, natural waifs, uncertain presence in marine 

waters.


21 

Table 3. Vectors transporting nonindigenous marine and estuarine species to the 

Hawaiian Islands 

Abbreviation 

Used in Table 1 

AN 

BW 

O 

R 

SB 

SBA 

SF 

Vector 

Ship anchor and anchor chains 

Ballast water 

Other, including carriage with commercial products, in airplane cabins, resting 

or egg stages with luggage and clothing, etc. 

Releases, intentional or accidental, by private individuals or government agencies, 

and species associated with such releases 

Ship boring (organisms boring into wooden ships, or living in such burrows) 

Solid ballast, including rock and sand ballast 

Ship fouling (organisms living on hulls, in sea chests, or other fouled areas) 

SYSTEMATIC ACCOUNT OF INTRODUCED AND CRYPTOGENIC SPECIES 

VIRUSES, BACTERIA, and FUNGI 

Carr (1996) reviewed the introduced and indigenous viruses, bacteria, and fungi associated with 

introduced penaeid shrimp in mariculture in the Hawaiian Islands. Several of the viruses (but none 

of the bacteria or fungi), including IHHNV (infectious hypodermal and hematopoietic necrosis 

virus), MBV (Penaeus monodon-type baculovirus), and RPS (rhabdovirus of American penaeids) 

were imported to the Islands with live penaeid shrimp. Whether they are established in the wild is 

not known; we thus do not count these viruses in our tabulations of introduced species. 

Zhu et al. (2008) investigated the very diverse microbial consortia associated with the introduced 

sponge Suberites aurantiacus in Pearl Harbor but demured from suggesting which if any of 

the species may not be native. 

FUNGI 

Introduced Marine Fungi 

Species Date Vector Native to 

Cytospora rhizophorae 1902 R Western Atlantic 

Etheirophora blepharospora 1902 R Western Atlantic 

Cytospora rhizophorae Kohlmeyer 

Introduced 

& Kohlmeyer, 1971 

Kohlmeyer (1969, as Cytospora sp.) reported this Western Atlantic marine mangrove fungus from 

Pearl Harbor from Rhizophora mangle, based upon 1968 collections. In describing this as a new 

species, Kohlmeyer & Kohlmeyer (1971) noted further material from Coconut Island, also collected 

in 1968. The Kohlmeyers suggested that this host-specific fungus was introduced in 1902 with 

Rhizophora from Florida. Cytospora rhizophorae has been implicated in widespread dieback and 

mortality of Rhizophora mangle in Puerto Rico (Wier et al., 2006).

22 


Etheirophora blepharospora Kohlmeyer 

Introduced 

& Kohlmeyer, 1965 

= Keissleriella blepharospora 

Kohlmeyer (1969) reported this host-specific fungus from the bark of seedlings and from proproots 

of the mangrove Rhizophora mangle based upon collections made in 1968 on Coconut Island and 

Anahulu River near Hale‘iwa, O‘ahu. Volkmann-Kohlmeyer & Kohlmeyer (1993) further discuss 

the manglicolous fungi of Hawai‘i. 

PROTOCTISTA 

Introduced and Cryptogenic Protozoa 


Introduced Species 

Eufolliculina lignicola 1961 SB Unknown 

Mirofolliculina limnoriae 1961 SB Indo-Pacific 

Cephaloidophora communis 1949 SF Unknown 

Perkinsus marinus 1972 R Western Atlantic 

Cryptogenic Species 

Zoothamnium sp./spp. 1935 SF, BW, R 

Halofolliculina annulata — SF, R 

Parafolliculina violaceae — SF, R 

Metafolliculina nordgardi — SF, R 

Metafolliculina andrewsi — SF, R 

Lagotia viridis — SF, R 

Ascobius simplex — SF, R 

We discuss here examples of introduced and cryptogenic protists. A thorough exploration of the 

Hawaiian marine and estuarine protists will reveal many additional introduced and cryptogenic 

species. We focus here on ciliates, which we treat here in three ecological groups: fouling sessile 

taxa, ectocommensal taxa, and endocommensal taxa. 

The absence of any foraminiferans (“forams”) from this list reflects a lack of biogeographic and 

systematic resolution, rather than the actual absence of nonnative forams in the Hawaiian near-shore 

and harbor environments. This is no more clearly illustrated by the resolution that 32 species of nonnative 

forams have been recognized recently in the eastern Mediterranean fauna (Meriç et al., 2007). 

Forams are often common in fouling communities and on oysters and are easily transported by human 

activities. The lagoonal taxa noted by Resig (1974) from O‘ahu that are not in the Hawaiian fossil 

record would be candidates for further investigation as potential human-mediated invasions, particularly 

species such as Trochammina inflata, noted by Resig as having a “broad latitudinal distribution 

in bays, lagoons, and marshes of both hemispheres,” as well as Haplophragmoides wilberti, H. manilaensis, 

and Lamellodiscorbis aguayoi, three taxa originally described from the Gulf of Mexico and 

Caribbean. 

DINOFLAGELLATA 

Stat & Gates (2008) report the endosymbiotic dinoflagellate Symbiodinium A1 (Symbiodinium 

microadriaticum, Freudenthal sensu stricto) in the native coral Acropora cytherea from the French 

Frigate Shoals in the Northwestern Hawaiian Islands based upon collections made in May 2006. 

They suggested that Symbiodinium “has been vectored into Hawai‘i via Cassiopea sp. from the Red 

Sea and/or Atlantic Caribbean and has spread in its distribution and formed a symbiotic association 

with A. cytherea.“ LaJeunesse et al. (2009) argued that rather than being introduced, the association 

was more likely to be a natural one.


23 

Parsons & Preskitt (2007) report upon a survey of the epiphytic dinoflagellates from coastal 

waters on the island of Hawai‘i. Twenty species were reported for the first time from the Islands, but 

biogeographic relationships of this microflora are insufficiently worked out to determine if any of 

these species have been introduced. 

Class Dinophyceae 

Perkinsidae 

Perkinsus marinus (Mackin, Owen, 

Introduced 

& Collier, 1950) 

= Dermocystidium marinum; = Labyrinthomyxa marina 

Kern et al. (1973) reported this well-known endoparasite, popularly known as “Dermo” from 

Atlantic oysters (Crassostrea virginica) in Pearl Harbor, based upon collections made in July 1972 

during and following a mass oyster mortality (see discussion at C. virginica). Ford & Tripp (1996) 

provide a review of the history and distribution of this lethal, disease-causing organism. It was presumably 

introduced with other imported oysters from the Atlantic coast of North America. We 

assume it is still present in the C. virginica population on O‘ahu. 

CILIOPHORA 

vorticellidae 

Zoothamnium sp. or spp. 

Cryptogenic 

We interpret the record of Edmondson & Ingram (1939: 256) of unidentified “colonial forms of 

Infusoria ... [which] may add somewhat to the friction of [the bryozoan] Bugula, to the stems and 

branches of which they often thickly adhere,” from Kāne‘ohe Bay (1935) likely to be peritrichous 

ciliates in the genus Zoothamnium. Grovhoug & Rastetter (1980: 257) recorded Zoothamnium sp. 

from O‘ahu in fouling. Carr (1996) notes records of Zoothamnium sp. (and Epistylis sp.) as ectocommensals 

of cultured penaeid shrimp. Ship fouling and oyster culture have transported these protists 

around the world. We regard the one or more species of Zoothamnium, as they occur in harbor fouling 

communities in Hawai‘i, as cryptogenic. 

Folliculinidae 

A number of ”cosmopolitan” fouling taxa of these famous “bottle animacules” are recorded from 

Kāne‘ohe Bay [summary in Phillips, 1977; earlier works are those of Andrews (1944) and Matthews 

(1962, 1963, 1964)]. Many of the species with which these folliculinids are associated are recognized 

as introduced fouling organisms. We regard all of the following as cryptogenic in the Hawaiian 

biota, as it seems probable that a number of taxa may have found their way to the Islands on sailing 

vessels, with aquaculture products, and by other means. However, the global biogeography of marine 

folliculinids is poorly known and precludes any certain conclusions at this time: 

Halofolliculina annulata (Andrews, 1944) 

Cryptogenic 

Parafolliculina violaceae (Giard, 1888) 

Cryptogenic 

Metafolliculina nordgardi (Dons, 1924) 

Cryptogenic 

Metafolliculina andrewsi (Giard, 1883) 

Cryptogenic 

Lagotia viridis Wright, 1858 

Cryptogenic 

Ascobius simplex (Dons, 1917) 

Cryptogenic 

Grovhoug& Rastetter (1980) also recorded Parafolliculina violaceae and Metafolliculina andrewsi 

in 1976–1977 from fouling in Pearl Harbor and Kāne‘ohe Bay. Rastetter & Cooke (1979) recorded 

folliculinids from Kāne‘ohe Bay (1977–1978). Bailey-Brock (1989) recorded Halofolliculina annulata, 

Lagotia viridis, and Ascobius simplex on PVC settlement plates on an artificial reef in open 

waters at 20 m, 2.4 km off the southeast coast of O‘ahu.

24 


Eufolliculina lignicola (Fauré-Fremiet, 1936) 

Introduced 

Matthews (1963) found this wood-associated folliculinid in 1961, “abundant in tracheids [of wood] 

opened by burrowing gribbles” [limnoriid isopods], probably on Coconut Island in Kāne‘ohe Bay. It 

likely occurs wherever harbor-dwelling limnoriids are found in Hawai‘i and should be looked for in 

sites such as Pearl Harbor. Fauré-Fremiet (1936) described this species from the Breton coast of 

France, originally in the genus Folliculinopsis. A small species (“fully extended its length does not 

exceed 350 microns”) with a pale blue color, it was associated with “Limnoria lignorum” (at the time 

the general name for most such wood-boring gribbles) and may be a microhabitat specialist. Fauré- 

Fremiet noted that “it builds its rudimentary shell in the pitted tracheids which are cut through by 

the hollowed galleries of these isopods in the pine wood of submerged wrecks” (translated from the 

French). We regard it as introduced based upon its association with introduced limnoriids. While first 

described from Europe, its biogeographic origin requires investigation, as it may of course have 

itself been introduced to the North Atlantic Ocean by shipping from elsewhere. 

Ectocommensal Protists 

Folliculinidae 

Mirofolliculina limnoriae (Giard, 1883) 

Introduced 

Matthews (1963) recorded this species on the pleotelson of the isopod (gribble) Paralimnoria 

andrewsi from samples collected in 1961 probably on Coconut Island in Kāne‘ohe Bay. We regard 

it as introduced with limnoriids. Its known hosts (both Paralimnoria in Hawai‘i and Limnoria elsewhere) 

appear to be rooted in the Indo-Pacific, where we regard this commensal to be native as well. 

Endocommensal Protists 

Cephaloidophoridae 

Cephaloidophora communis Mavrodiadi, 1908 Introduced 

Ball (1950) found this endoparasitic gregarine protozoan in the introduced barnacle Amphibalanus 

eburneus, presumably from O‘ahu, in 1949. It was found only in this barnacle and not in two other 

species of Balanus, nor in 30 species of decapods. Ball (1950) notes that this protozoan was previously 

known from both B. eburneus and other species of Balanus, and that “since B. eburneus has a 

very wide distribution over the world, the presence of C. communis is not surprising.” We regard it 

as introduced with A. eburneus. 

PORIFERA (sponges) 

Introduced and Cryptogenic Porifera 



Suberites aurantiacus 1902 SF Unknown 

Halichondria melanodocia 1960s [


25 

Introduced and Cryptogenic Porifera (continued) 


Callyspongia diffusa 1945 SF 

Chalinid sp. 1997 SF 

Toxiclona sp. 1997 SF 

Biemna sp. 1996 SF 

Clathria procera 1960s [

26 


In the following analysis, we treat as introduced or cryptogenic those taxa that appear to be 

either entirely restricted to, or primarily found in, the float-fouling communities of harbors, suggesting 

that they owe or may owe their presence in the islands to ship-borne introduction. For nomenclature 

and systematics, we follow the World Porifera Database at: http://www.vliz.be/Vmdcdata/ 

porifera/ [accessed December 2007]. 

Class Demospongiae 

Order Hadromerida 

Clionidae 

Cliona sp. or Pione sp. 

Cryptogenic 

= Cliona vastifica of de Laubenfels, 1950 and of Bergquist, 1977 

De Laubenfels (1950), under the name of the North Atlantic species Cliona vastifica Hancock, 1849 

(now Pione vastifica) recorded this boring sponge, based on 1947 collections, from Waialua Bay 

(northwest of Kāne‘ohe Bay) noting it was “common throughout the shallow waters of Hawaii but 

is always inconspicuous. If one breaks up almost any long-submerged calcareous material, such as 

shells or dead coral, one will find this boring sponge. It is very common throughout Kaneohe Bay.” 

De Laubenfels further noted that “It is an abundant species throughout the Old World, but quite 

uncommon (although present) in the New World....... Is vastifica perhaps a recent immigrant to 

North America, carried by barnacles on ship bottoms? Did it reach Hawaii in that same way too?.” 

Bergquist (1977) lists this as the only Cliona in Hawai‘i. Coles et al. (1999a) report it in 1996 collections 

from Pearl Harbor. 

It seems possible that the Clionidae in Hawai‘i may include both indigenous open-coast and 

coral reef taxa, as well as introduced species brought to the Islands by ships (as de Laubenfels suggests) 

and through commercial oyster culture (as discussed for Cliona in general by Carlton, 1979a). 

We treat harbor- and bay-dwelling species as cryptogenic. 

Tethyidae 

Tethya ornata Sarà, Bavestrello & Calcinai, 2000 Cryptogenic 

While we suspect that this species will eventually prove to be an introduction, we conservatively 

treat it here as cryptogenic, pending further knowledge of its distribution and phylogenetic relationships 

with other Tethya species. Sara et al. (2000) have described this sponge as a new species from 

a floating dock in Kāne‘ohe Bay, at Coconut Island, collected in 1985 by Peter Karuso. It remains 

unknown from elsewhere, but we predict it will be found to be an Indo-Pacific species. We consider 

it a probable ship-fouling introduction. It may occur in Pearl Harbor, Ke‘ehi Lagoon (Honolulu 

Harbor), and similar locations as well. 

Suberitidae 

Suberites aurantiacus (Duchassaing & Michelotti, 1864) Introduced 

= Suberites zeteki (de Laubenfels, 1936), described from Florida 

= Terpios zeteki 

This species appears in the Hawaiian literature as either Terpios zeteki or Suberites zeteki. De 

Laubenfels (1950,1951) recorded it from Kāne‘ohe Bay on O‘ahu (1947) and from Ka‘alualu and 

Honaunau, Island of Hawai‘i. De Laubefels (1954a) further recorded it growing in tanks in the 

Honolulu Aquarium, and in 1957 reported it from Waialua Bay (1947) and Ulumoku Pond in West 

Loch, Pearl Harbor (1947) and from boat fouling in Pearl Harbor. DeFelice et al. (2001) note that it 

is in all leeward harbors on O‘ahu and in Kāne‘ohe Bay, and in Nawiliwili Boat Harbor on Kaua‘i, 

nothing that it is “common as fouling in harbors, especially those with some estuarine conditions, 

and in Kāne‘ohe Bay, primarily on floating docks, dock pilings and mangrove roots, also on hulls of 

ships.” Hoover (1998, 2006) illustrates a specimen from Pearl Harbor. Coles et al. (1999b) report it 

from Ke‘ehi Lagoon (1988), Honolulu Harbor, and Ala Wai Harbor. 

De Laubenfels (1950, page 4) appears to refer to this species as one of the two Panamanian 

forms that “conceivably may have been brought to Hawaii on ship bottoms ... . Conversely, they may


27 

have been taken to Panama from Hawaii”; de Laubenfels further states that of these two Panamanian 

species, “at least one of them is abundant on those that are dry-docked at Pearl Harbor.” In his following 

text, the two Panamanian species referred to are Mycale cecilia and Terpios zeteki [Suberites 

aurantiacus]. 

Grovhoug & Rastetter (1980) recorded it from Pearl Harbor and Kāne‘ohe Bay (1976–1977). 

Coles et al. (1999a) report it from Pearl Harbor (1996 collections) and note that it (along with Mycale 

celia) was “so abundant at the power station outfall in East Loch that they form a virtual sponge reef.” 

It was collected, interestingly enough, in 1902 during the Albatross surveys in Pearl Harbor (R.C. 

DeFelice, pers. comm., 1999, based on examination of Smithsonian Institution collections). 

Suberites aurantiacus, first described from the Virgin Islands, occurs in both the Atlantic and 

Pacific. We regard it as introduced to the Hawaiian Islands. Zhu et al. (2008, as S. zeteki) report upon 

the microbial flora of Suberites aurantiacus at the Rainbow Marina, Pearl Harbor. 

Order Halichondrida 

Halichondriidae 

Halichondria (Halichondria) melanodocia de Laubenfels, 1936 Introduced 

Bergquist (1967) recorded this black western Atlantic tropical species on the floating pier at Coconut 

Island, based on material collected in the 1960s. It was previously known from the Caribbean (West 

Indies), Florida, and North Carolina (Hechtel, 1965). Bergquist considered it introduced. Coles et al. 

(1999a) report it from Pearl Harbor based on collections both in 1993 and 1996; it also occurs in Ala 

Wai Harbor (Coles et al., 1999b) and Kāne‘ohe Bay (Coles et al., 2002a). Hechtel (1965) noted its 

habitat in Jamaica as “abundant on pilings and mangrove roots.” We consider float-fouling Hali - 

chondria in Hawai‘i, because of their habitat and their absence in the open marine biota, to be introduced. 

Halichondria coerulea Bergquist, 1967 

Introduced 

Bergquist (1967) described this Halichondria, “a massive, spreading sponge ... found only on fouling 

surfaces” as a new species from the floating dock and stone pier at Coconut Is., Kāne‘ohe Bay 

(1960s collections). It has not been reported elsewhere (M. Kelly-Borges Shanks, pers. comm., 

1996). It was not found in recent surveys of Pearl or Honolulu Harbors (R.C. DeFelice, pers. comm., 

1999). We treat it here as introduced by ship fouling, and thus unknown in its area of its origin. 

Oceanapia sp. or Halichondria sp. 

Cryptogenic 

= Pellina sitiens of de Laubenfels (1957) [now Halichondria sitiens] 

= Pellina eusiphona of Bergquist, 1977 [now Oceanapia eusiphonia] 

De Laubenfels (1957) reported a single colony of a sponge under the name Pellina sitiens (Schmidt, 

1870), now placed in Halichondria, “from a concrete dry dock at Pearl Harbor” (collected in 1948) 

an identification he called “dubious.” Halichondria sitiens is a North Atlantic taxon, and thus a name 

applied with hesitation to Hawaiian material (M. Kelly-Borges Shanks, pers. comm., 1996). Another 

species, Pellina eusiphona Ridley, 1884, now placed in the genus Oceanapia, was described from 

the Indian Ocean and had been recorded by de Laubenfels from the Marshall Islands (de Laubenfels, 

1954b). De Laubenfels (1957) noted that the specimen from Pearl Harbor was intermediate between 

the published descriptions of these two species; Bergquist (1977) placed de Laubenfels’ use of the 

name (but not the actual species) “sitiens” in the synonymy of P. eusiphona. We consider it cryptogenic. 

Topsentia dura (Lindgren, 1897) 

Cryptogenic 

= Halichondria dura 

De Laubenfels (1951) reported “with some hesitation” this Indo-Pacific species in 2 m of water in 

Hilo, based on specimens collected in 1948. R.C. DeFelice (pers. comm., February 1998) reports 

material collected since 1996 in Pearl Harbor and Hilo.

28 


Topsentia sp. 

Cryptogenic 

Coles et al. (1999a) report this species as a new record for Hawai‘i, based on collections in 1996 

from Pearl Harbor. It may be Indo-Pacific in origin (R.C. DeFelice, pers. comm. February 1998). We 

regard it as cryptogenic. 

Order Haplosclerida 

Callyspongiidae 

Callyspongia (Cladochalina) diffusa (Ridley, 1884) Cryptogenic 

This Australasian and Indo-Pacific species was collected in 1947 on the dock of Coconut Island, 

Kāne‘ohe Bay, and also in Waialua Bay (de Laubenfels, 1950) and at Halapē in 1945 and Ka‘alualu 

in 1948 (de Laubenfels, 1951). Hildemann et al. (1980) noted that it “exists in abundance in large, 

ramose colonies” in Kāne‘ohe Bay, where it “thrives in shallow water (0.5 to 5 m) exposed to direct 

sunlight, and exhibits a uniform bright purple pigmentation throughout its soft tissues.” It still occurs 

in Kāne‘ohe Bay (Coles et al., 2002a). It was collected in Pearl Harbor in 1996 [Coles et al., 1999a; 

the record therein of a report from Pearl Harbor in 1947 by de Laubenfels (1957) is in error] and in 

Honolulu Harbor and Kewalo Basin (Coles et al., 1999b) and in Waikīkī (Coles et al., 2002b). 

Hoover (1998, 2006) illustrates a specimen from Coconut Island living in the coral Montipora capitata. 

We consider it cryptogenic. 

Chalinidae 

Haliclona (Soestella) coerulea (Hechtel, 1965) Introduced 

= Sigmadocia coerulea 

Coles et al. (1999a) reports this pale blue-green Caribbean species as a new record for Hawai‘i (and 

thus for the Pacific Ocean), based upon specimens collected in Pearl Harbor in 1996. They note that 

it “is common on artificial substrates of all harbors on the main Hawaiian Islands and Midway 

Atoll.” On O‘ahu it occurs in Kāne‘ohe Bay, Honolulu Harbor, Ke‘ehi Lagoon, Kewalo Basin, and 

Ala Wai Harbor; on Kaua‘i it occurs in Nawiliwili Harbor (DeFelice et al., 2001). DeFelice et al. 

(2001) note that it is unlikely that de Laubenfels or Bergquist would have overlooked it in earlier 

surveys, given its abundance in such localities as the floating docks on Coconut Island. It now occurs 

on pier pilings, floating docks, mangrove roots, and patch reefs and in dredged channels in the 

Hawaiian Islands. We regard it as introduced. When a dry dock left Pearl Harbor in 1999 for Guam, 

it sailed with this sponge as well. Hoover (1998, 2006) illustrates a specimen from Pearl Harbor. 

Hechtel (1965) reported it as “Common on pilings and mangrove roots on the outside of the 

Port Royal mangrove thicket,” and also from a “sandy turtle grass bed,” both in Jamaica. In addition 

to the Caribbean, H. coerulea is also known from the Pacific side of Panama; whether native to the 

Atlantic or Pacific Oceans remains unknown. 

Undescribed chalinid 

Cryptogenic 

Another cryptogenic chalinid sponge occurs in Ke‘ehi Lagoon (1997) and Ala Wai Harbor (1998) 

(Coles et al., 1999b: 117), in Kāne‘ohe Bay (Coles et al., 2002a) and in Waikīkī (Coles et al., 2002b). 

Toxiclona sp. 

Cryptogenic 

An apparently cryptogenic species of Toxiclona occurs in Honolulu Harbor (Coles et al., 1999b; collections 

of 1997), in Kāne‘ohe Bay (Coles et al., 2002a), and at Waikīkī (Coles et al., 2002b). 

Niphatidae 

Gelliodes fibrosa (Wilson, 1925) 

Introduced 

Coles et al. (1999a) report this blue-gray Philippine species from Pearl Harbor (1996 collections), as 

a new state record for Hawai‘i, “abundant on the hull of a floating dry dock [the Machinist] brought 

from the Philippines in 1992.” It now occurs and is established in Pearl Harbor and Honolulu Harbor


29 

on O‘ahu, apparently as a direct result of this introduction from the Philippines, on Maui (Kahului 

Harbor) and on Kaua‘i (Nawiliwili Harbor) (DeFelice et al., 2001), as well as in Kāne‘ohe Bay 

(Coles et al., 2002a) and Waikīkī (Coles et al., 2002b). Coles et al. (1999b) also list it from Ke‘ehi 

Lagoon, Ala Wai Harbor, Barbers Point Harbor, and Kewalo Basin, while Coles et al. (2004) report 

it from Kawaihae and Hilo Harbors, Hawai‘i. 

It is “mainly restricted to shallow-water fouling communities (i.e., pier pilings, floating docks) 

of the major harbors or associated disturbed habitats (i.e., dredged channels and artificial lagoons)... 

(and is) found on patch reefs… typically found encrusting the shaded underside of plate corals” in 

Kāne‘ohe Bay (DeFelice et al., 2001). 

Order Poecilosclerida 

Desmacellidae 

Biemna sp. 

Cryptogenic 

Coles et al. (1997, 1999a) report this species (as B. fistulosa Topent, 1897) as a new record for 

Hawai‘i, based on collections in 1996 from Pearl Harbor. It is also found in Kāne‘ohe Bay (Coles et 

al., 2002a). Hoover (1998, 2006) illustrates a specimen from Magic Island, O‘ahu, at 3 m depth. 

Coles et al. (1999b) report it from Honolulu Harbor, Ke‘ehi Lagoon, and Ala Wai Harbor. 

It appears to be similar to a species known from Zanzibar and the west central Pacific (Coles et 

al., 2002a; M. Kelly-Borges Shanks & R.C. DeFelice, pers. comm.). We regard it as cryptogenic. 

Microcionidae 

Clathria (Thalysias) procera of Bergquist, 1967 Cryptogenic 

Bergquist (1967) recorded Clathria procera (Ridley, 1884), a widespread species occurring from the 

Red Sea to the Indo-Pacific (Hooper, 1996) from Kāne‘ohe Bay, based on material collected in the 

1960s. We treat it as cryptogenic. 

Mycalidae 

Mycale (Carmia) cecilia de Laubenfels, 1936 

Introduced 

= Mycale maunakea de Laubenfels, 1951, synonymy fide Bergquist (1967, as M. manaukea). 

De Laubenfels (1950) noted that this Mycale was “very common in Hawaii, widely scattered in shallow 

water locations. Its greatest abundance, however, seems to be in Honolulu Harbor and in Pearl 

Harbor” (collections of 1947). He further noted that it “... conceivably may have been brought to 

Hawaii on ship bottoms... Conversely, they may have been taken to Panama from Hawaii.” De 

Laubenfels offered reasons to doubt this initial identification, but it is accepted by Bergquist (1967), 

who found it at the floating pier at Coconut Island in Pearl Harbor and who considered it introduced. 

De Laubenfels (1957) noted that Mycale cecilia occurred in Honolulu Harbor in December 1947 “on 

nearly every piling;” he further recorded it from Ulumoku Pond in the West Loch of Pearl Harbor 

(1947), and from barge bottoms in Pearl Harbor. Evans et al. (1972) recorded a Mycale sp. from 

Pearl Harbor (1971). Coles et al. (1999a) report it from Pearl Harbor (1996 collections), and note 

that it (along with Suberites aurantiacus) was “so abundant at the power station outfall in East Loch 

that they form a virtual sponge reef.” 

It also occurs in Kāne‘ohe Bay, Honolulu Harbor, Ke‘ehi Lagoon, Ala Wai Harbor, Barbers 

Point Harbor, Kewalo Basin, Kaua‘i, and Maui (Coles et al., 1999b, 2002a, 2006). A Caribbean 

species (Coles et al., 2002a), we regard it as introduced. Young (1967, as Mycale maunakea) noted 

that the native nudibranch Chromodoris aspersa feeds upon this species in Hawai‘i. 

Mycale (Mycale) grandis Gray, 1867 

Introduced 

= Mycale (Aegogropila) armata Thiel, 1903 

Coles et al. (1999a) report this bright red-orange species (as M. armata) as a new record for Hawai‘i, 

based on collections in 1996 from Pearl Harbor. It is now known from a number of stations around 

O‘ahu (Pearl Harbor, Honolulu Harbor, Ke‘ehi Lagoon, Barbers Point Harbor, and Kāne‘ohe Bay) and 

on Maui at Kahului Harbor (DeFelice et al., 2001). It occurs on pier pilings, floating docks, dredged

30 


channels, and lagoons, and on patch reefs in Kāne‘ohe Bay. Coles et al. (1999b) report it on O‘ahu from 

Honolulu Harbor, Ke‘ehi Lagoon, Ala Wai Harbor, and Barbers Point Harbor. Coles et al. (2006) report 

it from Kaua‘i as well, based upon 2003 specimens. Coles & Bolick (2007) note that M. grandis has 

overgrown reef corals in Kāne‘ohe Bay, where it was found to be smothering Porites compressa and 

Montipora capitata on back reef flats and slopes. Described originally from Ternate, an island in the 

Maluku Islands (Moluccas) in eastern Indonesia, it was previously known from Australia, Torres 

Straits, the Red Sea, and the general Indo-Malaysian region (DeFelice et al., 2001). 

Mycale (Zygomycale) parishii (Bowerbank, 1875) Introduced 

De Laubenfels (1950) recorded material from Kāne‘ohe Bay (1947) and further noted that “It is 

extremely abundant as a growth on vessels that remain for a year or so at harbor on the lee side of 

O‘ahu, as revealed by study of ship bottoms in the dry-docks at Pearl Harbor.” Bergquist (1967) 

recorded it again at the floating pier at Coconut Island and considered it to be introduced, a determination 

with which M. Kelly-Borges Shanks concurs (pers. comm., 1996). McCain (1975) found it 

on fouling panels in Pearl Harbor (1973). Coles et al. (1999a) report it based on collections in 1996 

from Pearl Harbor. Coles et al. (2002b) and DeFelice et al. (2001) record it on O‘ahu from Pearl 

Harbor, Honolulu Harbor, Ke‘ehi Lagoon, Kewalo Basin, Barbers Point, and Kāne‘ohe Bay, and on 

Maui at Kahului Harbor. They note that “this sponge is mainly restricted to shallow-water fouling 

communities (i.e., pier pilings, floating docks) of the major harbors on O‘ahu or associated disturbed 

habitats (i.e., dredged channels and artificial lagoons). In Kāne‘ohe Bay, it is found on patch reefs.” 

Mycale parishii is an Indo-Pacific species, previously known from the Indian Ocean, the East 

Indies (Indonesia), the Philippines, Australia, and the Pacific side of Panama. It has also been recorded 

from the Caribbean (West Indies [Jamaica]) and Brazil (Hechtel, 1965), to where (along with 

Pacific Panama) we regard it as introduced as well via ship fouling. 

Banner & Banner (1966) [repeated by Castro (1971)] reported that in Hawai‘i this sponge hosts 

in its spongocoel the alpheid shrimp Synalpheus streptodactylus Coutière, 1905. They provide no 

details in terms of location or exact collection date but note that it was between 1959 and 1963 

(Banner & Banner, 1966, pp. 145 and 158). This shrimp occurs in other locations in the Pacific and 

Indian Oceans as well but not in association with this sponge. The shrimp is presumably native. 

Crambeidae 

Monanchora sp. 

Introduced 

An encrusting sponge from the fouling community of Honolulu Harbor (1997) and Ke‘ehi Lagoon 

(1997) may be a species of the genus Monanchora [Coles et al., 1999b, identified therein as 

Neofolitispa unguiculata, but probably not that species (R.C. DeFelice, pers. comm., 2001)]. We 

regard it as introduced by ship fouling. 

Tedaniidae 

Tedania (Tedania) reticulata Thiele, 1903 

Cryptogenic 

= Tedania ignis of de Laubenfels, 1950 and Bergquist, 1967 

De Laubenfels (1950) provisionally identified a Tedania, described as “very common throughout the 

shallow waters of Hawaii including Kaneohe Bay” (1947) as the well-known West Indian fire 

sponge T. ignis (Duchassaing & Michelotti, 1864). De Laubenfels (1951) noted material collected in 

1945 at Halape; under the name T. ignis, he further reported this species from shallow subtidal waters 

near Koko Head, near Hanauma Bay (1948) and in dredgings 2–3 km south of Pearl Harbor (1948). 

Bergquist (1967), in studying material from the floating pier at Coconut Island considered it to be 

introduced, and noted that of the species she considered introduced, only this species “is common 

elsewhere in Hawaii.” While de Laubenfels tentatively identified this species as T. ignis, Bergquist 

uses this name without further comment. It appears to be T. reticulata of the Indo-Pacific (M. Kelly- 

Borges Shanks, pers. comm., 1998). We consider it cryptogenic. 

Coles et al. (1999a) report it as T. reticulata from Pearl Harbor (1996 collections), and Coles et 

al. (1999b) report it from Ke‘ehi Lagoon, based on 1997 collections. Hoover (1998, 2006) illustrates 

a specimen from Coconut Island, Kāne‘ohe Bay.


31 

Sims & Irei (1979) report that this “fire sponge” occasionally causes severe human dermatitis 

in Hawai‘i, a phenomenon noted earlier by de Laubenfels in 1950. 

Tedania sp. 

Cryptogenic 

= Tedania macrodactyla of Bergquist, 1967 

Bergquist (1967) recorded a Tedania species [as T. macrodactyla (Lamarck, 1813)] from the floating 

pier and stone pier on Coconut Island, Kāne‘ohe Bay (collections of the 1960s); the taxon T. 

macrodactyla, at the time, was previously otherwise known in the Pacific Ocean from the East 

Indies. 

The applicability of this name to Indonesian or to Hawaiian material is questionable (M. Kelly- 

Borges Shanks, pers. comm., 1996). The identity of this fouling Tedania on O‘ahu thus remains to 

be determined. We regard it as cryptogenic. 

Raspailiidae 

Echinodictyum asperum Ridley & Dendy, 1886 Cryptogenic 

Coles et al. (1999a, 1999b) report this species as a new record for Hawai‘i, based on collections in 

1997 from Pearl Harbor. It is known from the Indo-Pacific (R.C. DeFelice, pers. comm. February 

1998). We regard it as cryptogenic. 

Order Dictyoceratida 

Dysideidae 

Dysidea sp. 

Cryptogenic 

A sponge in the Pearl Harbor fouling community was collected in 1996 by M. Kelly-Borges Shanks 

and R.C. DeFelice that appears to be a new species of Dysidea, although similar to the well-known 

tropical western Atlantic “heavenly sponge” Dysidea etheria de Laubenfels, 1936. It is also found in 

Honolulu Harbor (Coles et al., 1999b) and Kāne‘ohe Bay (Coles et al., 2002a) and in Ke‘ehi Lagoon 

(Coles et al., 2002a). We treat it here as cryptogenic. 

Dysidea sp., cf. D. avara (Schmidt, 1862) 

Cryptogenic 

De Laubenfels (1950) reported a single specimen collected at Coconut Island in Kāne‘ohe Bay in 

1948 “on the east side of the dock, near the concrete bridge which connects the dock with the island,” 

in 2 m of water. It remains in Kāne‘ohe Bay (Coles et al., 2002a). It is also found in Pearl Harbor 

(Coles et al., 1999a; 1996 collections), Honolulu Harbor (Coles et al., 1999b) and in Waīkikī (Coles 

et al., 2002b). Hoover (1998, 2006) illustrated a specimen growing on a mangrove root on Coconut 

Island in one meter. Originally described from the Mediterranean, by the time of de Laubenfels’ 

(1950) report it had been reported from “throughout the Old World, Europe to Australia... even into 

the near Arctic,” which suggests either a species complex or a taxon widely distributed for centuries 

by ships. We regard it as cryptogenic. 

Dysidea sp., cf. D. arenaria Bergquist, 1965 

Cryptogenic 

Coles et al. (1999a) report this sponge, previously known from Palau, as found in Pearl Harbor 

(1996) collections. It is also found in Honolulu Harbor (Coles et al., 1999b), Kāne‘ohe Bay (Coles 

et al., 2002a), and Waikīkī (Coles et al., 2002b). We regard it as cryptogenic. 

Spongiidae 

Hyattella intestinalis (Lamarck, 1814) 

Cryptogenic 

Coles et al. (1999a) report this Indo-Pacific sponge as found in Pearl Harbor (1996) collections; it is 

a new record for Hawai‘i. It was also collected on the hull of the floating dry dock the Machinist 

brought from the Philippines in 1992 (R.C. DeFelice, pers. comm., 2001). Coles et al. (1999b) report 

it from Honolulu Harbor, Ke‘ehi Lagoon, and Barbers Point Harbor. Although said to have been 

from the Mediterranean, the type locality was probably the Indian Ocean (Hooper & Wiedenmayer, 

1994). We regard it as cryptogenic.

32 


Class Calcarea 

Order Leucosolenida 

Heteropiidae 

Heteropia glomerosa (Bowerbank, 1873) 

Cryptogenic 

Coles et al. (1999a) report this cryptogenic species from Pearl Harbor (1996 collections). It also 

occurs in Kāne‘ohe Bay (Coles et al., 2002a). Coles et al. (1999b) report it from Honolulu Harbor, 

Ke‘ehi Lagoon, Ala Wai Harbor, and Kewalo Basin. Material was collected in 1955 on fouling test 

blocks in Honolulu Harbor by C.H. Edmondson (R.C. DeFelice, pers. comm., 1999). It was previously 

known from the Indian Ocean, South Africa, and Australia (M. Kelly-Borges Shanks & R.C. 

DeFelice, pers. comm., 1999). 

CNIDARIA 

Class Hydrozoa (hydroids) 

Introduced and Cryptogenic Hydrozoa 



Cordylophora caspia 1974 SF, BW Ponto-Caspian 

Turritopsis nutricula 1972 SF, BW Atlantic? 

Garveia sp. 1972 SF, BW Unknown 

Bougainvillia muscus 1967 SF, BW Atlantic 

Cladonema radiatum 1972 SF, BW North Atlantic 

Pennaria disticha 1928 SF, BW Unknown 

Obelia dichotoma 1972 SF, BW Unknown 

Obelia bidentata 1946 SF, BW Unknown 


Moerisia horii 1972 SF, BW 

Eudendrium sp. 1972 SF, BW 

Halecium beanii 1943 SF, BW 

Halecium sibogae 2003 SF, BW 

Ventromma halecioides 1999 SF, BW 

Plumularia floridana 1999 SF, BW 

Plumularia strictocarpa 1999 SF, BW 

Plumularia setacea 1972 SF, BW 

Antennella secundaria 1999 SF, BW 

Halopteris diaphana 1970s SF, BW 

Halopteris polymorpha 1999 SF, BW 

Lytocarpia phyteuma 2001 SF, BW 

Anthohebella parasitica 1999 SF, BW 

Clytia latitheca 1999 SF, BW 

Clytia hemisphaerica 1972 SF, BW 

Thyroscyphus fruticosus 1973 SF, BW 

Synthecium megathecum 1972 SF, BW 

Dynamena crisioides 1972 SF, BW 

Dynamena cornicina 1972 SF, BW 

Dynamena quadridentata 2001 SF, BW 

Sertularella areyi 1999 SF, BW 

Sertularella tongensis 1999 SF, BW 

Sertularella diaphana 1972 SF, BW 

Tridentata loculosa 1972 SF, BW 

Tridentata hupferi 1972 SF, BW 

Tridentata turbinata 2001 SF, BW 

Tridentata distans 2001 SF, BW 

Tridentata marginata 2001 SF, BW 

Sertularia subtilis 

Additional Taxon Treated and Its Status 

Status: Record doubtful; see discussion.


33 

As early as 1933, C.H. Edmondson speculated that some species of hydroids may have been transported 

to the islands in ship fouling (Edmondson, 1933). Cooke (1977a) remarked on the presence 

of many cosmopolitan species in the Hawaiian hydroid fauna, stating, “It should be noted ... that 

many of the forms reported were collected from artificial habitats such as floats, pilings, and so forth, 

or from disturbed areas such as Kāne‘ohe Bay. This would also tend to favor the establishment of 

colonizing (in the ecological, not morphological sense) species, which in general are quite widely 

distributed.” Cooke (1975) earlier noted, relative to the Enewetak hydroid biota, that the “many hundreds 

of ships and barges [that] visited Enewetak Atoll in the later part of World War II and during 

the period of atomic bomb testing” provided “a ready method ... for the introduction of so many cosmopolitan 

species.” 

While occasionally recognizing ships as a transport mechanism for harbor-dwelling hydroids, 

the hydrozoan systematic and biogeographic literature, in general, does not attempt to identify or 

suggest which species in subtropical and tropical harbor biotas, or in adjacent habitats in the open 

ocean, may be introduced, presumably in deference to the largely undocumented (and often unlikely) 

possibility of natural dispersal as oceanic drifters. 

We treat several species here as introduced, but most taxa are of necessity cryptogenic. Chu & 

Cutress (1955) reported Syncoryne mirabilis (Agassiz, 1862) (now known as Sarsia tubulosa [(M. 

Sars, 1835); Calder (1988)] as causing dermatitis in swimmers in Hawai‘i. There are no further 

reports of this or any similar hydroid from Hawai‘i, and D. Calder (pers. comm., July 2003) considers 

the report of Chu & Cutress to be a misidentification. We do not further consider it. 

As with a number of other groups, we note that many more species of shallow-water hydroids 

from the islands could be considered cryptogenic. Thus, the species listed here are examples of cryptogens, 

and our treatment is not meant to imply that we consider all other hydroids known from the 

islands to be native. 

Corydendriidae 

Cordylophora caspia (Pallas, 1771) 

Introduced 

= Cordylophora lacustris Allman, 1844 

Cooke (1977a) reported this species in an anchialine (18 to 32 ‰) pond (Hālua Pond) at Cape Kinau, 

Maui, growing on the alga Caulerpa serrulata, collected by Diana Wong and John Maciolek. A 

Ponto-Caspian species native to the Black and Caspian Seas, Cordylophora has been widely distributed 

by ships in the Atlantic and Pacific Oceans. Wong (1975) notes the record; her material was collected 

in the summers of 1974 and 1975. 

Turritopsis nutricula McCrady, 1857 

Introduced 

Cooke (1977a) found Turritopsis growing on the hydroid Halocordyle (= Pennaria) stems, “in harbors 

such as Ala Wai Yacht Harbor” in 1972 (BPBM-D456). Cooke further suggested that “it is likely 

that the medusa figured by Edmondson (1946a); [see Fig. 1b (sic, lapsus for 13b)] represents a 

maturing T. nutricula medusa.” Edmondson’s figure is of a medusa “occasionally taken in the tow 

net on Waikiki reef and in Pearl Harbor, Oahu” (Edmondson,1946a; the same figure and statement 

appears in Edmondson, 1933). However, Edmondson’s medusa is not Turritopsis nutricula (C. Mills, 

pers. comm., June 2003), and it may or may not be a campanulariid, as Edmondson suggested. 

Grovhoug & Rastetter (1980) reported T. nutricula from 1976–1977 collections in Pearl Harbor 

and Kāne‘ohe Bay. It still occurs in Kāne‘ohe Bay (Coles et al., 2002a) and doubtless still occurs in 

Pearl Harbor as well. 

This species, possibly North Atlantic in origin, now also occurs on the Pacific coast of America, 

and in South Africa, Papua New Guinea, and New Zealand (Schuchert, 1996), which we suggest is 

due to ship-borne transport. We thus also consider it introduced to Hawai‘i. 

It is often cited as being described in 1859 (e.g., Calder, 1988), but the date of publication was 

1857 (D. Calder, pers. comm., June 2003).

34 


Moerisiidae 

Moerisia horii (Uchida & Uchida, 1929) 

Cryptogenic 

=Ostroumovia horii 

Cooke (1977a) reported this species, as Ostroumovia horii, otherwise known from Japan, from a 

brackish pond in a lava flow near Honokohau, island of Hawai‘i (collected in 1972; W. Cooke, pers. 

comm., July 1996). He noted that the “recording of this species from both Japan and Hawaii is quite 

interesting, since most species in this family have a severely restricted distribution, unless transported 

by human activity.” We regard it as cryptogenic, pending further taxonomic work. No specimens 

from this collection in the Hawaiians Islands exist in the Bishop Museum collections, but Cooke 

(1977a, fig. 1) provides a figure of Hawaiian material. 

Bougainvilliidae 

Garveia sp. 

Introduced 

We regard an unidentified Garveia found in O‘ahu fouling communities as introduced by ships. 

Cooke (1977a) reported it on the hydroid Pennaria at Ala Wai Yacht Harbor, based upon collections 

in 1972 (W. Cooke, pers. comm., July 1996). Grovhoug & Rastetter (1980) reported it as “Garveia 

humilis (Allman, 1877)” from Pearl Harbor and Kāne‘ohe Bay based on 1976–1977 collections, an 

identification we regard as requiring confirmation. We predict that once identified, it will be found 

to be a species with a disjunct, harbor-based distribution. 

Bougainvillia muscus (Allman, 1863) 

Introduced 

= B. ramosa (van Beneden, 1844), a junior homonym; see Calder, 1988 and Schuchert, 1996). 

Tusov & Davis (1971) recorded a “Bougainvillia sp.” from Kāne‘ohe Bay, O‘ahu in the summer of 

1967. Cooke (1977a) reported B. ramosa from Kāne‘ohe Bay in the form of dense colonies growing 

on the stems of the hydroid Pennaria and noted that it was “more common in temperate waters, but 

known to penetrate subtropically to Brazil and Australia.” Bishop Museum collections contain material 

from Kāne‘ohe Bay collected in October 1972 by T. Gosliner. We regard it as introduced by shipping, 

and, given the history and pattern of records of the species, as native to the Atlantic Ocean, and 

introduced to ports in the Pacific, such as in Australia, Papua New Guinea, and New Zealand 

(Schuchert, 1996). 

Eudendriidae 

Eudendrium sp. 

Cryptogenic 

Cooke (1977a) reported a Eudendrium species (lacking gonophores and thus not resolvable to 

species) from Honolulu Harbor and Kāne‘ohe Bay based upon material collected 1972 (W. Cooke, 

pers. comm., July 1996). We regard it as cryptogenic, as Eudendrium species are also frequently 

found in natural communities worldwide. 

Cladonematidae 

Cladonema radiatum Dujardin, 1843 

Introduced 

This well-known North Atlantic hydroid (Calder, 1988) was first collected in the Pacific Ocean in 

1949, when it was described as a new species from Wellington, New Zealand; it was later reported 

from Japan (Schuchert, 1996). W. Cooke collected it on O‘ahu in June, 1972 (BPBM-D461) who 

noted that “the medusae are often found on the alga Ulva in relatively quiet waters” (Cooke, 1977a). 

It has since been introduced to Puget Sound, Washington (Kozloff, 1996; Wrobel & Mills, 1998). 

Pennariidae 

Pennaria disticha Goldfuss, 1820 

Introduced 

= Halocordyle disticha (following Gibbons & Ryland, 1989) 

= Pennaria tiarella Ayres, 1854 (following Vervoort, 1968) 

= Corydendrium splendidum Boone, 1938 (following Cooke, 1977a) 

Gibbons & Ryland (1989) have argued for the retention of the name Pennaria, with the genus 

Halocordyle as a junior synonym; Schuchert (1996) follows this usage. We regard this common foul-


35 

ing hydroid as introduced with shipping. Paulay et al. (2002) treat this species as cryptogenic in 

Guam, while Cranfield et al. (1998) treat it as introduced to New Zealand, with a European origin. 

It is reported, with various synonyms, from warm-water seas worldwide (Vervoort, 1968; Cooke, 

1975; Cutress, 1977; Gibbons & Ryland, 1989). Cooke (1975) noted that it “is a very cosmopolitan 

species found in temperate and tropical regions of all oceans. It is often found on man-made structures, 

although it is also common on marine angiosperms in temperate waters.” Its origin remains 

unknown. 

The earliest identified Bishop Museum material is from 1929 from Pearl Harbor. Edmondson 

(1933: 23) reported it (as Pennaria tiarella) in Pearl Harbor and Kāne‘ohe Bay, “attached to stones, 

pilings of old wharves, buoys, and other floats.” He noted that it appeared to be identical to the 

Atlantic Ocean P. tiarella McCrady “and may have been transported to Hawaii on the bottoms of 

ships.” 

Boone (1938) redescribed P. disticha as a new species, Corydendrium splendidum, from 

Kāne‘ohe Bay (synonymized by Cooke, 1977a), collected as a single large colony in 2 m (“1 fathom”) 

of water at low tide, in 1928. Edmondson & Ingram (1939) reported it (as Pennaria tiarella) 

in 1935 and subsequent years in fouling in Kāne‘ohe Bay. Hiatt (1954) noted that “there are certain 

areas (Pearl Harbor and Kaneohe Bay) where a species of Pennaria is found abundantly attached to 

almost any solid substrate.” It was further recorded by Evans et al. (1972) from Pearl Harbor (1971, 

as Pennaria sp.), by Long (1974) on offshore and Pearl Harbor panels (1968–1972), and by Brock 

(1995) from Pearl Harbor (1993). Cooke (1977a) reported it from Kāne‘ohe Bay, Ala Wai Yacht 

Harbor, Kewalo Basin, Honolulu Harbor, Ke‘ehi Marina, on pilings, submerged lines, and boat bottoms. 

Cooke noted that it is known to sting when handled. Coles et al. (1999a) record its continued 

presence in Pearl Harbor (1976 collections). DeFelice et al. (2001) note that it now occurs on all the 

main Hawaiian Islands. DeFelice et al. (1998) report it from Midway Atoll, and DeFelice et al. 

(2002) report it from French Frigate Shoals. 

Studies of the feeding biology of Pennaria (as P. tiarella) by Pardy & Lenhoff (1968), and on 

the paths and rates of food distribution by Rees et al. (1970) were based upon specimens from floating 

docks at Coconut Island, Kāne‘ohe Bay. 

Hoover (1998, 2006) illustrates specimens from Lāna‘i Lookout, O‘ahu, from approximately 7 

m depth. He notes that Pennaria disticha is “the most common large hydroid in Hawai‘i” and that it 

grows to its largest size (about 30 cm height) in protected locations such as Kāne‘ohe Bay. 

Haleciidae 

Halecium beanii (Johnston, 1838) 

Cryptogenic 

De Oreo (1946) reported this widespread tropical and temperate hydroid, whose origin is unknown, 

to cause human dermatitis in Hawai‘i, based upon a series of cases beginning in June 1943 in the 

vicinity of Honolulu, with details of further cases in 1945. It is tempting to interpret these first 

reports of Halecium-induced dermatitis as the timing of its introduction during World War II, but 

confirmation of this will await a more detailed study of the chronological appearance of this hydroid 

around the Pacific theater. Cooke (1977a) recorded it from “rubble at a depth of 30 feet in Maunalua 

Bay”, based upon material collected in 1972. He earlier recorded it at Enewetak on pier pilings and 

on rubble from a patch reef in shallow water (Cooke, 1975). The earliest Bishop Museum material 

is from 1945 (Coles et al., 1999b) from Honolulu Harbor, on a wood block at 5 m depth (BPBM 

D260). It was also found in 1998 at Barbers Point Harbor, O‘ahu (Coles et al., 1999b). 

Halecium sibogae Billard, 1929 

Cryptogenic 

Specimens of this Indo-Pacific species were collected in 2003 on the Mahi wreck in O‘ahu in 20 to 

30 m of water by Steve Coles and identified as this species by Dale Calder as a new record for 

Hawai‘i (S.L.Coles, pers. comm., February 2004).

36 


Kirchenpaueriidae 

Ventromma halecioides (Alder, 1859) 

Cryptogenic 

This hydroid was first found in 1999 in Kāne‘ohe Bay (Coles et al., 2002a). It is circumglobal in 

tropical and warm temperate waters (D. Calder, identification and pers. comm. to S.L. Coles, 2001). 

Plumulariidae 

Plumularia floridana Nutting, 1900 

Cryptogenic 

This hydroid was first found in 1999 in Kāne‘ohe Bay and in 2000 in Waikīkī (Coles et al., 2002a, 

2002b). Coles et al. (2006) report it from the island of Hawai‘i, based upon 2003 collections. It is 

reported from all seas (Calder, 1997). 

Plumularia strictocarpa Pictet, 1893 

Cryptogenic 


2002b). Coles et al. (2006) report it from Maui, based upon 2003 collections. It is reported from all 

seas (Calder, 1997). 

Plumularia setacea (Linnaeus, 1758) 

Cryptogenic 

= (?) Plumularia milleri Nutting, 1905, from Hawai‘i (1902) (see Calder, 1991, and discussion below) 

= (?) Plumularia corrugata Nutting, 1900 as reported by Nutting (1905) (see below) 

Cooke (1977a) reported P. setacea from Kāne‘ohe Bay, on a variety of substrates. He suggested that 

the material referred to by Edmondson (1946a, Fig. 13d) is “this species (or one similar).” Bailey- 

Brock (1989) recorded it on PVC settlement plates on an artificial reef in open waters at 20 m, 2.4 

km off the southeast coast of O‘ahu. Plumularia setacea is regarded as a temperate and tropical cosmopolitan 

species (Calder, 1997). Cranfield et al. (1998) concluded that P. setacea was native to 

Europe and introduced to the Pacific Ocean, including New Zealand. We treat it as cryptogenic, 

whose origin remains unclear. 

Calder (1997) concurred with earlier taxonomic conclusions that a hydroid described from 

Hawai‘i, Plumularia milleri, by Nutting (1905) is one of many synonyms of P. setacea, and that 

Plumularia corrugata Nutting, 1900, was also a synonym. However, reminiscent of the situation 

with Sertularella diaphana and S. torreyi, we note that P. milleri was described from 173 m off the 

north coast of Maui, a habitat unusually deep for harbor-dwelling fouling hydroids. Plumularia corrugata, 

as reported by Nutting (1905), was from 223 m off the north coast of Maui, a habitat also 

deep for the genre of shallow-water fouling species considered here. Re-examination of this deeperwater 

material from Hawai‘i would be of value. 

Halopterididae 

Antennella secundaria (Gmelin, 1791) 

Cryptogenic 


2002b); it also occurs in Kanunakakai Harbor, Moloka‘i (Coles et al., 2004). It is circumglobal in 

tropical and warm temperate waters of the Atlantic, Pacific, and Indian Oceans (Calder, 1997). 

Halopteris polymorpha (Billard, 1913) 

Cryptogenic 


2002b); it also occurs in Kahului Harbor, Maui (Coles et al., 2004). It was previously known from 

the Indo-West Pacific and Brazil (D. Calder, identification and pers. comm. to S.L. Coles, 2001). 

Halopteris diaphana (Heller, 1868) 

Cryptogenic 

Cooke (1977a) reported this hydroid, cosmopolitan in tropical seas, from a variety of substrates in 

Kāne‘ohe Bay. In more recent years it has been found in Kahului Harbor, Maui (Coles et al., 2004). 

Aglaopheniidae 

Lytocarpia phyteuma (Kirchenpauer, 1876) 

Cryptogenic 

This species was first found in 2001 in Waikīkī (Coles, 2002b).


37 

Lafoeidae 

Anthohebella parasitica (Ciamician, 1880) 

Cryptogenic 


2002b). It was previously known from the eastern Atlantic Ocean and the Indo-West Pacific (D. 

Calder, identification and pers. comm. to S.L. Coles, 2001). 

Campanulariidae 

We regard several Hawaiian campanulariids found largely in fouling communities as introduced by 

ships. Edmondson & Ingram (1939) reported unidentified campanulariids in Kāne‘ohe Bay in 1935 

and later years, doubtless representing one or more of the following species. Long (1974) reported 

an Obelia sp. found on fouling panels outside of Pearl Harbor. 

Obelia dichotoma (Linnaeus, 1758) 

Introduced 

This fouling hydroid, possibly of Atlantic origin, has been reported from Pearl Harbor and Kāne‘ohe 

Bay by several workers since the 1970s (Cooke, 1977a; Grovhoug, 1979; Grovhoug & Rastetter, 

1980; Coles et al., 2002a). Bishop Museum collections contain material (BPBM-D458) collected in 

October, 1972, in Kāne‘ohe Bay by T. Gosliner. It has likely been in the Islands for a great many 

years. Cornelius (1975) did not cite Hawaiian records. 

Obelia bidentata Clark, 1875 

Introduced 

= Laomedea bicuspidata 

This fouling hydroid, also possibly of Atlantic origin, was listed (as Laomedea bicuspidata) by 

Vervoort (1946) from Hawai‘i, with no further details [Cornelius (1975) repeated this citation]. 

Grovhoug (1979) recorded it from Pearl Harbor (1978 collections) as Obelia bidentata (?). It is not 

listed by Cooke (1977a). Coles et al. (2002a) list it for Kāne‘ohe Bay. 

Clytia latitheca Millard & Bouillon, 1973 

Cryptogenic 

This hydroid was first found in 1999 in Kāne‘ohe Bay (Coles et al., 2002a). It was previously known 

from the Indian Ocean and Bermuda (D. Calder, identification and pers. comm. to S.L. Coles, 2001). 

Clytia hemisphaerica (Linnaeus, 1767) 

Cryptogenic 

Cooke (1977a) recorded this fouling species from many different sites in Kāne‘ohe Bay, based upon 

collections made in 1972 (W. Cooke, pers. comm., July 1996). Grovhoug (1979) reported it from 

Pearl Harbor (1978) and Grovhoug & Rastetter (1980) recorded it from both Pearl Harbor and 

Kāne‘ohe Bay (1976–1977). It is to be expected in nonfouling facies in Hawai‘i, considering reports 

of it on sea grasses on intertidal sandflats and other habitats in Fiji (Gibbons & Ryland, 1989), if correctly 

identified there. Calder (1991) has provided a long list of synonyms and taxonomic discussion. 

Thyroscyphidae 

Thyroscyphus fruticosus (Esper, 1793) 

Cryptogenic 

This Indo-Pacific species was first collected in 1973 by W. Cooke in Honolulu Harbor, at 0.3 m “on 

rubble”, where it still occurs (Coles et al., 1999b, as T. fructicosus). It also occurs in Nawiliwili 

Harbor, Kaua‘i, and is very abundant in Hilo Harbor (Coles et al., 2004). 

Syntheciidae 

Synthecium megathecum Billard, 1924 

Cryptogenic 

= Synthecium tubithecum of authors, not Allman, 1877 

This cryptogenic species was reported by Cooke (1977a, as Synthecium tubitheca) from Kāne‘ohe 

Bay in 1–2 m of water as moderately common on coral rubble (based on collections made in 1972; 

W. Cooke, pers. comm., July 1996); it still occurs there (Coles et al., 2002a). It also occurs at Waikīkī 

(Coles et al., 2002b). Nutting (1905, as Synthecium tubithecum) much earlier reported this species

38 


from Hawai‘i based upon collections made in 1902 by the R/V Albatross. These collections were off 

the south coast of Moloka‘i (in 128 m), and of the northeast coast of Hawai‘i (in 53 m). As with 

Sertularella tongensis and Plumularia setacea, the specimens upon which these deeper water 

records of S. tubithecum are based bear reexamination, and we use 1972, rather than 1902, as the 

first date of record. 

Relative to the specific name, D. Calder (pers. comm., June 2003) advises using S. megathecum 

for Pacific populations, and reserving at this time the name S. tubithecum for Atlantic populations. 

Sertulariidae 

Dynamena crisioides Lamouroux, 1824 

Cryptogenic 

This cryptogenic species was reported by Cooke (1977a) from Kewalo Basin and Honolulu Harbor, 

O‘ahu, “where [it is] common on substrates of rock and rusting iron.” Bishop Museum collections 

contain material collected in October 1972 by W. Cooke in Honolulu Harbor. It also occurs in Ke‘ehi 

Lagoon and Barbers Point Harbor (Coles et al., 1999b) and in Kāne‘ohe Bay (Coles et al., 2002a). 

Coles et al. (2006) report it from Kaua‘i, based upon 2003 collections. 

As with the previous species, it is widely recorded from the tropical Atlantic, Pacific, and Indian 

Oceans (Cooke, 1977a; Calder, 1991) and may also represent one or more species. This is primarily 

an intertidal species, apparently (Calder, 1991), and Gibbons & Ryland (1989) note that in Fiji it 

is “the only hydroid to occur on coral boulders that endure long emersion.” Cooke (1975) made similar 

remarks about this hydroid’s intertidal adaptations. 

Dynamena cornicina McCrady, 1859 

Cryptogenic 

This cryptogenic species was reported by Cooke (1977a) from Coconut Island, Kāne‘ohe Bay, based 

upon material collected in 1972 (W. Cooke, pers. comm., July 1996). It is also recorded from the 

tropical Atlantic, Pacific, and Indian Oceans (Cooke, 1977a) and may represent one or more species. 

Gibbons & Ryland (1989) report this species as “widely distributed on sublittoral rock” in Fiji up to 

about the low water mark. However, Calder (1991) notes that this species may be synonymous with 

Dynamena disticha (Bosc, 1802). It is also reported from Nawiliwili and Port Allen Harbors, Kaua‘i, 

Kaunakakai Harbor, Moloka‘i, and Kahului Harbor, Maui (Coles et al., 2004). 

Dynamena quadridentata (Ellis & Solander, 1786) Cryptogenic 

This hydroid was first found in 2001 in Waikīkī (Coles et al., 2002b). It occurs widely in the Atlantic, 

Indian, and Pacific Oceans (Calder, 1991). 

Sertularella areyi Nutting, 1900 

Cryptogenic 

This hydroid was first found in 1999 in Kāne‘ohe Bay and in 2000 in Waikīkī (Coles et al., 2002b). 

It was previously known from the western Atlantic Ocean, Japan, and Korea (D. Calder, identification 

and pers. comm. to S. Coles, 2001). 

Sertularella tongensis Stechow, 1919 

Cryptogenic 

This hydroid was first found in 1999 in Kāne‘ohe Bay (Coles et al., 2002a, 2002b). It was previously 

known from the western Pacific Ocean (D. Calder, identification and pers. comm. to S.L. Coles, 

2001). 

Sertularella diaphana (Allman, 1885) 

Cryptogenic 

= Sertularella speciosa Congdon, 1907 

= (?) Sertularella torreyi Nutting, 1905, from Hawai‘i (1902) (see Calder, 1991, and discussion below) 

Cooke [1977a, as S. speciosa (type locality, Bermuda) but noting the probable synonymy with S.


39 

diaphana (type locality, Australia)] extended the range of this species to the Hawaiian Islands by 

reporting it from Kāne‘ohe Bay “from shallow water”, based upon material collected in 1972 (W. 

Cooke, pers. comm., July 1996). Hoover (1998, 2006) illustrates specimens from Palea Point, O‘ahu, 

in 7 m of water. 

Based upon many synonyms, this is a cosmopolitan hydroid occurring in the Atlantic, Pacific, 

and Indian Oceans (Calder, 1991), or it is perhaps a species complex. Calder (1991) concurred with 

earlier taxonomic conclusions that a hydroid described from Hawai‘i, Sertularella torreyi, by 

Nutting (1905) is another synonym of S. diaphana. We note, however, that Nutting’s material was 

from deep water (128 m) off the south coast of Moloka‘i, as well as from between 742 and 1372 m 

off Kaua‘i, suggesting a re-evaluation of shallow-water and deep-water taxa may be of value. 

Sertularia subtilis Fraser, 1937 

[See Discussion] 

Cooke (1977a) extended the range of this western tropical Atlantic Ocean (previously known from 

Puerto Rico) species into the Pacific Ocean by reporting it from Kahe Point, O‘ahu. However, S. subtilis 

is a poorly known species, and Cooke’s specimens are regarded as a misidentification of 

Tridentata hupferi (D. Calder, pers. comm., June 2003). 

Tridentata loculosa Busk, 1852 

Cryptogenic 

= Sertularia ligulata Thornely, 1904 

This widespread tropical Atlantic, Pacific, and Indian Ocean hydroid was reported by Cooke [as 

Sertularia ligulata (Lamouroux, 1816)], from the surface of Porites lobata in 2 m of water in 

Kāne‘ohe Bay (material collected in 1972; W. Cooke, pers. comm., July 1996). It was found again 

in 1999 in Kāne‘ohe Bay and in 2000 in Waikīkī (Coles et al., 2002a, 2002b, the latter as Tridentata 

ligulata). 

Tridentata hupferi (Broch, 1914) 

Cryptogenic 

= Sertularia subtilis of Cooke (1977a), not Fraser, 1937 

Cooke (1977a) reported this species (as Sertularia subtilis) from 2 m of water in coral rubble at Kahe 

Point reef, based upon material collected in 1972 (W. Cooke, pers. comm., July 1996). Cooke’s identification 

of S. subtilis is regarded as a misidentification of Tridentata hupferi by D. Calder (pers. 

comm., June 2003). It still occurs in Kāne‘ohe Bay, based on 1999 collections identified by D. 

Calder (Coles et al., 2002a, as T. humpferi). It was previously known from West Africa, the 

Seychelles, and Fiji (D. Calder, identification and pers. comm. to S.L. Coles, 2001). 

Tridentata turbinata (Lamouroux, 1816) 

Cryptogenic 

This hydroid was first found in 2001 in Waikīkī (Coles et al., 2002b). It is widely distributed in the 

Indian, Pacific, and Atlantic Oceans (Calder, 1991). 

Tridentata distans (Lamouroux, 1816) 

Cryptogenic 

This species was reported by Cooke (1977a, as Sertularia distans gracilis) from coral rubble at 2 m 

at Kahe Point Reef, O‘ahu (material collected in 1972; W. Cooke, pers. comm., July 1996). It was 

reported again in 2001 in Waikīkī (Coles et al., 2002b). It is widely distributed in the Indian, Pacific, 

and Atlantic Oceans (Calder, 1991). 

Tridentata marginata (Kirchenpauer, 1864) 

Cryptogenic 

This hydroid was first found in 2001 in Waikīkī (D. Calder pers. comm. June 2002 to S.L. Coles). It 

is widely distributed in the Indian, Pacific, and Atlantic Oceans (Calder, 1991).

40 


Class Anthozoa (corallimorpharians, corals and anemones) 

Introduced and Cryptogenic Anthozoa 



Culicia rachelfitzhardingeae 1983 SF, BW Indo-Pacific 

Carijoa riisei 1972 SF, BW Indo-Pacific 

Diadumene leucolena 1950s SF, BW, R Northwest Atlantic 

Diadumene lineata 1999 SF, BW, R Japan 

Diadumene franciscana 1998 SF, BW Unknown 


Sarcothelia sp. 2000 SF, BW 

Additional Taxa Treated and Their Status 

Discosoma nummiforme 1997 R Indo-Pacific 

Status: Released not established 

Dendronephthya sp. 1980s BW? Indo-Pacific 

Status: Failed 

Montipora turgescens 

Status: Native 

Order Corallimorpharia (corallimorpharians) 

Stichodactylinae 

Discosoma nummiforme Leuckart in Rüppell, 1828 Released; Not Established 

= Actinodoscus nummiformis 

Coles et al. (1999b, p. 47) reported that this anthozoan was first found in the Ala Wai Yacht Harbor 

in December 1997 by Waikīkī Aquarium staff. The specimens “were artificially attached to a piece 

of coral rubble, indicating that the organisms had been discarded from a salt-water aquarium. Three 

small clumps were still growing at the site in April 1999, but the organism has not been observed 

elsewhere in or outside of the harbor.” It is considered a private release (D. Gulko, pers. comm., 

2003). 

The colonies expanded despite a few attempts to remove them between 1999 and 2004, at 

which time a formal eradication was launched; no further colonies or individual polyps were detected 

as of fall 2006 (A. Montgomery & S. Pelleteri, pers. comm., February 2007). 

Order Scleractinia (stony corals) 

A thorough study of the pre- and post-World War II coral diversity in the Islands, and a comparison 

with the islands fossil record, remains to be undertaken. Grigg (1981) noted a hypothesis that 

Acropora species “may have been introduced by man via vessel traffic between Hawaii and other 

islands where Acropora exists” but concluded that “In light of the geological record, it is unlikely 

that Acropora was introduced to Hawaii by man. We know from the reef core at French Frigate 

Shoals that Acropora was in the Hawaiian Islands at least 100 years before Western contact. There 

is a remote possibility that Acropora could have been introduced by the early Polynesians on the bottoms 

of their canoes. However, if this were true, Acropora should be present in the high islands 

where the majority of the Hawaiians settled.” Maragos et al. (2004) noted seven species of Acropora 

were present in the Northwestern Hawaiian Islands and were nearly absent from the main islands. 

Specimens of living and dead corals from the Indo-Pacific and not otherwise known from 

Hawai‘i occasionally are encountered in shallow waters of O‘ahu and other islands. These are usually 

discarded specimens from aquaria, or may on occasion represent actual attempts to establish 

new colonies of “exotic” (in the aquarium trade sense) species.


41 

Rhizangiidae 

Culicia rachelfitzhardingeae Cairns, 2006 

Introduced 

= Culicia sp., cf. C. tenella of Fitzhardinge (1985, 1993) not of Dana, 1846 

This small (4 mm in diameter and 1.5 mm in height) ahermatypic coral was first recorded in 1983 

by Fitzhardinge (1985, 1993) from Kāne‘ohe Bay, where it exists both as solitary polyps and as 

“pseudocolonies” of asexually budded clones. Fitzhardinge recorded it on the underside of coral rubble 

and beneath coral colonies; Culicia also settled on experimental concrete blocks. Fitzhardinge & 

Bailey-Brock (1989) reported it settling on artificial reef materials in Kāne‘ohe Bay and noted that 

this coral recruited in all seasons. 

In 2002–2003 Pakki Reath collected additional specimens in fouling communities on Kaua‘i 

(the main pier at Nawiliwili Harbor, 2002, and at the main docks at Port Allen, 2002) and on 

Moloka‘i (Kaunakakai Dock, 2003); it was also collected at Maui and Hawai‘i in 2002–2003 (Coles 

et al., 2004; Cairns, 2006). It may have been overlooked in Pearl Harbor fouling surveys in 1996 

because of its small size and cryptic nature (S.L. Coles, pers. comm., January 2004). 

This fouling coral was described as a new species from Hawai‘i in 2006; it is known not from 

elsewhere. The last shallow-water endemic Hawaiian coral was described in 1907 (S. Cairns, pers. 

comm., November 2005). Although this is a small species, it is not likely to have been overlooked 

prior to the 1980s, as many of the same habitats and locations were well-collected and well-explored 

prior to this time. It is restricted to docks and harbors or to very shallow water in locations known 

to be highly invaded, such as Kāne‘ohe Bay. 

Species of Culicia are widespread through the Indo-Pacific (Maragos, 1974; Maragos & Jokiel, 

1978; Veron, 1986; Cairns et al., 1999; Randall, 2003). We believe this coral was introduced to the 

Hawaiian Islands in ship hull fouling. 

Acroporidae 

Montipora turgescens Bernard, 1897 

Native 

Coles (1998) discovered this widespread Indo-Pacific coral in the Hawaiian archipelago in 1997 at 

the “Reef Hotel” in eastern Midway Atoll lagoon. Originally described from the Great Barrier Reef, 

it occurs from the Red Sea to the southern islands of Japan, and throughout the Indo-Pacific to the 

islands of French Polynesia and the Caroline and Phoenix Islands. Nonetheless, the closest populations 

to Midway are over 3500 km to the west and southwest in southern Japan (Coles, 1998). Coles 

thus noted that the “most likely origin by natural colonization is Japan, with planulae having been 

transported eastward on the North Pacific current”, but further commented that “as an alternative to 

natural transport by currents, M. turgescens may have been transported to Midway in the last century 

as hull fouling or in ballast water of ships utilizing the Midway harbor.” Maragos et al. (2004) 

found this coral more widely distributed in the Northwestern Hawaiian Islands, increasing in occurrence 

along the northwest half of the chain, and noted that the Hawaiian form of this species has not 

been observed outside Hawai‘i and does not conform to published photographs, suggesting that a 

cryptic endemic species may be involved. 

Subclass Octocorallia 

Order Alcyonacea (soft corals) 

Nephtheidae 

Dendronephthya sp. 

Failed 

Gulko (1998) reproduced a photograph (derived from a video tape) of a colony of “Dendronephthya 

sp.”, a genus of soft coral previously unknown in the Hawaiian Islands, that was taken “a couple of 

years ago” in approximately 70 m of water [“a couple hundred feet’], “growing on the side of a 

sewage outfall off of O‘ahu.” Gulko further remarked that, “This probably represents an accidental 

introduction.” This appears to be the same colony tracked by Richard Brock “over an approximately 

7 year period” (while he watched it grow in size) in the 1980s, but searches in the 1990s failed to 

find it, perhaps because of the impacts to the area where it occurred by Hurricane Iniki in September 

1992 (R. Brock, pers. comm. March 2005 to S. Kahng). The colonies were located near Honolulu 

Harbor and may have been a ship-mediated introduction.

42 


Xeniidae 

Sarcothelia sp. 

Cryptogenic 

Coles et al. (2002a) report this xeniid soft coral from Kāne‘ohe Bay (2000) as a first record from 

Hawai‘i and as a cryptogenic species. It is not the same as Sarcothelia edmondsoni (Verrill, 1928), 

a species known only from the Hawaiian Islands. 

Clavulariidae 

Carijoa riisei (Duchassaing & Michelotti, 1860) Introduced 

= Telesto riisei 

This “orange soft coral” (Henderson, 1990) or “snowflake coral” (Gulko, 1998; Hoover, 1998, 2006) 

was long thought to be native to the western Atlantic Ocean, from Florida to Brazil, but has now been 

shown to be native to the Indo-West Pacific (Kahng, 2005; S. Kahng, pers. comm., November 2005). 

It was first described from the Caribbean, presumably introduced there in the 19th century by shipping 

from the Pacific Ocean. 

Carijoa riisei was first found in the Hawaiian Islands in 1972 by William J. Thomas in the fouling 

community in Pearl Harbor (Evans et al., 1974; Thomas, 1978, 1979). Devaney (1977) noted that 

it was also in Honolulu Harbor and “several other areas along the leeward Oahu coast.” Henderson 

(1990) reported it on the USS Arizona in Pearl Harbor (1986 collections); Brock (1995) again reported 

it from Pearl Harbor (1993); Coles et al. (1999a) record its continued presence there (1996 collections). 

It also occurs in Kāne‘ohe Bay (Coles et al., 2002a). Coles & Eldredge (2002) review its 

apparent spread from harbors to outer coast areas. 

Additional records include Honolulu Harbor, Hawai‘i Kai Marina, Port Allen Harbor, Kaua‘i, 

Kaunakakai Harbor, Moloka‘i, Kahului and Mā‘alaea Harbors and Mala Wharf, Maui, and Hilo 

Harbor (Coles et al., 1999b, 2002b, 2004, and 2006). 

Bishop Museum collections include material collected on July 27, 1972 by Mike Lee, in about 

7 m of water (“20 feet”) in Pearl Harbor. Colonies can also be found in high energy subtidal sites 

around O‘ahu (C. Zabin, pers. comm., February 2000). Russo (1994) stated that it occurs to 50 m in 

the Hawaiian Islands; Grigg (2003) recorded it to 120 m in Hawai‘i. DeFelice et al. (2001) note that 

it is now throughout the main Hawaiian Islands. Eldredge & Englund (2001) list Kaua‘i, O‘ahu, 

Moloka‘i, Maui, and Hawai‘i. Kahng & Grigg (2005) note that it “is commonly observed on hard 

substrata in shaded habitat with moderate current flow. In the clear oceanic water surrounding 

Hawai‘i, C. riisei is restricted to shaded habitat at depths shallower than 40 m”, and was “not 

observed ... deeper than 115m.” 

Hoover (1998, 2006) illustrates specimens from Mākua, O‘ahu, in 6 m of water, noting that it 

“forms dense colonies in cavities along vertical walls or on the ceilings of caves and overhangs 

where current is strong, or under docks in harbors were plankton is plentiful. It is abundant on 

wrecks. It now occurs in all the main Hawaiian Islands and at Midway.” Color photographs of this 

species are also in Gulko (1998) and Fenner (2005). 

Grigg (2003) reported that in 2001 C. riisei was very abundant at depths of 75 to 100 m on the 

“Maui Black Coral Bed”, where up to 90% of the black coral colonies of Antipathes dichotoma and 

A. grandis were dead, having been overgrown by C. riisei. Grigg (2004) further discussed the impact 

of Carijoa on the harvest of the black coral Antipathes. Kahng & Grigg (2005) noted the extensive 

smothering of black coral at depths between 80 and 105 m, and found more than 60% of all black 

coral colonies “were at least partially overgrown.” 

Concepcion et al. (2008) report a possible second species of Carijoa in the Hawaiian Islands. 

The taxon in question is a transparent species, and a combination of ecological, morphological, and 

genetic data suggest that it may be a distinct species: these include the absence of green flourescent 

pigments, always present in C. riisei, and the presence of a “unique concatenated mtDNA haploytpe 

and a unique nDNA SRP54 allele.”


43 

Order Actiniaria (sea anemones) 

As with many of the Hawaiian corals, several widespread Indo-Pacific sea anemones occur in the 

Hawaiian Islands. While some of these are potential candidates as introduced species, genetic work 

on possible source regions, an understanding of their potential (or lack thereof) for long-distance 

planktonic or drifting dispersal, as well as verification that some of these taxa could interface with 

human-mediated dispersal vectors (such as fouling or ballast) would be necessary before assigning 

taxa to a cryptogenic category. An example is the phymanthid anemone Heteranthus verruculatus 

Klunzinger, 1877, known in Kāne‘ohe Bay “on sills of concrete spillways between fish ponds” 

(Cutress, 1977), and otherwise known from the Red Sea and Eastern Australia. It was not recorded 

by Verrill (1928) but nor were other apparently native shallow-water actinians reported by Cutress 

(1977). 

Another example is Gyractis excavata Bovieri, 1893 [=Actinogeton sesere (Haddon & 

Shackleton, 1893) fide England, 1987]. Dunn (1974), upon newly finding this anemone on O‘ahu (in 

1973, fide England, 1987, p.255) asked “whether this actinian has arrived in the islands only recently, 

or whether it was overlooked or not recorded” in previous studies. It occurs in Kāne‘ohe Bay on 

“firm substrates, usually on pieces of dead coral, intertidally and in shallow water on sandy shores 

and reef flats” (Dunn, 1974) and also on “exposed coasts” at Manana Island, O‘ahu (Cutress, 1977). 

It is possible to visually lump this anemone in the field with the zoanthiniarian Palythoa, and this 

potential confusion may account for the lack of earlier records (D. Fautin, pers. comm., 18 February 

1998). It is known from the Torres Strait (the type locality of A. sesere), and by synonymy with other 

species and by additional collections from Sri Lanka, Zanzibar, South Africa, Western Australia, 

Aden, Goa, Singapore, and Fiji ( England, 1987). Fautin (pers. comm.,1998) notes that she has further 

collected it in Madang, Papua New Guinea. If the species England identified as synonyms do in 

fact encompass one biological species, then this is a widespread anemone of the South Atlantic and 

Indian Oceans. 

If taxa such as Heteranthus and Gyractis are associated with ship-bottom fouling communities, 

or could be dispersed as planulae or as newly settled stages on floating debris in ballast water, then 

such distributions could have been achieved long ago, and they should be regarded as cryptogenic. 

Diadumenidae 

Diadumene leucolena (Verrill, 1866) 

Introduced 

This small translucent anemone was probably collected as early as the 1950s in O‘ahu. Hiatt (1954) 

listed “Diadumene (leucolena?)” in his key to Hawaiian invertebrates. Hiatt would have been familiar 

with D. leucolena in his graduate student years at the University of California at Berkeley, as this 

anemone was used in zoology classes at the time and is common on the eastern shore of San 

Francisco Bay. He described his D. leucolena (1 cm x 4 cm) as having white catch tentacles, with a 

green-black column distinctly divided into regions and with green or white tentacles. Diadumene 

leucolena, as described by Cutress (1977) has the column distinctly divided into two regions, with a 

color varying from a dark, dirty green to greenish brown to dirty white. Hiatt’s tentacle description 

matches that of Hand (1956). Cutress (1977), definitely identifying this anemone, recorded it from 

Pearl Harbor, Honolulu Harbor, and the Ala Wai Canal. It also occurs in Honolulu Harbor (1997, 

Coles et al., 1999b) and in Waikīkī (2000, Coles et al., 2002b). 

Diadumene leucolena is native to the Northwest Atlantic Ocean as far south as the West Indies 

and Caribbean (Cutress, 1977) and has also been introduced to the Pacific coast of America (Carlton, 

1975, 1979b). Cutress suggested it could have been introduced with commercial oyster culture, 

although introduction by ships is possible as well. As warmer-adapted genotypes would fair better 

in Hawai‘i, it may be that this is another Caribbean element in the Hawaiian biota. 

Godwin (2003b) notes that D. leucolena was present on a barge towed from San Diego to O‘ahu 

in 1999; D. leucolena is a nonnative species in California as well.

44 


Diadumene lineata (Verrill, 1869) 

Introduced 

= Haliplanella luciae (Verrill, 1898) 

Chela Zabin discovered a population of this well-known orange-striped sea anemone on and among 

Pacific oysters (Crassostrea gigas) on intertidal pilings on the south shore of Kāne‘ohe Bay on 15 

February 1999 (Zabin et al., 2004). In 2002 and 2003 additional specimens were found at Coconut 

Island in Kāne‘ohe Bay in and around empty tests of the barnacle Amphibalanus reticulatus on intertidal 

fouling panels and under a pier on a piece of intertidal coral rock (Zabin et al., 2004). 

In November 2000, about 100 individuals of D. lineata were found by S. Godwin on a drift net 

in the lagoon at Pearl and Hermes Reef, Northwestern Hawaiian Islands (Zabin et al., 2004). 

Whether this net drifted in from other Hawaiian Islands, or from elsewhere in the Pacific, is 

unknown, although if the latter, it would constitute the first evidence of transoceanic dispersal of a 

neritic sea anemone. 

Native to Japan, it was introduced to the North Atlantic Ocean in the 1890s and to the Pacific coast 

of North America in the early 1900s (Carlton, 1979a). It has also been introduced to Indonesia, New 

Zealand, and Brazil (Zabin et al., 2004). This anemone may have arrived in the Hawaiian Islands 

between the 1970s and late 1990s, as it would not likely have been overlooked by C.E. Cutress 

(Cutress, 1977), an experienced sea anemone field naturalist and systematist. Possible transport vectors 

include fouling on commercial or recreational vessels, ballast water, or with imported oysters. 

Diadumene franciscana Hand, 1956 

Introduced 

Coles et al. (1999b) reported this anemone from Ala Wai Yacht Harbor in Honolulu based upon specimens 

collected in 1998. It was previously known from bays along the central California coast, to 

where it is also considered introduced (Carlton, 1979a). Specimens in Ala Wai were found near the 

Waikīkī Yacht Club and near the outflow into the harbor from Hilton Lagoon. A specimen of what 

was likely D. franciscana was found on the south shore of Kāne‘ohe Bay in May 2001 by Chela 

Zabin and J.T. Carlton (Zabin et al., 2004). 

The origin of this anemone is not known, but it may be from the western or southwestern 

Pacific Ocean, Australasia, or the Indian Ocean (Carlton, 1979b). Shipping has likely led to its presence 

in the Hawaiian Islands, where it is doubtless more widespread than indicated here. 

Classes Scyphozoa (jellyfish) and Cubozoa (cubomedusae) 

Introduced Scyphozoa and Cubozoa 


Scyphozoa 

Aurelia sp. 1953 SF, BW Indo-Pacific 

Cassiopea andromeda 1950 SF, BW Indo-Pacific 

Cassiopea sp. 2000 SF, BW Indo-Pacific 

Phyllorhiza punctata 1933 SF, BW Indo-Pacific 

Anomalorhiza shawi 1983 SF, BW Philippines 

Cubozoa 

Carybdea sivickisi 1996 BW Indo-Pacific 

Scyphozoa 

Ulmaridae 

Aurelia sp. 

Introduced 

= Aurelia labiata of Hawaiian authors, not of Chamisso & Eysenhardt, 1821 

This often abundant shallow-water moon jellyfish was unknown to C.H. Edmondson (1933, 1946a), 

the premier marine invertebrate zoologist of the Islands from the 1920s to the 1950s, and would not


45 

have escaped his notice. It first came to general attention in the islands when Chu & Cutress (1954) 

reported it (as Aurelia labiata) to be “common the year round in bays and salt-water canals”, relative 

to reports of human dermatitis (“swimmer’s itch”) in Hawai‘i. Based upon Chu & Cutress’s 

report, presented at the 1953–1954 annual meeting of the Hawaiian Academy of Sciences, we interpret 

the first record to be in 1953. Devaney & Eldredge (1977), reporting it as “possibly A. labiata,” 

note that “it is seen, sometimes frequently, in harbors or inshore areas.” 

Wrobel & Mills (1998) note it as A. labiata from Honolulu, and Gershwin (2001, p. 115) also 

provisionally assigned Hawaiian Aurelia to this species. Dawson et al. (2005), based upon genetic 

comparisons of world-wide Aurelia populations, assign Hawaiian populations to a probably 

unnamed Aurelia (“Aurelia species 4”) from the Indo-Pacific and specifically endemic to the region 

around eastern Borneo and Palau. 

W. Cooke (pers. comm., July 1996) alerted us to this introduction, noting that given the biology 

and life history of both the adult and juvenile stages it could not survive transoceanic transport to 

the Islands naturally. We regard it as introduced, either as ship-fouling scyphistomae or as ephyrae 

in ballast water. Bishop Museum material includes 1963–1964 collections from Pearl Harbor, 

Kāne‘ohe Bay, and Kewalo Basin, all on O‘ahu. Hoover (1998, 2006) illustrates a specimen (as 

Aurelia aurita) from Ala Wai Boat Harbor, Honolulu, a location also noted by Coles et al. (1999b). 

Cassiopeidae 

Cassiopea andromeda (Forsskål, 1775) 

Introduced 

Pacific basin Cassiopea have been reported from Hawai‘i under two separate names, Cassiopea 

medusa Light, 1914 and Cassiopea mertensii Brandt, 1835, both of which are synonyms of C. 

andromeda (reviewed in Holland et al., 2004). Cooke (1984) noted that these Cassiopea, with “their 

pseudobenthic habits are the most improbable adult immigrants.” As C. medusa, Chu & Cutress 

(1954) noted that it was “common the year round in bays and salt-water canals.” Cutress (1961) considered 

it to be introduced from the Philippines by ships as hull-fouling scyphistomae to Pearl 

Harbor between 1941–1945. Carlton (1985) suggested that an alternative mechanism of introduction 

was as ephyrae in ballast water. It was restricted to Pearl Harbor until about 1950 when it appeared 

in Honolulu Harbor and Ala Wai Canal (Cutress, 1961). As Cassiopea mertensii Brandt, 1835, 

Uchida (1970) reported it from “the sandy bottom at a depth of 2 feet from Kaneche [Kāne‘ohe] 

Bay,” O‘ahu. This material was collected in February 1964 (Bishop Museum collections). Cassiopea 

were seen in the early 1990s in fishponds on Moloka‘i and in fishponds in Waikoloa area of the 

island of Hawai‘i (G. Crow, pers. comm., 2000). Hoover (1998, 2006) illustrates a specimen (as C. 

medusa) from the lagoon at Magic Island, O‘ahu. DeFelice et al. (2001) note that it now occurs 

“throughout main Hawaiian Islands.” 

Holland et al. (2004) confirm the identity of C. andromeda based upon molecular evidence 

from the Hawaiian Islands. Cassiopea andromeda is widely distributed in the Indo-West Pacific; distant 

records are the Red Sea (from where it was described in the 18th century) and the western 

Atlantic Ocean (Florida Keys and Bermuda), to both of which regions it may be introduced. 

Cassiopea sp. 

Introduced 

Holland et al. (2004) report that a second genetically distinct species of introduced Cassiopea 

(“Cassiopea species 3”) also occurs in the Hawaiian Islands. This species of Cassiopea also occurs 

in the Papua New Guinea region. Hawaiian material was collected on windward O‘ahu from Kahuku 

fish pond, the Wedding Chapel, Mid Pacific Golf Course pond, and Kualoa Ranch. The date of collection 

of this material was 2000 (B.S. Holland, pers. comm., January 2007). 

Magistiidae 

Phyllorhiza punctata von Lendenfeld, 1884 

Introduced 

= Cotylorhizoides pacificus of Cutress in Doty, 1961 (see Devaney & Eldredge, 1977, p. 114, and Cooke, 

1984, p. 587) 

Under the name Cotylorhizoides pacificus, Cutress (1961) suggested that this Indo-Pacific jellyfish 

was introduced from the Philippine Islands, as ship-fouling scyphistomae, into Pearl Harbor between

46 


1941–1945. It was, however, present in Pearl Harbor by 1933, as described and photographed by 

Edmondson (1933; see Devaney & Eldredge, 1977, who noted that the undetermined rhizostomid of 

Edmondson “certainly appears to be P. punctata”; Edmondson’s species was also noted as being P. 

punctata by W.J. Cooke in an unpublished bibliography of Hawaiian coelenterates compiled in 1980 

and on file in the Bishop Museum’s Department of Invertebrate Zoology reprint library. Edmondson 

(1933) wrote a detailed description of the jellyfish along with a photograph, and noted that “At certain 

times of the year, usually during the winter months, a large undetermined species of jellyfish is 

abundant in Pearl Harbor, Oahu ... large specimens exceed 12 inches in diameter.” 

It may have been restricted to Pearl Harbor until about 1950; in 1953–1954 it appeared in 

Kāne‘ohe Bay (Cutress, 1961). Carlton (1985) suggests that an alternative mode of introduction 

would be as ephyrae in ballast water. Cooke (1984) feels that the taxonomy of the Hawaiian population 

was unresolved, and that it should be referred to as a “mastigid,” but that “any attempt at more 

precise identifications... would be unwise.” We provisionally retain the name used in Devaney & 

Eldredge (1977). 

Clarke & Abey (1998) note that in Kāne‘ohe Bay on “several occasions during 1968–1970 .... 

approximately 20 [to] 40 cm [in diameter] medusae were visible roughly every 10 m across open 

areas of the bay, usually with a cluster of small yellowtail scad [Atule mate] associated with each 

one. The rather sudden disappearance of P. punctata from Kāne‘ohe Bay in the early 1970s thus 

could have a major effect.” 

It was recorded again in Pearl Harbor in 1971–1973 (Evans et al., 1974, as P. punctata). De - 

vaney & Eldredge (1977, as P. punctata) noted that it “occurs, sometimes commonly, in Pearl 

Harbor, Honolulu Harbor, and Kaneohe Bay.” They further note that the undetermined rhizostomid 

of Edmondson (1933, 1946a) abundant in Pearl Harbor during the winter “certainly appears to be P. 

punctata.” 

Hoover (1998, 2006) illustrates a specimen from the Waikīkī Aquarium, noting that “these 

impressive jellyfishes can be common in Honolulu Harbor, Pearl Harbor and Kāne‘ohe Bay, O‘ahu, 

mainly during the winter months.” DeFelice et al. (2001) listed Pearl and Honolulu Harbors, Ala Wai 

Canal and Yacht Harbor, and Kāne‘ohe Bay as the known O‘ahu localities. 

Graham et al. (2003) have reviewed the global invasion history of P. punctata, and taxonomic 

challenges associated with this jellyfish. It has also been introduced to California, the Mediterranean 

Sea, the southwest Atlantic Ocean, and the southern Caribbean Sea, and, since 2000, the Gulf of 

Mexico. Dense populations in the northern Gulf of Mexico in the summer of 2000 impacted shrimp 

fisheries. 

Anomalorhiza shawi Light, 1921 

Introduced 

Cooke (1984) noted that this species, otherwise known from the Philippine Islands, was collected in 

1983 and 1984 in Kāne‘ohe Bay, O‘ahu. He suggested it was introduced as scyphistomae in ships’ 

fouling communities. Carlton (1985) suggested it may also have been introduced as planktonic 

ephyrae in ships’ ballast water. In February 2001 another specimen was collected in Kāne‘ohe Bay, 

approximately 60 cm in diameter; a photograph appeared in The Honolulu Advertiser for February 

16, 2001. This jellyfish remains unknown from any other locations in the Islands and is considered 

“extremely rare” (T. Theeger, pers. comm., February 1998). 

Class Cubozoa (cubomedusae) 

Carybdeidae 

Carybdea sivickisi Stiasny, 1926 

Introduced 

This small (14 mm) but distinctive cubomedusa (box jellyfish) was first collected in 1996 at Waikīkī 

in O‘ahu (Matsumoto et al., 2002). We suggest that it would not have been overlooked by 

Edmondson, who collected and noted gelatinous zooplankton between the 1920s and 1950s. 

Described from the Philippines, it has since been found in Thailand, Japan, New Zealand 

(Matsumoto et al., 2002), Vietnam (Hartwick, 1991), Australia (Hartwick, 1991), and Guam 

(Gershwin, 2003). On O‘ahu it is found, curiously enough, in a saltwater swimming pool known as 

the Natatorium (opened in 1927), fed by ocean water. It has also been found outside of Yokohama


47 

Bay on the west coast of O‘ahu (1998). Crow et al. (2006) report it from Mā‘alaea Boat Harbor, 

Maui, based upon night-lighting collections in 2005 and 2006. 

Carybdea sivickisi, which can cause a painful sting, is benthic during the day and in the water column 

at night (Hartwick, 1991). We suggest ballast water transport, perhaps facilitated by the uptake of 

medusae during the periodic presence of breeding swarms (http: //www.ucmp.berkeley.edu/ 

cnidaria/C_sivickisi.html). 

CTENOPHORA (comb jellyfish) 

Introduced Ctenophora 


Vallicula multiformis 1992 BW Caribbean 

Coeloplanidae 

Vallicula multiformis Rankin, 1956 

Introduced 

The report of Eldredge & Miller (1995) of a nonindigenous ctenophore in Hawaiian waters is based 

upon the collection in 1994 of this species in Hawai‘i by Charles Galt. He found this benthic, creeping 

platyctene comb jelly in Kāne‘ohe Bay, associated with the algae Acanthophora, Bryopsis, 

Kappaphycus, and Caulerpa (C. Galt, in litt., August 1994). In the summer of 1994, they reached 

densities of over 3,000 per square meter. It was present on the dock algae at Coconut Island at least 

by 1992 (C. Fiedler, pers. comm. to C. Galt; C. Galt, in litt., August 1996). Vallicula occurs both in 

shallow fouling communities and is known to be transported with algae in the aquarium trade. It also 

appeared in San Diego, California in 1997 (G. Matsumoto, pers. comm., 1997). Vallicula multiformis 

was originally described from the Caribbean, in Jamaica. 

NEMATODA (roundworms) 

Introduced and Cryptogenic Nematoda 



Camallanus cotti 1993 R Asia 


Spirocamallanus istiblenni 1962 R? 

Order Secernentea 

Camallanidae 

Camallanus cotti Fujita 1927 

Introduced 

This Asian nematode—a “pernicious helminth” (Font,1997a)—occurs in the introduced euryhaline 

mosquitofish Gambusia affinis and the introduced freshwater poecilid (swordtail) Xiphophorus helleri 

in Hawai‘i (Font, 1997a, 1997b; Font & Tate, 1994). Records include it occurring in X. helleri 

in June 1993 in Hakalau Stream, Hawai‘i. Englund et al. (2000a) reported it from additional fish 

hosts in Pearl Harbor, including brackish water sites. 

Spirocamallanus istiblenni Nobel, 1966 

Cryptogenic 

Nobel (1966) described this nematode as a new species from the native zebra blenny (pao‘o), 

Istiblennius zebra, collected on O‘ahu in 1962 and 1963. It also occurs in the native sleeper Eleotris

48 


sandwicensis (Font,1997a). While S. istiblenni was assumed to be native, Rigby & Font (1997) subsequently 

found it in Moorea and Fiji in marine fishes, including the bluestripe snapper (ta‘ape) 

Lutjanus kasmira. Because ta‘ape were introduced to Hawai‘i, “it is problematic whether (this nematode) 

should be regarded as a native or exotic species” in the Hawaiian islands (Font, 1997a). Font 

(1997b) treated it as “native.” Font & Rigby (2000) subsequently found this nematode in Lutjanus 

kasmira from Hilo Bay and conclude that it is cryptogenic. They suggest that examination of museum 

specimens of native freshwater fish from Hawai‘i collected prior to the introduction of ta‘ape in 

1958 may be able to resolve whether this parasite is native or not. 

ROTIFERA (rotifers) 

Cryptogenic rotifers in brackish water of Maui 

(data from Jersabek, 2003; designation as cryptogenic herein) 

Species Location* Notes 

Brachionus plicatilis (Müller, 1786) CP, KeP, KaP A probable species complex 

Colurella adriatica Ehrenberg, 1831 

CP KaP 

Hexarthra oxyuris (Sernov, 1903) KeP Previously recorded from an 

O‘ahu brackish pool by Hauer, 

1941, as Pedalia fennica 

(Levander, 1892) (Jersabek, 

2003). 

Lecane hastata (Murray, 1913) 

CP 

Proalides sp., cf. P. wulferti Sudzuki 1959 CP, KaP A Palaearctic (Europe, North 

Africa, and Asia north of the 

tropics) species. 

* CP, Coot Pond; KeP, Kealia Pond; KaP, Kanahā Pond 

Turner (1996) and Jersabek (2003) present lists of the rotifers of the Hawaiian Islands. Many of these 

are “cosmopolitan” taxa and bear European names. It seems probable that a number of these are 

introduced, but data remain too limited to make such determinations. We offer a table here of five 

brackish-water species (from Jersabek’s work) that may be considered cryptogenic. All are considered 

“cosmopolitan” with the exception of Proalides wulferti, which was “previously known from 

the Palaearctic region only but may have been confused with P. tentaculatus de Beauchamp” 

(Jersabek, 2003). All but Hexarthra oxyuris were new records for the islands, based upon 2001 collections, 

but all of these species were doubtless present much earlier. As Jersabek notes, possible 

transport vectors include intentional releases of nonnative fish, clams, or ornamental plants, as well 

as the movement of resting stages with tourist luggage and clothing. To these vectors we add the 

usual shipping vectors for marine and brackish water species. 

PLATYHELMINTHES (flat worms) 

Introduced and Cryptogenic Platyhelminthes 



“Turbellaria” 

Taenioplana teredini 1938 SB Unknown 

Trematoda 

Ascocotyle tenuicollis 1993 R North America 

Monogenea 

Salsuginus seculus


49 

Introduced and Cryptogenic Platyhelminthes (continued) 



“Turbellaria” 

Convolutriloba sp. 1970s BW 

Additional Taxon Treated and Its Status 

Trematoda 

Centrocestus formosanus 

Status: Presence in brackish-marine waters in Hawai‘i not known; see discussion 

Class “Turbellaria” (flatworms) 

Poulter (1987) lists several species of marine flatworms with “cosmopolitan” or disjunct distributions 

(for example, “Brazil and Hawaii”, or “Red Sea and Hawaii”). Karling et al. (1972), for example, 

newly reported the microturbellarian Gyratrix hermaphroditus Ehrenberg, 1831, a “holeuryhaline 

(sic) cosmopolite and ubiquist” from the Pacific Ocean, from the American Pacific coast (as 

“Karling, unpublished’), and from Kāne‘ohe Bay, “on coral reefs and in sand in the tidal zone” at 

Coconut Island, based upon 1969 collections. 

The distribution and systematics of most marine turbellarians are poorly known, and it is thus 

difficult to recognize either introduced or cryptogenic species in the Hawaiian fauna. That noted, 

there has been and remains striking potential for nonnative shallow water flatworms to be transported 

to the Hawaiian Islands in fouling, in ballast water, in ballast rock or sand, and with commercial 

oysters, as well as by other means, and careful attention needs to be paid to the probability of synanthropic 

dispersal for a great many species. 

Order Acoela 

Convolutidae 

Convolutriloba sp. (?) 

Cryptogenic 

Poulter (1987) noted the presence of “?Convoluta sp.” in marine laboratory aquaria in Kewalo Basin, 

as well as in the laboratory aquaria at Coconut Island and in the wild in Kāne‘ohe Bay. Michael 

Hadfield first noted it between 1973 and 1978, but it may have been here much earlier (M. J. Hadfield, 

pers. comm., February 2000). This small (to 6 mm) green-brown-orange worm contains symbiotic 

algae. It is of interest to note that the acoel Convolutriloba longifissura was found in a seawater aquarium 

in Austria which had been set up with “material from the Pacific” (Gschwentner et al., 1999). 

Given the potential for convolutid acoel flatworms to be transported transoceanically (Rivest et 

al., 1999) the human-mediated dispersal of this distinctive, sometimes abundant species is possible. 

Although this conspicuous flatworm was not likely to have been overlooked by C.H. Edmondson in 

his explorations of the fauna for many years between the 1920s and 1950s, we conservatively regard 

it as cryptogenic. 

Order Polycladida 

Euplanidae 

Taenioplana teredini Hyman, 1944 

Introduced 

Hyman (1944) described this commensal flatworm as a new species from Teredo burrows (species 

not identified) from Honolulu Harbor. Edmondson (1945a) reported it as very abundant in Honolulu 

Harbor during 1943–1944, but “conspicuously absent during 1945.” Edmondson (1945b) noted that 

it was first observed in 1938, being recovered from burrows of shipworms in test blocks in Honolulu 

Harbor; he provides a detailed description of the worm and some notes on its biology. Poulter (1987) 

reports it in burrows of Teredo (species not identified) in Honolulu Harbor, Waikīkī, and Kāne‘ohe 

Bay. Bishop Museum collections contain material collected by C.H. Edmondson between 1941 and 

1944 (the latter being the date of the paratypes). Hyman (1944) noted that the entire life cycle of this

50 


flatworm is completed inside the shipworm burrows, where the eggs are laid as well. She speculated 

that Taenioplana, as a shipworm predator, “may constitute an important enemy of this pest.” It is 

likely found wherever shipworms are common and thus should be expected in Pearl Harbor as well. 

Now widely reported from the Atlantic and Pacific Oceans, its origin remains unknown. 

Class Trematoda (flukes) 

Martin (1958) reported the Asian heterophyid trematodes Centrocestus formosanus (Nishigori, 

1924) and Haplorchis taichui (Nishigori, 1924) from freshwater snails in Hawai‘i. Both may use 

poeciliid fish and snails as hosts in their life cycle. To our knowledge neither of these flukes has been 

reported in either brackish or freshwater introduced poeciliids in Hawai‘i. Centrocestus formosanus 

specifically uses the mosquito fish Gambusia affinis and the snail Melanoides tuberculata as hosts 

(Mitchell et al., 2005), both of which are found in brackish waters in Hawai‘i. 

Order Opisthorchiida 

Heterophyidae 

Ascocotyle tenuicollis Price, 1935 

Introduced 

Font (1997a, 1997b) reported this North American helminth parasite from the heart of the introduced 

mosquito fish Gambusia affinis, from specimens from the Island of Hawai‘i, presumably based upon 

1993 collections. This trematode uses introduced melanid snails as their first intermediate host, fishes 

as the second intermediate host, and birds as the definite hosts for the adult parasite. This parasite 

has been found in fish at Seaside, in Hilo, in salinities of 5 ‰ (W. Font, pers. comm., June 2003). 

Class Monogenea (flukes) 

Order Pseudophyllidea 

Dactylogyridae 

Salsuginus seculus (Mizelle & Arcadi, 1945) 

Introduced 

Font (1997a) reported this North American parasite from the gill filaments of the introduced mosquito 

fish Gambusia affinis, from specimens from Hawai‘i (no specific locality records or dates are 

given). This parasite infects mosquito fish in brackish estuaries in Louisiana (W. Font, pers. comm., 

June 2003), and since Gambusia occurs in both freshwater and brackish water in Hawai‘i, we 

include this parasite here. 

Capsalidae 

Neobenedenia melleni (MacCallum, 1927) 

Introduced 

This ectoparasite was reported from introduced tilapia (Oreochromis mossambicus) acclimated to 

full marine water and held in experimental cages in Kāne‘ohe Bay, based upon specimens first noted 

in 1981 (Kaneko et al., 1988). The original fish were collected from “fresh to brackish water” on 

O‘ahu. Kaneko et al. (1988) considered that tilapia, “a euryhaline freshwater fish is not a normal host 

of N. melleni, which apparently maintains itself on natural hosts in Kaneohe Bay” (which have not 

been identified). Based upon the global distributions reported by Whittington & Horton (1996), 

which identify two geographic clusters for this species (the Western Atlantic and the Eastern 

Pacific), with outlier populations in the Red Sea and Hawai‘i, we regard this fluke as not native. 

Class Cestoda (tapeworms) 

Bothriocephalidae 

Bothriocephalus acheilognathi Yamaguti,1934 Introduced 

The well-known Asian fish tapeworm has been moved around the world with carp (Cyprinus carpio) 

aquaculture and by the introduction of poeciliid fishes (Gambusia) for mosquito control (Dove 

et al., 1997; Font, 1997a, 1997b; Font & Tate, 1994). It has been found on the island of Hawai‘i in 

the introduced poeciliids Xiphophorus helleri (swordtail) and Poecilia reticulata (Font & Tate, 1994) 

based upon collections made in 1993. As Poecilia reticulata also occurs in brackish water, we 

include this parasite here.


51 

ANNELIDA (true worms) 

Introduced and Cryptogenic Annelida 



Polychaeta 

Ophryotrocha adherens 1973 SF, BW Europe 

Neanthes arenaceodentata 1950s SF, BW Northwest Atlantic 

Neanthes succinea 1941 SF, BW, R Northwest Atlantic 

Myrianida pachycera 1959 SF, BW Indo-West Pacific 

Eumida sanguinea 1966 SF, BW, R Unknown 

Polydora nuchalis 1988 FO Northeast Pacific 

Polydora websteri 1940s FO Northwest Atlantic 

Streblospio benedicti 1977 SF, BW, R Northwest Atlantic 

Chaetopterus sp. 1960 SF, BW Unknown 

Branchiomma japonica 1946 SF, BW Japan 

Sabellastarte spectabilis 1946 SF, BW Philippines 

Ficopomatus enigmaticus 1937 SF, BW Australia 

Hydroides brachyacanthus 1939 SF, BW Eastern Pacific 

Hydroides cruciger 1936 SF, BW Eastern Pacific 

Hydroides diramphus 1900 SF, BW Western Atlantic? 

Hydroides elegans 1929 SF, BW Indo-Pacific? 

Pomatoleios kraussii 1967 SF, BW Indo-Pacific 

Salmacina tribranchiata 1935 SF, BW Eastern Pacific 

Serpula watsoni 1936 SF, BW Indo-West Pacific 

Janua pagenstecheri 1960s SF, BW Northeast Atlantic 

Pileolaria militaris 1960s SF, BW Northeast Atlantic? 


Oligochaeta 

Bathydrilus adriaticus 1986 BW, SBA 

Pectinodrilus rectisetosus 1987 BW, SBA 

Pectinodrilus molestus 1986 BW, SBA 

Thalassodrilides gurwitschi 1987 BW, SBA 

Smithsonidrilus minusculus 1987 BW, SBA 

Limnodriloides rubicundus 1987 BW, SBA 

Tectidrilus bori 1986 BW, SBA 

Polychaeta 

Spinther japonicus 1976 SF, BW 

Lumbrineris sphaerocephala 1987 SF, BW 

Mesonerilla fagei 1997 BW, SBA 

Platynereis abnormis 1950s? SF, BW 

Namalycastis abiuma 1995 BW, SBA 

Namalycastis hawaiiensis 1900 BW, SBA 

Namanereis amboinensis 1987 BW, SBA 

Namanereis littoralis 1970s BW, SBA 

Capitella sp. 1975 SF, BW 

Malacoceros sp. 1978 SF, BW 

Minuspio sp. 1970s BW 

Armandia intermedia 1979 SF, BW 

Eulaeospira orientalis 1960s SF, BW 

Leodora knightjonesi 1960s SF, BW 

Neodexiospira foraminosa 1960s SF, BW 

Neodexiospira nipponica 1960s SF, BW 

Neodexiospira pseudocorrugata 1960s SF, BW 

Pileolaria pseudoclavus 1960s SF, BW 

Simplicaria pseudomilitaris 1960s SF, BW 

Spirorbis marioni 1960s SF, BW 

Vinearia koehleri 1960s SF, BW

52 




Polychaeta 

Namalycastis brevicornis 1942 BW, SBA Atlantic Ocean 

Status: Deleted 

Namalycastis senegalensis 1942 BW, SBA Atlantic Ocean 


Boccardia proboscidea 1990 R Northeast Pacific 

Status: Intercept 

Clitellata: Leeches 

Myzobdella lugubris 

Status: See discussion 

Class Clitellata 

Oligochaetes 

Erséus & Davis (1989) were the first to suggest that certain members of the marine oligochaete fauna 

of the Hawaiian Islands may have been introduced by human activities, referring specifically to three 

species previously known largely from the warm waters of the Western Atlantic Ocean: Smithson - 

idrilus minusculus (treated in Erséus & Davis as Limnodriloides claviger), L. rubicundus, and 

Tectidrilus bori. However, Erséus & Davis also note the considerable uncertainty in the biogeography 

and systematics of these and other taxa. We thus consider these three, and several others, as 

cryptogenic species. Sand ballast may have been a particularly effective mechanism in the 19th and 

early 20th centuries for the introduction of some of these species. We follow here the updated 

nomenclature for Hawaiian oligochaetes in Erséus et al. (2005). 

Tubificidae 

Bathydrilus adriaticus (Hrabe, 1971) 

Cryptogenic 

Erséus & Davis (1989) were the first to record this sand-dwelling species from Hawai‘i; it was previously 

known from the western Atlantic (Caribbean), Mediterranean, and Persian Gulf. Hawaiian 

records are from O‘ahu (Kāne‘ohe Bay, Mōkapu Point, Maunalua Beach) and Maui (Olowalu) all 

from intertidal sites or shallow subtidal (to 1.5 m), with the exception of a collection at 30 m from 

Mokapu Point. Erséus & Davis consider it “probably cosmopolitan.” The material was collected in 

1986–1987. 

Pectinodrilus rectisetosus Erséus, 1979 

Cryptogenic 

Erséus (1988) reported this species (as Phallodrilus rectisetous) from Paikō Beach, Maunalua Bay, 

O‘ahu (1987 material) from “barely subtidal mixed sand.” It was previously known from Italy, the 

Atlantic coast of France, and Saudi Arabia. 

Pectinodrilus molestus Erséus, 1988 

Cryptogenic 

Erséus (1988) reported this species (as Phallodrilus moletus) from O‘ahu from three stations: Paikō 

Beach, Maunalua Bay (1987) from “barely subtidal mixed sand”, from off Ke‘ehi Lagoon, Hono - 

lulu, in about 70 m in medium sand (1986) and from Kawaiku‘i Beach Park, Maunalua Bay (1987). 

It was previously known from the Caribbean (Belize and Barbados), Florida, Bermuda, Great Barrier 

Reef, and Fiji. 

Thalassodrilides gurwitschi (Hrabe, 1971) 

Cryptogenic 

Erséus & Davis (1989) newly recorded this intertidal to shallow subtidal brackish water oligochaete, 

“known from heavily polluted situation(s)”, from O‘ahu (Maunalua Bay, 1987). It was previously 

known from the Black and Mediterranean Seas, the Caribbean, Persian Gulf, and southern China. 

Demopoulos et al. (2007) record it (as “cf. gurwitschi”) from Moloka‘i, as part of the mangrove 

infauna.


53 

Smithsonidrilus minusculus (Erséus, 1983) 

Cryptogenic 

This species was first recorded from Hawai‘i by Erséus & Davis (1989), as Limnodriloides claviger, 

based upon specimens “tentatively identified” as this species and discovered in shallow muddy sand 

in Maunalua Bay, O‘ahu (1987). It is also known from the Caribbean (Bermuda and Belize), Aus - 

tralia, and Hong Kong (C. Erséus, pers. comm., 4 January 2008). 

Limnodriloides rubicundus Erséus, 1982 

Cryptogenic 

As with S. minusculus, Erséus & Davis (1989) tentatively extended the range of this western Atlantic 

species to Hawai‘i, based upon specimens from O‘ahu (intertidal and shallow waters at Maunalua 

Bay and Kāne‘ohe Bay, and from 70 m off Ke‘ehi Lagoon, Honolulu; material collected in 1987). 

Limnodriloides rubicundus was previously recorded from Delaware south to Venezuela and 

Bermuda, with another new record in Australia (Erséus & Davis, 1989, p.97). 

Tectidrilus bori (Righi & Kanner, 1979) 

Cryptogenic 

As with the group of species of Limnodriloides, Erséus & Davis (1989) report this western Atlantic 

species (Florida to the Caribbean, and Bermuda) in the Pacific Ocean based upon Hawaiian material 

collected in intertidal and shallow waters at Maunalua Bay and in 70 m off Ke‘ehi Lagoon, 

Honolulu, in 1986 and 1987. The identification of T. bori is considered to be less tentative than that 

of the other two species. 

Leeches 

Piscicolidae 

Myzobdella lugubris Leidy, 1851 


Font & Tate (1994) and Font (1997a, 1997b) reported the presence of this North American freshwater 

leech on introduced freshwater fish in Hawai‘i. We note it here because of their speculation that 

the private release of blue crabs, Callinectes sapidus, from the Gulf of Mexico, and on which the egg 

cocoons of this leech are found cemented to the carapace, may have led to the introduction of this 

leech (see also Font, 1997a). However, unless a large number of parasitized crabs have been released 

into Hawaiian waters, the number of Callinectes actually found in the wild in Hawai‘i would appear 

to be too few to support this means of introduction of Myzobdella. 

Class Polychaeta (polychaetes) 

The majority of Hawaiian intertidal and fouling polychaetes have not been previously analyzed in 

terms of potential candidates for introduced species, and we can present only a preliminary assessment 

here. Decades and centuries of ship movements can create distributions from the Red Sea and 

Africa to the Hawaiian Islands to South America and into the Atlantic Ocean that would appear to 

represent “natural” dispersal or ancient distributions. In short, a great many taxa must eventually be 

considered cryptogenic. We take a generally conservative view here. A large number of species, even 

those regarded as endemic and known only from the Hawaiian Islands, may also bear further consideration 

as cryptogens, especially for those taxa found in association largely with other introduced 

species. Many nonnative species around the world have first been described in the region to which 

they were introduced. 

Bailey-Brock (1976) has noted that in the Hawaiian Islands, “boat harbors and lagoons have a 

typically rich fauna due to introduction of benthic invertebrates on the bottoms of boats. Such habitats 

remain reservoirs of introduced species, which are important in the geographical distribution of 

tube worms within the islands.” She further noted that in addition to the introduced tubeworm 

Ficopomatus (discussed as Mercierella), “the cosmopolitan fouler Hydroides norvegica has been 

dispersed by boats, and it is quite possible that Hydroides lunulifera, Pileolaria militaris, and others 

have been spread to and between the islands in the same manner.” We discuss these species below. 

Poorly represented in our treatment here are species of Syllidae, Phyllodocidae, and Sabellidae, 

and not represented at all are species of Eunicidae and Cirratulidae. All five of these families are

54 


often well-represented in ships’ fouling communities and ballast water and on commercial oysters, 

making these families rich fields for exploration for cryptogenic and introduced species. Examples 

of genera that should be particularly examined for introduced and cryptogenic species are Phyllo - 

doce, Brania, Exogone, Typosyllis, Marphysa, Cirriformia, and Euchone. 

The Bishop Museum “Progress Report of the Marine Zoology Department” for 1965 stated that 

information was provided to the State Quarantine Station on the “potential harmfulness of Nereis 

japonica and Lumbriconereis japonica if established in Hawaii.” The reason for this concern is not 

given, but it may be that these species were being imported, or were being considered for importation, 

as fish bait. 

Spintheridae 

Spinther japonicus Imajima & Hartman, 1964 

Cryptogenic 

This species was collected in 1976–1977 from sponges in Pearl Harbor and Kāne‘ohe Bay (Grov- 

houg & Rastetter, 1980; Bailey-Brock & Hartman, 1987). Coles et al. (1999b) report it from Ke‘ehi 

Lagoon. It is otherwise known from Japan. We consider it cryptogenic. 

Lumbrineridae 

Lumbrineris sphaerocephala (Schmarda, 1861) Cryptogenic 

Bailey-Brock & Hartman (1987) note that this Indo-Pacific species is “found commonly among fouling 

communities” in Kāne‘ohe Bay. We consider it cryptogenic. Hawaiian material should also be 

compared to the Caribbean Lumbrineris perkinsi (L. Harris, pers. comm., January 2009). 

Nerillidae 

Mesonerilla fagei Swedmark, 1959 

Cryptogenic 

Bailey-Brock (1999) reported this tiny sand-dwelling species, previously known from the Northeast 

Atlantic Ocean (English Channel and Irish Sea) based upon numerous specimens collected in 1997 

from coarse sand in 3 to 5 m in Honolulu Harbor. This remarkably disjunct distribution suggests possible 

human-mediated dispersal, perhaps with sand ballast of the 19th or early 20th centuries. 

Genetic comparisons with European material would be of value. 

Dorvilleidae 

Ophryotrocha adherens Paavo, Bailey-Brock 

Introduced 

& Åkesson, 2000 

Paavo et al. (2000) and Bailey-Brock et al. (2002) reported this species from Sand Island and 

Barbers Point sewage outfalls on O‘ahu (first collections 1973). It also occurs at the Hawai‘i Kai 

Marina (B. Paavo, pers. comm., June 2003). This very small (1.2 mm) worm has likely been long 

overlooked because of its size and because of its potential to be mistaken for the juveniles of other 

dorvilleids. It is otherwise known from the Mediterranean (in the harbor of Kyrenai, Cyprus) and 

from the harbor of Las Palmas, in the Canary Islands. A protandric simultaneous hermaphrodite, it 

has a free-swimming larval stage lasting only up to five days in the water; this, combined with its 

close association with inshore, disturbed, and harbor habitats, suggests transport by shipping to the 

Hawaiian Islands. Populations from O‘ahu have been successfully interbred with European worms, 

indicating that they are the same species (Paavo et al., 2000) further suggesting a lack of genetic 

divergence (as might be expected from natural colonization and genetic isolation) and thus modernday 

human-mediated transport. Bailey-Brock et al. (2002) provide population data off Sand Island 

at 70 m for the period 1990–1998. 

Nereididae 

Neanthes arenaceodentata Moore, 1903 

Introduced 

= Neanthes caudata of authors 

= Neanthes arenaceodonta of authors 

This now widely distributed pileworm (Day, 1967) which may have originated in the North Atlantic 

Ocean, was reported by Hartman (1966) from coral sand in Kāne‘ohe Bay. This material was likely


55 

collected between 1946 and 1966. It was recorded in Pearl Harbor in 1971–1973 (Evans et al., 1974). 

Bailey-Brock & Hartman (1987) note that it “may have been introduced.” Bailey-Brock et al. (2002) 

report it from off the Sand Island (O‘ahu) open ocean sewer outfall, from where it “virtually disappeared” 

after 1993. 

Neanthes succinea Frey & Leuckart, 1847 

Introduced 

This is another Atlantic pile worm, which has been carried around the world with commercial oysters 

and in ship fouling and perhaps in more modern times in ballast water. Material from O‘ahu in 

the Bishop Museum dates from 1941 (O‘ahu), 1945 (off Waikīkī, in about 6 m) and 1944–1945 

(Honolulu Harbor), but it was not reported in the literature until Bailey-Brock & Hartman (1987) 

reported it from the Ala Wai Canal, where it “is abundant in the mud along the banks.” We consider 

it introduced. 

Platynereis abnormis (Horst, 1924) 

Cryptogenic 

This species is known in sponge colonies and coral mud in Kāne‘ohe Bay, likely based upon material 

collected between 1946 and 1966 (Hartman, 1966). Coles et al. (2004) report it from 1997 collections 

by R.C. DeFelice in Mā‘alaea Harbor, and Coles et al. (2006) report it from the island of 

Hawai‘i, based upon 2003 collections. Bailey-Brock & Hartman (1987) note it otherwise “has a 

broad Indo-West Pacific distribution.” We consider it cryptogenic. 

Subfamily Namanereidinae 

Glasby et al. (1998) reported four species of supralittoral namanereids from Hawai‘i. We include 

them here because they may be found in brackish water, such as on mudflats in the mangrove zones 

of estuaries (Glasby, 1999). Because of their potential to have been carried in ship’s shore ballast or 

in modern ballast water on floating algae or bits of small debris, all four species are here considered 

cryptogenic (despite being widespread throughout the Pacific basin). 

Namalycastis abiuma (Grube, 1872) 

Cryptogenic 

Glasby et al. (1998) reports that this is a common species “of the supralittoral zone of mud flats in 

the tropics and subtropics around the world, often associated with mangroves and decaying vegetation 

such as Nypa palms, coconut husks and Enteromorpha (now called Ulva) overlying mud flats. 

On Moloka‘i the species was found [in 1995] in mixed gravel and detritus along the stream edge at 

8–9 m elevation.” Glasby notes that this is the first actual record of this species in Hawai‘i, previous 

records of N. albiuma probably referring to Namalycastis hawaiiensis . 

Namalycastis hawaiiensis (Johnson, 1903) 

Cryptogenic 

Glasby et al. (1998) notes that this species is the “most abundant freshwater namanereid species” in 

the islands. Glasby et al. (1998) noted earlier records from a spring near Honolulu (Johnson, 1903; 

material probably collected about 1900) and, many years later, at the ‘Ewa Plantation on O‘ahu and 

on the Kona coast of Hawai‘i. Glasby et al. (1998) remark that in Hawai‘i “the species occurs in mud 

to muddy-sand sediments of streams, swamps, aquaculture ponds and on the Kona coast in closed 

coastal anchialine ponds; preferred salinities range from fresh to very slightly brackish water... (it) 

is often associated with leaf litter, under stones, coconut husks and under the bark of floating wood 

in areas of heavy nutrient load together with talitrid amphipods, the oligochaete Branchiura sowerbyi 

Beddard and other unidentified oligochaetes.” Whether N. hawaiiensis arrived in the Islands by 

natural drifting or by shipping is not yet known; if it is introduced, it is another example of a species 

first described from other than where it is native. 

Namalycastis hawaiiensis is also known from eastern Asia (Hong Kong and Ryukyu Island), 

southeast Asia (Sumatra and Java) and the western Pacific Ocean (New Guinea; Palau Islands, 

Guam). Glasby et al. (1998) examined specimens from O‘ahu (1987), Kaua‘i (1995), Hawai‘i (1987, 

1995), Moloka‘i (1994, 1995), and Maui (1995).

56 


Namalycastis brevicornis (Audouin & Edwards, 1833) Deleted 

Glasby (1999) notes that this Atlantic species, known from France, French Guiana, and Brazil, is 

represented by verified specimens in the Paris Museum from “Iles Sandwich” which had also been 

examined by Pierre Fauvel in 1942. Glasby noted that the record “probably refers to the Hawaiian 

Islands.” If the record is a valid one, it would appear to be a case of an introduction, possibly by 

ship’s shore ballast or in ballast water. 

Namalycastis senegalensis (Saint-Joseph, 1900) Deleted 

Glasby (1999) notes that this Atlantic species, known from West Africa and Brazil, is represented by 

verified specimens in the Paris Museum from “Îles Sandwich” which had also been examined by 

Pierre Fauvel, probably about 1942. Glasby notes that the record “probably refers to the Hawaiian 

Islands” (see Glasby, 1999, pp. 42 and 70). If the record is a valid one, it would appear to be another 

case of an introduction, possibly by ship’s shore ballast or in ballast water. 

Namanereis amboinensis (Plugfelder, 1933) 

Cryptogenic 

This worm, already widely recorded from the upper littoral zone of the tropics and subtropics 

(Glasby et al., 1998) was newly discovered in the Hawaiian archipelago with a collection in 1987 of 

specimens in He‘eia Stream, O‘ahu. It is “found in the upper littoral zone of mangroves (together 

with Namanereis littoralis) and under the bark of logs floating in fresh-brackish water (together with 

talitrid amphipods and Namalycastis hawaiiensis )” (Glasby et al., 1998). The record is also noted 

by Glasby (1999). 

Namanereis littoralis (Grube, 1872) 

Cryptogenic 

Glasby et al. (1998) note that this “cosmopolitan” species was previously recorded from open ponds 

on the Kona coast of Hawai‘i as Namalycastis sp., based upon material collected in the early 1970s. 

“The present material extends its Hawaiian distribution to O‘ahu where it occurs behind the mangrove 

(Rhizophora) zone in muddy sand with surface detritus together with the more numerous 

species Namanereis amboinensis . On the Kona coast of Hawai‘i the species occurs in anchialine 

ponds under stones at the water’s edge.” Glasby et al. (1998) reported specimens from near 

Anaeho‘omalu Bay, 1987 (Anaeho‘omalu Stream, pond) and from He‘eia Stream, O‘ahu, 1987. 

Glasby (1999) also notes this record. 

Syllidae 

Myrianida pachycera (Augener, 1913) 

Introduced 

= Myrianida crassicirrata Hartmann-Schröder, 1965, described from O‘ahu 

Nygren (2004) has suggested, based upon molecular genetic data, that this spectacular Western and 

Indo-Pacific worm has been introduced to various harbors around the world (outlier sites include 

Hawai‘i and Fort Pierce, Florida). It has been collected in ship’s fouling in Los Angeles Harbor as 

well (Nygren, 2004). It was first collected in 1959 in Kāne‘ohe Bay, where it is described as “common 

... (in) coral rubble, mud, algae, and living sponges” (Friend, 1987). It is also known from Pearl 

Harbor (photograph by J. Grovhoug, “swimming over fouling community,” on Plate 3 in Devaney 

& Eldredge, 1987), where it was first collected in 1976–1977 (Grovhoug & Rastetter, 1980). We 

regard it as a post-World War II invasion by ship fouling or ballast water. 

Phyllodocidae 

Eumida sanguinea (Oersted, 1843) 

Introduced 

= Eulalia sanguinea 

Hartman (1966) reported this species as being collected in June, 1966 in blisters in the introduced 

Atlantic oyster Crassostrea virginica in Pearl Harbor; the record is repeated by Bailey-Brock & 

Hartman (1987). Coles et al. (1999a) report it again from Pearl Harbor based on 1996 collections. 

Coles et al. (2002a) record it from Kāne‘ohe Bay. 

As this species appears to be harbor-restricted and occurs in a location heavily subjected histor-


57 

ically to both the introduction of oysters and shipping, we consider it introduced. It is a very widely 

reported cosmopolitan species (Day, 1967), suggesting that more than one species is likely involved. 

Capitellidae 

Capitella sp. 

Cryptogenic 

Ward (1978) first recorded Capitella capitata from Hawai‘i on the basis of larvae from the Ala Wai 

Canal and Kāne‘ohe Bay collected in 1975 or 1976. Bailey-Brock (1984) found it in densities up to 

11,600 per square meter in the sand beds of the onuphid worm Diopatra leuckarti at Niu Valley on 

the south shore of O‘ahu. Bailey-Brock & Hartman (1987) noted that adult Capitella capitata have 

“been found in large numbers in sediments on fringing reefs near stream outlets in Kāne‘ohe Bay, at 

Niu near Aina Haina, in oyster culture ponds, and from other areas around O‘ahu.” Bailey-Brock 

(1990) reported it as co-occurring in Hawai‘i with Polydora nuchalis in oyster raceways and phytoplankton 

ponds at an oyster pond (“The capitellid is very hardy and survived in damp cracks and fissures 

in the floor of the ponds when they were drained and left to dry out.”). Coles et al. (2002b) 

report it from Waikīkī. Bailey-Brock et al. (2002) provide population data off Sand Island at 70 m 

for the period 1990–1998. 

It is curious that there are no earlier records of this taxon in the Islands, especially from Pearl 

Harbor. Capitella capitata is now regarded as a species complex, and we use the designation Capi - 

tella sp. We consider it cryptogenic. 

Spionidae 

Polydora nuchalis Woodwick, 1953 

Introduced 

Bailey-Brock (1990) reports that this species, previously known from central California to the Gulf 

of California, was found in June 1988 at Kahuku, O‘ahu, in an Atlantic Oyster (Crassostrea virginica) 

farm. The oysters were being grown in the effluent water from a commercial shrimp 

(Litopenaeus vannamei) farm. The worms may have been introduced either with the oysters (both C. 

virginica and the Pacific oyster C. gigas having been imported from both the east and west coasts of 

North America, respectively) or with shrimp (from north of Guaymas in the Gulf of California and 

other locations in Central America and Asia). 

Polydora websteri Hartman, 1943 

Introduced 

= Polydora ciliata of authors 

This well-known Northwestern Atlantic “mudworm” was first found in the Hawaiian Islands by D.P. 

Abbott (1946) who reported it (as Polydora ciliata) from Wailupe Pond, O‘ahu; he also noted that it 

was very abundant at Kuapā Pond. No dates are given, but the material would have been collected 

in the 1940s. Hartman (1966) recorded it from mud blisters in shells of the Atlantic oyster 

Crassostrea virginica collected in 1966 in Pearl Harbor. Bailey-Brock & Hartman (1987), citing 

Smith et al. (1977) noted that large numbers of larvae were found in the stomach of the sergeant fish 

Abudefduf in Kāne‘ohe Bay. 

Bailey-Brock & Ringwood (1982) reported that this worm periodically infested a land-locked 

intensive Crassostrea gigas oyster farm at Kahuku, O‘ahu where it “may have been inadvertently 

introduced to the Kahuku system with oysters transplanted from Kāne‘ohe Bay or with oyster spat 

imported from U.S. west coast hatcheries” The oyster farm eventually ceased operation (Eldredge, 

1994). Ward (1987) reviewed these records, noting that it was “detrimental to the oyster mariculture 

industry in Hawaii”, and further reported it from coral rock in Kāne‘ohe Bay and Ala Wai Canal, as 

well as from mud in Nu‘upia Ponds, O‘ahu; planktonic larvae were found in both Kāne‘ohe Bay and 

the Ala Wai Canal. 

Boccardia proboscidea Hartman, 1940 

Intercept 

This North Pacific worm was found in March 1990 in newly imported Atlantic Oysters (Crassostrea 

virginica) from Maine at an oyster culture facility in Keahole, Hawai‘i (Bailey-Brock, 2000). The 

oysters were held in open raceways with flowing seawater released to the sea, but the fate of these 

worms (which included egg capsules with larvae) is not known. Curiously, B. proboscidea was not

58 


previously known from the Atlantic Ocean, and is thus likely an introduction to Maine as well. It is 

also introduced to Australia and possibly to Japan (records reviewed in Bailey-Brock, 2000). 

Streblospio benedicti Webster, 1879 

Introduced 

This introduced species was found in 1977 (Ward, 1981) in mud in Hālawa Stream, draining into 

Pearl Harbor (Ward, 1981, 1987). Englund et al. (2000a) reports it from additional estuarine streams 

in Pearl Harbor. It may have been introduced from either the Atlantic coast of America or from the 

Pacific coast, to where it has also been introduced (Cohen & Carlton, 1995). 

Malacoceros sp. 

Cryptogenic 

Ward (1987) reports a Malacoceros species from estuarine mud banks in the Ala Wai Canal (collected 

in 1978, fide Bishop Museum collections). This location is characterized by numerous invasions. 

We regard it as a cryptogenic species. 

Minuspio sp. 

Cryptogenic 

Ward (1987) reports a species of Minuspio from Kāne‘ohe Bay, Barbers Point, Pearl Harbor, and 

from sediment and coral rubble in experimental microcosm tanks at Mōkapu Peninsula. These are 

presumably based upon collections made in the mid- to late 1970s. Englund et al. (2000a, p. 109) 

report what is likely the same species from estuarine conditions in Pearl Harbor. 

Opheliidae 

Armandia intermedia Fauvel, 1902 

Cryptogenic 

We consider this opheliid to be cryptogenic, pending its resolution as conspecific with the Atlantic 

species. Bailey-Brock (1987) notes it from bottom samples in Honolulu Harbor and “among mangrove 

roots on a south shore fringing reef where densities reach 589/m 2 ” and in plankton in 

Kāne‘ohe Bay. Englund et al. (2000a, p.109) record it from estuarine waters in Pearl Harbor. Coles 

et al. (2002b) report it from Waikīkī. It is considered to be widely distributed in the Indo-West 

Pacific and Atlantic (Bailey-Brock, 1987) and Australia (Hutchings, 2000). 

Chaetopteridae 

Chaetopterus sp. 

Introduced 

= Chaetopterus variopedatus of Hawaiian authors 

Bailey-Brock (1976) noted that Chaetopterus sp. was abundant in Kāne‘ohe Bay on the alga Dictyo - 

sphaeria cavernosa. Bailey-Brock & Hartman (1987, as C. variopedatus) noted it was also “a frequent 

component of fouling communities.” Grovhoug & Rastetter (1980) report it based on 1976– 

1977 collections from Pearl Harbor and Kāne‘ohe Bay; Coles et al. (1999a) record its continued 

presence in Pearl Harbor (1996 collections). It also occurs in Honolulu Harbor, Ke‘ehi Lagoon, and 

Barbers Point Harbor (Coles et al., 1999b). Hoover (1998, 2006) presents a color photograph of a 

population in Pearl Harbor at 3-m depth. The earliest specimen from the Islands appears to be that 

collected by L.G. Eldredge in May 1960 on Coconut Island, Kāne‘ohe Bay (BPBM R-260). 

Curiously, this large worm was not reported by Hartman (1966), or in the lengthy papers on 

O‘ahu fouling communities by C.H. Edmondson, or in Edmondson’s several books on the Hawaiian 

marine fauna, the latter covering aspects of the biota of the Islands up to the 1940s. These lack of 

early reports of a relatively large (tubes up to 13 cm long), conspicuous, shallow-water invertebrate 

in easily-accessed habitats, combined with its presence in fouling communities, suggest to us that it 

is an introduction. Given the lack of historical records, it may have been introduced in ship fouling 

since World War II. 

Sabellidae 

Branchiomma japonica (McIntosh, 1883) 

Introduced 

= Branchiomma nigromaculata of Bailey-Brock & Hartman, 1987, p. 411, not of Baird, 1865; not Sabella 

havaica Kinberg, 1867, and not Branchiomma cingulata (Grube, 1870) of Hartman, 1966, fide P. Knight- 

Jones, pers. comm., February 2000. 

This medium-sized sabellid of Hawaiian fouling communities was long thought to be the same as 

Sabella havaica Kinberg, 1867, described from the Hawaiian Islands (and collected in 1852), which


59 

would have made it one of the earlier records of an introduced marine invertebrate in the archipelago. 

However, while poorly known, S. havaica is a different species (per the above synonymy), and 

thus the first specimens of B. japonica in the Hawaiian Islands are likely those of D. P. Abbott, collected 

“from scrapings of bottom of naval vessel” in Pearl Harbor (Hartman, 1966, p. 235). No dates 

of collection are given, but Abbott was collecting in the islands from the early to mid 1940s (Abbott, 

1941, 1946). Hartman (1966), under the name “?Branchiomma cingulata“, also noted material collected 

by R. W. Hiatt in 1946 from Halape “in holes, crevices and in matted algae at outer edge of 

rocky shore, 0.0 to 1.0’” (BPBM R504). 

Bailey-Brock (1976) noted that it was only common in boat harbors and lagoons (“abundant on 

hard substrates, piers, buoys, and floating docks in relatively calm waters”) in the Hawaiian Islands, 

although, reminiscent of Hiatt’s record, it occurred at Kapoho, Hawai‘i, on a lava bench with living 

coral. Bailey-Brock (1976) further recorded it from a number of stations around O‘ahu, including 

the Ala Wai Canal. Grovhoug & Rastetter (1980, (as B. cingulata) collected it in 1976–1977 from 

Pearl Harbor and Kāne‘ohe Bay; Henderson (1990) records it in 1986 from the USS Arizona in Pearl 

Harbor, and Coles et al. (1999a) report its continued presence in Pearl Harbor (1996 collections). 

Phyllis Knight-Jones (pers. comm., February 2000) noted that “In Hawai‘i we found it at Koko and 

Coconut Harbors under pontoons, pipes, and fauna-covered ropes.” Coles et al. (1999b) report it as 

B. nigromaculata from Honolulu Harbor, Ke‘ehi Lagoon, Ala Wai Harbor, Ala Wai Harbor, Barbers 

Point Harbor, and Kewalo Basin, and Coles et al. (2002a) record it for Waikīkī. Coles et al. (2006) 

report it from Kaua‘i, based upon 2003 collections. 

Branchiomma japonica is “abundant in Japan” (P. Knight-Jones, pers. comm., February 2000). 

We regard it as introduced, given its largely harbor and fouling habitat in the Islands and the highly 

disjunct distribution between Japan and Hawai‘i. 

Sabellastarte spectabilis (Grube, 1878) 

Introduced 

= Sabellastarte sanctijosephi of Hawaiian authors, not Gravier, 1906, vide Knight-Jones & Mackie, 2003. 

= Sabellastarte indica Savigny,1822 (name pre-occupied, vide Knight-Jones & Mackie, 2003) 

This large and distinctive fanworm (also known as a feather-duster worm) has had a complex taxonomic 

history. Curiously, it seems not to be mentioned in the Hawaiian biota until after World War 

II: there is no material in Bishop Museum collections prior to 1946, nor is there any mention of it in 

the literature until 1966. It does not appear (either by name or by description) in Edmondson’s books 

(1933, 1946a), nor is it mentioned in the early fouling studies of Pearl Harbor, an environment with 

which Edmondson was very familiar. On the assumption that such a large, colorful, and conspicuous 

fouling species would not be overlooked, we treat it here as introduced. 

It has been treated in the Hawaiian literature as either Sabellastarte indica (in Hartman, 1966) 

or Sabellastarte sanctijosephi (in Bailey-Brock, 1987). Knight-Jones & Mackie (2003) reidentify 

Hawaiian material as S. spectabilis, a species from the Philippines. Hartman (1966) noted that R.W. 

Hiatt collected it in 1946 at Halapē, Hawai‘i, from a wide variety of coastal habitats (for example, 

“in holes, crevices, and in matted algae at outer edge of rocky shore;” from the interstices of the coral 

Pocillopora meandrina; from under boulders in quiet water, in crevices in lava in open coast tide 

pool, and even from a tidal channel exposed to heavy surf). Hartman (1966, p. 239, also as S. indica) 

referred material from “Puata Bay and Waialea (?) [sic], O‘ahu” collected in 1902 by the U.S.S. 

Albatross, but given the absence of any other records prior to 1946, as noted above, and its failure 

to be detected by Edmondson and others, we regard this material as requiring re-examination. 

Bailey-Brock (1976) noted that it was “found in pockets and crevices in the reef flat. It is especially 

abundant along the edges of reefs that have been dredged to make small-boat harbors and 

swimming areas, as at Ala Moana and Fort Kamehameha, O‘ahu; it may be an indicator of waters 

with high sediment content.” She further characterized it (Bailey-Brock, 1976) as a species of “calm 

waters in harbors and marinas, along the edges of reef pockets, in dredged areas.” A specimen of this 

worm was found on the snail Conus quercinus collected in 2 m at Sand Island Reef, Kāne‘ohe (A. 

J. Kohn in Bailey-Brock, 1976). 

Bailey-Brock & Hartman (1987, p. 220) repeats the fact that “the walls of dredged channels in

60 


shallow or intertidal waters are frequently populated” by this worm, and noted (p. 416) that it was 

“abundant on Oahu’s south shore reefs, and in Pearl Harbor and Kaneohe Bay at shallow depths, 

especially in dredged areas that receive silt-laden waters. They have also been found at depths of 25 

to 30 m off Molokini Island, near Maui.” Grovhoug & Rastetter (1980) record it from Pearl Harbor 

and Kāne‘ohe Bay, based on 1976–1977 collections. Coles et al. (1999a) record it from Pearl Harbor 

as well. Coles et al. (1999b) report it from Honolulu Harbor, Ke‘ehi Lagoon, Ala Wai Harbor, Bar - 

bers Point Harbor, and Kewalo Basin. Coles et al. (2006) report it from Kaua‘i, based upon 2002 

collections, and from 2003 collections in Kaunakakai Harbor, Moloka‘i, Kahului Harbor, Maui, and 

Kawaihae and Hilo Harbors, Hawai‘i (Coles et al., 2004). 

Bybee et al. (2006a, b, 2007) describe the reproductive and larval biology of Sabellastarte in 

Kāne‘ohe Bay, noting that it is a sequential protandrous hermaphrodite. 

Walsh et al. (2003) noted that between 1976 and 2003, no fewer than 741,949 specimens of this 

brightly-colored worm were harvested for the aquarium trade, for a value of $860,362. Bybee et al. 

(2006b) remark that it “has become one of the most harvested marine ornamental species” in Hawai‘i, 

and that “corals and other components of the reef community are often damaged by collectors.” 

Bailey-Brock & Hartman (1987, plate 3.II.4.b) present a color photo of the tentacular crown. 

Hoover (1998, 2006) also presents a color photograph of individuals amongst corals, at approximately 

12 m at Kewalo Pipe, O‘ahu. 

Serpulidae 

Bastida-Zavala (2008) noted that additional species, which he formally recorded as now occurring 

in Hawai‘i, were found on incoming vessels (Godwin, 2003a; Godwin et al., 2004). These included 

Pomatoceros minutus Rioja, 1941 (Mexico to Peru) and Protula atypha Bush, 1905 (California to 

Mexico). These are intercepts, and we do not further treat them here. 

Long (1974) reported Hydroides sanctaecrucis Kroyer in Morch, 1863, a Caribbean species, 

from panels deployed a short distance offshore from O‘ahu, in 15 m, between 1968 and 1972. Lewis 

et al. (2006) note that the introduction of H. sanctaecrucis to the Pacific coasts of Panama and 

Mexico, to Australia and to Singapore “adds some credibility” to Long’s record. It has not been further 

reported in Hawai‘i, despite extensive studies by M.G. Hadfield and his students on the serpulids 

of Pearl Harbor. We await further material, but at this time do not further consider it. 

Ficopomatus enigmaticus (Fauvel, 1923) 

Introduced 

= Mercierella enigmatica 

This often abundant tubeworm first appeared in the northern hemisphere in the early 1920s in both 

San Francisco Bay, California, and in western France (Carlton, 1979a). It is thus of interest to note 

its establishment within about 15 years of these dates in Hawai‘i. The Hawaiian populations may 

have been derived from San Francisco Bay; a genetic comparison of these two populations would be 

of interest. Straughan (1969a) noted records from Pearl Harbor in 1937, Kewalo Basin (1947), and 

Alai [Ala] Wai Canal (1947), based on Bishop Museum collections. Hartman (1952) had earlier 

reported it from the Ala Wai Canal based on 1948 collections, and later (Hartman, 1966) recorded 

again the 1947 material noted by Straughan from the Canal. Edmondson (1946a, p. 114) noted “an 

undetermined species of the genus Mercierella ... abundant on the bottom of a boat at Waikiki” (this 

statement does not appear in Edmondson’s first edition in 1933). Edmondson’s figure (p. 116, fig. 

53d) shows a distally asymmetrical operculum with a ring of 22 short, straight spines around the 

outer edge. This compares reasonably well with the variable opercular spination of Ficopomatus 

enigmaticus and its characteristic operculum with a “distal eccentrically placed concavity” (ten Hove 

& Weerdenburg, 1978). Evans et al. (1972) also reported it from Pearl Harbor. 

Bailey-Brock (1976) noted that Ficopomatus is present along the length of the Ala Wai Canal: 

“the most extensive population is seen in the section between the Manoa-Palolo channel and 

Kapahulu.” She also reported Ficopomatus on O‘ahu at the beach park at Paiko, in canals of Hawai‘i 

Kai, and in a drainage canal at Kahala. Bailey-Brock (1976) further reported that “On Maui, extensive 

masses of Mercierella were seen at a commercial fish farm located at Keālia Pond. It is believed


61 

that Mercierella was introduced to Maui with oysters sent from Pearl Harbor to stock the pond. 

These serpulids may pose a threat to aquaculturists because they are so easily introduced and are able 

to withstand considerable exposure to freshwater. They frequently clog the pipes used for circulation 

between the holding ponds and cover settlement sites.” 

Hydroides brachyacanthus Rioja, 1941 

Introduced 

= Hydroides brachyacantha 

This serpulid is known from Mexico to Ecuador and the Hawaiian Islands (Bastida-Zavala & ten 

Hove, 2003), and perhaps elsewhere, such as the Caribbean, Brazil, Australia, Chuuk, and other Indo- 

Pacific locations (Straughan, 1969a; Bailey-Brock & Hartman, 1987; H. ten Hove, pers. comm., 

1996). Bastida-Zavala & ten Hove (2003) question its circumtropical and circumsubtropical status; a 

species complex may be involved. 

Records are scattered and few in the Islands: 1939 (Black Point; Straughan, 1969a), Halapa 

(1946), and Kahulai, Maui (1973), all representing Bishop Museum material. Bailey-Brock (1976) 

notes that she did not encounter it and that it “is probably rare.” Bastida-Zavala & ten Hove, 2003 

examined Straughan’s 1972 collections from O‘ahu. Coles et al. (1999b) record it from Honolulu 

Harbor, Coles et al. (2002a) from Kāne‘ohe Bay, and Coles et al. (2004) from Kahului Harbor, Maui. 

We consider this species to be introduced by ship-fouling from the warm waters of the Eastern 

Pacific; it was found alive on a vessel arriving in Australia that had likely become fouled in Panama 

or Mexico (H. ten Hove, pers. comm., 1996). Bastida-Zavala & ten Hove (2003) note morphological 

differences from mainland populations. These differences could conceivably arise if H. brachyacanthus 

has been long isolated in the islands. 

Hydroides cruciger (Mörch, 1863) 

Introduced 

= Hydroides crucigera 

A species otherwise known from Mexico to Colombia (Bastida-Zavala & ten Hove, 2003), we regard 

this as another of the introduced Hydroides “facies.” Straughan (1969a) notes records from Pearl 

Harbor (1937, 1938), Kāne‘ohe Bay (1936, 1937, 1938) and Coconut Island (1968). Long (1974) 

found it on “offshore” and Pearl Harbor panels (1968–1972). Coles et al. (2002a) report it from 

Waikīkī. Coles et al. (2004, 2006) report it from Maui, based upon 2003 collections. Bastida-Zavala 

(2008) noted it as present on a vessel inbound from the American Pacific coast to Hawai‘i. 

Bailey-Brock (1976) noted that it has been found as an epizoic on the native snail Charonia tritonis 

in 6 m of water off Waikīkī, O‘ahu, and that it was also found on rubble subtidally and was less 

common than H. norvegica and H. lunulifera. 

Hydroides diramphus Mörch, 1863 

Introduced 

= Hydroides lunulifera, H. lunifera, and H. lunifer of Hawaiian authors; = Eupomatus lunifer (see Bailey- 

Brock & Hartman, 1987). 

This is a species possibly of tropical American Atlantic origin (H. Zibrowius, pers. comm., 1996; see 

also Zibrowius, 1973); it now occurs world-wide circumtropically and circumsubtropically. In the 

islands, Edmondson & Ingram (1939) reported it from Kāne‘ohe Bay in fouling (1935, as H. lunulifera). 

Hartman (1966, as Eupomatus lunifer) reported a “new record” as follows: “Honolulu yacht 

harbor, May, 1900, collected by Dr. Loye Miller” (these specimens were examined again by Bastida- 

Zavala & ten Hove, 2003). Straughan (1969a, as H. lunulifera) reports records from Pearl Harbor 

(1929, 1937), Kāne‘ohe Bay (1936–1937), Kewalo Basin (1943) and Coconut Island (1968). Long 

(1974) found it on both “offshore” and Pearl Harbor panels (1968–1972); McCain (1975) further 

reported it on fouling panels in Pearl Harbor (1973), where it continues to be found (Coles et al., 

1999a). Additional Bishop Museum material includes specimens from the Ala Wai Canal (1968). 

Coles et al. (1999b) record it from Honolulu Harbor, Ke‘ehi Lagoon, Ala Wai Harbor, and Barbers 

Point Harbor, while Coles et al. (2004) record it from Nawiliwili Harbor, Kaua‘i and Hilo Harbor. 

This serpulid has been found alive on a vessel after a sea voyage (H. ten Hove, pers. comm., 

1996). Bailey-Brock (1976) notes that it is “quite possible” that it may “have been spread to and

62 


between the islands” by ships. Bastida-Zavala (2008) reports it fouling on inter-island barges in 

Hawai‘i. 

Hydroides elegans (Haswell, 1883) 

Introduced 

= Hydroides norvegica of authors, not of Gunnerus, 1768 

Tubeworms under the name “Hydroides norvegica” were recorded around the world until the 

species-level taxonomy was first clarified by Zibrowius (1973). Edmondson & Ingram (1939) 

reported H. elegans (as Hydroides norvegica) from Kāne‘ohe Bay (1935), and also noted its presence 

in vessel fouling. Straughan (1969a, as H. norvegica) reported collections from Pearl Harbor 

from 1929 to 1948, and from Coconut Island (1968). Bastida-Zavala & ten Hove (2003) examined 

material collected in 1936 in Kāne‘ohe Bay. It remains common in Pearl Harbor and Kāne‘ohe Bay 

(Long, 1974; Evans et al., 1974; McCain, 1975; Grovhoug, 1979; Grovhoug & Rastetter, 1980; 

Rastetter & Cooke, 1979; Coles et al., 1999a, 2002a). Bailey-Brock (1976) also reports it from the 

Ala Wai Canal and notes that in general it is a “dominant fouling organism” in Hawaiian harbors. 

Coles et al. (1999b) record it from Barbers Point Harbor. 

Biological and ecological work on this species in Hawai‘i has been conducted for many years 

in the laboratory of M.G. Hadfield and his students. In Pearl Harbor, submerged fouling panels “typically 

accumulated 100% cover of H. elegans within a few weeks, throughout the year” (Walters et 

al., 1997). Walters et al. (1997) further note that these 75mm long worms in this habitat may reach 

reproductive maturity in 4 to 6 weeks. Hurlbut (1991c) reported on settlement and juvenile survival. 

Almost all “Tall Ships” arriving in Sydney Harbour in Australia after the “Bicentennial Race” 

had this worm on their hulls. It may have originated in the Indo-Pacific if not specifically in Australia 

(H. ten Hove, pers. comm., 1996). 

Pomatoleios kraussii (Baird, 1865) 

Introduced 

We consider this tropical Indo-Pacific species (Straughan, 1969b) to be introduced, and in particular 

possibly a post-World War II invasion. It also occurs in South Africa and Japan and has been 

introduced to the Levant region in Israel and Lebanon (H. Zibrowius, pers. comm., 1996). There 

appear to be no records or museum material prior to the 1960s. Straughan (1969a: 235–236) collected 

it only in Kāne‘ohe Bay at Coconut Island (1967–1968); Straughan (1969b) noted again that it 

was found only within Kāne‘ohe Bay and speculated that it might be in Pearl Harbor. Grovhoug & 

Rastetter (1980) reported it from Pearl Harbor and Kāne‘ohe Bay, based on 1976–1977 collections. 

Bailey-Brock (1976) noted that it was a “dominant” serpulid that “forms a distinct zone on the shoreward 

side of some leeward reefs and sea walls of Coconut Island” (citing Straughan), that it “occurs 

in the rocky intertidal and shallow subtidal on all the islands” (specific records are cited for O‘ahu, 

Maui, and Hawai‘i), and that it is a “dominant intertidal species not found below low tide.” These 

records indicate an expansion beyond the harbors and shallow bays. Coles et al. (1999a) note its continued 

presence in Pearl Harbor. Coles et al. (1999b) report it from Honolulu Harbor, Ke‘ehi Lagoon, 

and Barbers Point Harbor. Coles et al. (2006) report it from Kaua‘i, based upon 2002 collections; it 

also occurs in Kahului Harbor from 2003 collections (Coles et al,. 2004). Alan Miller (in Straughan, 

1969b) reported that the native gastropod Morula granulata will prey on it. 

Salmacina tribranchiata (Moore, 1923) 

Introduced 

= Salmacina dysteri of Hawaiian authors [in part], not of Huxley, 1855 [a European species]. 

Bastida-Zavala (2008) identified specimens of a Salmacina fouling an inter-island only cargo barge 

as this well-known Eastern Pacific (Alaska to Mexico) species. The material was collected in 2003 

(Godwin et al., 2004). Names of European species were first applied to many Pacific Ocean species, 

and this is such a case. 

Edmondson & Ingram (1939) found it in fouling beginning in 1935 in Kāne‘ohe Bay. Long 

(1974) reported it on “offshore” and Pearl Harbor fouling panels (1968), and Hartman (1966) noted 

that it occurred as “massed tubes fouling harbor installations during summer and fall months at Pearl 

Harbor and Kaneohe Bay.” Henderson (1990) reported it on the USS Arizona in Pearl Harbor (1986).


63 

Straughan (1969a) noted Bishop Museum records from Kāne‘ohe Bay (Coconut Island) from 1936 

to 1966 and from Black Point in 1937. Bailey-Brock (1976) noted that it was also present in the Ala 

Wai Canal, with further stations on O‘ahu, Maui, and Hawai‘i. Grovhoug & Rastetter (1980, based 

upon 1976–1977 collections) and Coles et al. (1999a, based upon 1996 collections) note its continued 

presence in Pearl Harbor. Bailey-Brock & Hartman (1987) reported that it is “common on fouling 

panels and hard substrates in sheltered waters, e.g., Pearl Harbor and Kaneohe Bay, Oahu.” 

Bailey-Brock (1989) recorded it on PVC settlement plates on an artificial reef in open waters at 20 

m, 2.4 km off the southeast coast of O‘ahu. Coles et al. (1999b) record it from Honolulu Harbor, 

Ke‘ehi Lagoon, Ala Wai Harbor, and Barbers Point Harbor. Coles et al. (2004, 2006) report it from 

Kaua‘i, based upon 2002 collections, and Coles et al. (2004, 2006) note 2003 collections from Maui 

and Hilo Harbor. We consider harbor and fouling populations of this Salmacina to be S. tribranchiata. 

Bailey-Brock (1976) reported its distribution in Hawaiian waters as “from shallow depths to 

200–600 meters,” and as being found across a wide variety of habitat in the Islands (rocky intertidal, 

reef flats on algae, anchialine lava ponds, and reef slopes to depth). This range of habitats suggests 

that additional Salmacina species may occur here. 

Bailey-Brock & Hartman (1987, plate 3.II.4.c) present a color photo of this species on a pier 

piling in Pearl Harbor. Hoover (1998, 2006) also gives a color photograph of specimens at 5 m at 

Kahe Point, O‘ahu. 

Serpula watsoni Wiley, 1905 

Introduced 

= Serpula vermicularis of Hawaiian authors, not of Linnaeus, 1767 

Bastida-Zavala (2008) reports this Indo-Pacific serpulid based upon specimens collected in 2003 

(Godwin et al., 2004) from an inter-island cargo barge operating solely in the Islands. We interpret 

this as the resolution of the identification of a common fouling Serpula recorded since 1936. 

Straughan (1969a: 231) recorded a serpulid (as the North Atlantic Ocean S. vermicularis 

Linnaeus, 1767) from Kāne‘ohe Bay as early as 1936, and from Pearl Harbor from 1938 on, noting 

that of these and other specimens, “none ... were collected from natural substrates.” It is curiously 

not listed by Hartman (1966) in her review of previous and new Hawaiian collections of polychaetes, 

although a fair amount of harbor- and bay-material was available to her, suggesting that it has perhaps 

never been common. Bailey-Brock (1976, as S. vermicularis) noted that in the islands it is “subtidal 

(and) relatively uncommon” and was found at stations on O‘ahu and the island of Hawai‘i. 

Coles et al. (1999a) note (as Serpula sp.) its continued presence in Pearl Harbor; Coles et al. (1999b) 

record it from Honolulu Harbor, and Coles et al. (2004, 2006) report 2002 collections in Nawiliwili 

Harbor and 2003 collections in Kaua‘i and Hilo Harbors. Serpula watsoni occurs from the Indian 

Ocean to southern Japan, including Australia and Micronesia (Bastida-Zavala, 2008), all of which 

range may not be natural. 

Spirorbid Tubeworms (Spirorbinae) 

The tiny coiled tubeworms historically placed in the genus Spirorbis, but now assigned to many different 

genera, interface with a large number of human-mediated dispersal vectors: they may occur 

in ship fouling, on commercial oyster shells, as juveniles or adults on small pieces of seagrass or 

floating debris taken into ballast tanks, on semisubmersible exploratory oil platforms, and so forth. 

In turn, many species of spirorbids are reported as occurring in harbors worldwide. Vine et al. (1972, 

p. 177), for example, note that the spirorbids Simplicaria pseudomilitaris and Neodexiospira pseudocorrugata 

settled abundantly on fouling panels in Kāne‘ohe Bay and that within about 30 days “had 

many fully mature adults of these two species, with well-developed embryos in their opercula.” Such 

reproductive strategies are particularly conducive to weedy dispersal. A number of authors have 

noted that ship dispersal may have been integral to the modern-day distribution of spirorbids 

(Knight-Jones et al., 1975; Bailey-Brock, 1976, p. 72). Bailey (1969) noted that no less than four 

species of spirorbids, Simplicaria pseudomilitaris, Pileolaria heteropoma, Janua pagenstecheri, and 

Neodexiospira pseudocorrugata (as Spirorbis corrugatus) occurred on the hull of a local fishing vessel 

on the island of Chios in the Aegean Sea.

64 


“Early” records of spirorbids in Hawai‘i include the report of Edmondson & Ingram (1939) of 

unidentified spirorbids from fouling in Kāne‘ohe Bay (1935 and later years). However, if spirorbids 

have been introduced here by ship fouling, such species likely arrived decades or centuries ago. 

Examination of historical materials (such as oyster shells from Hawaiian estuaries) for spirorbids 

with dried tissue in the shells, may prove fruitful in establishing earlier records. Thus, for example, 

dried specimens of Pearl Harbor spirorbids (some with their spiny opercula visible) are found on 

oyster shells (Ostrea sandvicensis) collected as early as 1919 in the mollusk collections of the 

Bishop Museum. Rehydration or genetic examination of this material should be possible. 

Bailey-Brock (1976) noted that “floating mangrove fruits, coconuts, driftwood, and other debris 

are often encrusted with spirorbines in Hawaiian waters, such debris providing an efficient dispersal 

mechanism for these gregarious worms.” Vine et al. (1972, pp. 178–179) also noted “the ease with 

which many adult Spirorbinae species may be transported on floating algae and the stones or shell 

fragments associated with these, and on crustacean carapaces, turtle shells, driftwood, ships’ bottoms, 

and perhaps on algae or barnacles attached to cetaceans.” Noting the few examples of 

endemism in the spirorbid fauna of the islands, they suggest that “Evidently there have been continuous 

invasions of species from other regions” (we interpret this to mean largely natural invasions, 

that is, range expansions) and conclude that “the widespread distribution of species can perhaps be 

explained by the ease with which adults may be transported.” We note that there appear to be no 

records of shallow-water spirorbids collected on drift material on the high seas. 

Knight-Jones et al. (1975) have gone into some detail on the evolutionary origin of different 

spirorbid clades and their means of dispersal. They suggest, based on diversity and climatic adaptations, 

that Janua pagenstecheri for example is native to the northeast Atlantic Ocean, whereas 

Neodexiospira brasiliensis (= Janua brasiliensis) [not yet recorded from Hawai‘i], Neodexiospira 

foraminosa (= Janua steueri), and Neodexiospira pseudocorrugata are native to the Indo-West 

Pacific or West Pacific. Regarding the now-widespread distribution of J. pagenstecheri and N. 

pseudocorrugata, they wrote: 

“As for methods of dispersal, most Janua may attach themselves to algae, including some of those 

algae which float when detached. In Europe pseudocorrugata is often found on Cystoseira, whilst 

pagenstecheri sometimes occurs sparsely on Fucus vesiculosus, but we have seen that these two most 

widely distributed of Spirorbinae are never found on algae in tropical waters. Probably their wide distribution 

has been brought about by long distance transport of shells lodged in the holdfasts of large 

floating algae such as Ecklonia and Macrocystis. Near Sydney (Australia) pseudocorrugata is often 

found on the algal part of such holdfasts. Self-fertilisation is possible, in pagenstecheri at least... so 

even a single adult thus transported to a littoral pool may establish this species in a new locality.” 

While there is no question that within northern or southern ocean basins dispersal of spirorbids on 

seaweeds must occur, little is known of the potential for such dispersal between oceans or across the 

equator; in particular, discovery of living spirorbids of any of the species treated below on drift material 

in mid-ocean (although such occur commonly near islands) would be of great interest. 

Regardless, as most of these species also occur in ship fouling, no presumption can be made as to 

the probability that drifting algae play or played a greater role. As a result, many spirorbids must now 

be considered cryptogenic. Spirorbid pelagic larval life is a few hours long (Bailey-Brock & 

Hartman, 1987), and thus larval transport to the islands is not possible. We conservatively treat only 

three species as introduced, in one case because of primary association with harbors and fouling 

communities, and in two cases because of a possible or presumed center of origin in the Atlantic 

Ocean. Other species in the Hawaiian Islands that occur on open reefs or even in deeper waters may 

of course also be cryptogenic or introduced, but we present a conservative list here. We treat one 

species, Vinearia koehleri, as a cryptogenic example based upon a potential dispersal vector, as discussed 

below.


65 

Eulaeospira orientalis (Pillai, 1960) 

Cryptogenic 

Bailey-Brock (1976) first reported this species from the Hawaiian Islands, based on collections made 

in the 1960s–1970s. The species was found on the green alga Dictyosphaeria cavernosa in Kāne‘ohe 

Bay (O‘ahu), as was Chaetopterus sp., and on rocks in Honokohau Harbor (Hawai‘i) (Bailey-Brock 

& Hartman, 1987). It appears to otherwise be known from the Red Sea, Madagascar, and Indian 

Ocean (Sri Lanka), as well as the Pacific Island groups of Fiji and Tonga (Bailey-Brock, 1985; 

Bailey-Brock, 1987). We regard it as cryptogenic. 

Janua pagenstecheri (Quatrefages, 1865) 

Introduced 

Vine et al. (1972) recorded material from intertidal stones on Sand Island (Kāne‘ohe Bay), based on 

collections made in the 1960s or 1970 and noted it has been recorded from many locations in the 

Mediterranean, Atlantic, and Pacific. Indeed, it is one of the most widely distributed of all spirorbids: 

“It is found in all oceans and extends from the equator to (in Norway) latitude 70°” (Knight- 

Jones et al., 1975). Knight-Jones et al. (1975) regard it as of European origin because of the breadth 

of its physiological and ecological adaptations to the climatic conditions of the northeast Atlantic 

Ocean. They also suggest (as noted above) that it may have gained entry to the Pacific Ocean by 

means of floating algae or other floating materials. However, the lack of physiological breadth noted 

by Knight-Jones et al. (1975) for Pacific populations would argue for relatively recent transport, and 

there is no reason to believe that this would be the case for natural drift algae. Considering the proposed 

European origin, we thus suggest that it has most likely been introduced by ships to the Pacific 

Ocean: Knight-Jones et al. (1975, pp. 123–124) noted that it occurs in ship fouling but does not survive 

on algae in tropical waters. Thus its long dispersal at sea on floating algae through the tropics— 

as opposed to quicker ship passages—would appear not to be likely (see earlier discussion, above). 

Leodora knightjonesi (de Silva, 1965) 

Cryptogenic 

= Janua knightjonesi 

One of a number of spirorbids reported by Vine et al. (1972) originally from Sand Island (Kāne‘ohe 

Bay) on intertidal stones but now more widely recognized around the islands (Bailey-Brock & 

Hartman, 1987) thus note that it is “very common on basalt rocks of the intertidal region of the Kona, 

Hawaii coast.” First Hawaiian records were in the 1960s or 1970s. Bailey-Brock (1976) reported that 

“an abundance [of this species] forms white encrustations on the lava rocks at Waiulua Bay, 

Anaehoomalu, Hawaii.” First described from the Indian Ocean (Sri Lanka), its origin is unclear, as 

there are isolated records in the Atlantic (West Indies) and the Indian and Pacific Oceans (Sri Lanka, 

Australia, Tonga, Hawaiian Islands) (Bailey, 1970; Vine et al., 1972; Knight-Jones et al., 1975; 

Bailey-Brock, 1987). We suggest it may be native to the Indo-Pacific and introduced to the West 

Indies and Hawaiian Islands. We treat it conservatively, however, as cryptogenic; it may have been 

introduced with ship rock ballast or in ship fouling. 

Neodexiospira foraminosa (Moore & Bush, 1904) Cryptogenic 

= Janua steueri Sterzinger, 1909 

Vine et al. (1972) reported specimens from 8 m depth on algae at Ma‘ili Point (O‘ahu) based upon 

material collected in the 1960s or 1970s. It was also noted by Grovhoug & Rastetter (1980) for 

Kāne‘ohe Bay and Pearl Harbor (1976–1977 collections. Bailey-Brock & Hartman (1987) noted that 

it “occurs in bays and harbors, often on algae attached to floating docks.” Bailey-Brock (1989) 

recorded it on PVC settlement plates on an artificial reef in open waters at 20 m, 2.4 km off the 

southeast coast of O‘ahu; N. foraminosa was one of the dominant colonists. Coles et al. (2006) report 

it from Kaua‘i and Maui, based upon 2003 collections. 

Neodexiospira foraminosa, first described from Japan (Honshu on red algae at 62 m; Knight- 

Jones et al., 1975) “has a Pacific distribution and has been collected at Johnston Atoll, Fiji, and 

Tonga” (Bailey-Brock & Hartman (1987). It is also reliably known from the Red Sea, Sri Lanka, and 

Florida (Knight-Jones et al., 1975) and New South Wales, Australia (Hutchings & Rainer, 1979). 

Knight-Jones et al. (1975) do not repeat or map the early record of it (as J. steueri) from the

66 


Caribbean (Bailey, 1970; Vine et al., 1972). Knight-Jones et al. (1975) consider it a species originating 

in the Western Pacific. Knight-Jones & Knight-Jones (1984) note the possibility that this species 

could be distributed in ship fouling (where it is often found), and that it has been found in fouling 

assemblages on test plates. We consider it cryptogenic in the Hawaiian biota. 

Neodexiospira nipponica (Okuda, 1934) 

Cryptogenic 

= Janua nipponica 

We consider this species cryptogenic based upon its disjunct distribution between Japan and Hawai‘i 

(Vine et al., 1972; Knight-Jones et al., 1975 (who note that South African material formerly assigned 

to this species is Janua brasiliensis) (Bailey-Brock & Hartman, 1987). It was collected, in the 1960s 

or 1970s from intertidal red algae at Nanakuli (O‘ahu). Coles et al. (2006) report it from Kaua‘i and 

Maui, based upon 2003 collections. 

It may have been introduced with ship fouling. Knight-Jones et al. (1975) suggest that N. nipponica 

may prove to be Neodexiospira brasiliensis (Grube, 1872). 

Neodexiospira pseudocorrugata (Bush, 1905) 

Cryptogenic 

= Janua pseudocorrugata; = Spirorbis corrugata of authors 

Vine et al. (1972) reported this spirorbid from intertidal stones at Sand Island and on fouling panels 

on Coconut Island (Kāne‘ohe Bay). Vine et al. (1972, p. 177) further noted that it was particularly 

abundant on the fouling plates at Coconut Island and that, along with Spirorbis marioni, is one of 

the few species able to tolerate upper shore conditions. Straughan (1977) documented its occurrence 

in upper intertidal rock pools on O‘ahu, where it is abundant on live and hermit crab-occupied snail 

shells. It was also noted by Grovhoug & Rastetter (1980) for Kāne‘ohe Bay and Pearl Harbor 

(1976–1977 collections). 

Although with a type locality in northwest France, and with a broad range from the British Isles 

to the Mediterranean, Knight-Jones et al. (1975) argued for an Indo-West Pacific origin and noted 

that it also occurs in ship fouling; we suggest here it may have been brought to Europe from the 

Pacific by the earliest sailing vessels. In the Pacific-Indian Ocean theaters it is also known from 

Australia, Mozambique, Sri Lanka, Tonga, Japan, and Catalina Island (California) (Knight-Jones et 

al., 1975; Bailey-Brock, 1987). The date of collection at O‘ahu is apparently in the 1960s or in 

1970s. Bailey-Brock & Hartman (1987) noted that it “is cosmopolitan in temperate and warm seas 

and is very common in shallow waters of Hawaii and at Johnston Atoll.” We regard it as cryptogenic. 

Pileolaria militaris (Claparède, 1868) 

Introduced 

Vine et al. (1972) noted this species on an oyster shell (= Isognomon sp.?, fide Vine et al., 1972, p. 

176) intertidally at Coconut Island, Kāne‘ohe Bay. Bailey-Brock & Hartman (1987) note that it is 

“found commonly in harbors and protected bays in Hawaii, and less frequently on exposed shores.” 

Bailey-Brock (1976, p.72) noted that it was “quite possible” that P. militaris may “have been spread 

to and between the islands” by ships. Vine et al. (1972, p. 176) also describe it as a species that “settles 

on a calcareous red alga which encrusts the undersides of platelike coral structures.” The date of 

first collection in the Hawaiian Islands is apparently in the 1960s or 1970s. It was also noted by 

Grovhoug & Rastetter (1980) for Kāne‘ohe Bay and Pearl Harbor (1976–1977 collections). 

It is widespread in the North Atlantic Ocean and the Mediterranean (Vine et al., 1972; Bailey, 

1969). If this is its area of origin, it is introduced to regions such as Mexico, and to the Pacific Ocean, 

where it is known from Hawai‘i, Australia, Tonga, and other regions (Vine et al., 1972; Bailey- 

Brock, 1987; Bailey-Brock & Hartman, 1987). We regard it as introduced. 

Pileolaria pseudoclavus Vine, 1972 

Cryptogenic 

= Pileolaria semimilitaris Vine, 1972 

This species was described from fringing reefs in the Red Sea and in the same year from fouling 

plates in “shallow” water at Coconut Island, Kāne‘ohe Bay (Vine, 1972; Bailey-Brock & Hartman, 

1987). The date of collection is apparently in the 1960s or in 1970 (Vine, 1972, p. 140). We regard 

it as cryptogenic; it may have been introduced with ship fouling.


67 

Simplicaria pseudomilitaris (Thiriot-Quiévreux, 1965) Cryptogenic 

= Pileolaria pseudomilitaris 

Vine et al. (1972) reported this species, originally described from the Mediterranean Sea, from 

stones and fouling panels on Sand Island (Kāne‘ohe Bay) and Koko Head from the intertidal to 3 m. 

The date of collection is apparently in the 1960s or in 1970. Grovhoug & Rastetter (1980) reported 

it from Pearl Harbor and Kāne‘ohe Bay (1976–1977). Bailey-Brock & Hartman (1987) note that it 

is “very common in harbors and shallow reefs of the Hawaiian Islands and at Johnston Atoll.” Vine 

et al. (1972, p. 176) note that it is “an abundant intertidal species usually found on the undersides of 

stones on the lower shore and in the shallow sublittoral zone.” Knight-Jones & Knight-Jones (1984) 

note that this species may owe its distribution “to transport on ships’ hulls” from which it has been 

recorded. Vine et al. (1972, p. 177) further note that it was particularly abundant on fouling plates at 

Coconut Island. It is now known from widespread stations in the Mediterranean and Atlantic (including 

the Aegean Sea and West Indies) and Pacific (Galapagos Islands, Australia, and New Zealand). 

We consider it cryptogenic. 

Spirorbis marioni (Caullery & Mesnil, 1897) 

Cryptogenic 

This species has been reported on Nanakuli, O‘ahu (at Mokoli‘i Islet) and Hawai‘i, on intertidal 

igneous rocks (Vine et al., 1972; Bailey-Brock, 1976; Bailey-Brock & Hartman, 1987), based upon 

specimens collected in the 1960s or 1970. It also occurs in Panama (the type locality), Mexico, the 

Galapagos Islands, Easter Island, and southern California (Vine et al., 1972; Bailey-Brock & 

Hartman, 1987). It is often “associated with harbours and thus ship transport” (Phyllis Knight-Jones, 

pers. comm., August 1996). Vine et al. (1972, p. 176) further note that it is a species of the “shallow 

rock pools in the upper shore and splash zone…(where)... it is presumably better adapted to avoid 

desiccation and salinity fluctuations than are most of the opercular-brooding species.” This would 

also suggest the potential to colonize ship hulls at the waterline. We consider it cryptogenic. 

Vinearia koehleri (Caullery & Mesnil, 1897) 

Cryptogenic 

= Pileolaria koehleri 

Vine et al. (1972) reported this species, described from the Mediterranean, from Mā‘ili Point (O‘ahu) 

and Hilo (Hawai‘i). On O‘ahu they were found at 8 m on stones; on Hawai‘i they were abundant 

intertidally and in shallow water on lava rocks. The date of first collection is apparently in the 1960s 

or in 1970s. Bailey-Brock (1989) recorded it on PVC settlement plates on an artificial reef in open 

waters at 20 m, 2.4 km off the southeast coast of O‘ahu. Vine et al. (1972, p. 176) note that it was 

“found intertidally on stones in rock pools but appears to favor settlement below tidemarks and can 

live in the intertidal zone only when it is permanently covered by water.” It has since been recorded 

from the West Indies, the Red Sea, New Zealand, and Australia, as well as at Johnston Atoll, Fiji, 

and Tonga (Vine et al., 1972; Bailey-Brock & Hartman, 1987).

68 


MOLLUSCA 

Class Gastropoda (snails) 

Introduced and Cryptogenic Gastropoda 



Diodora ruppelli 1962 SF, BW Indo-Pacific 

Tarebia granifera 1856 R Indo-Pacific 

Melanoides tuberculata 1994 R? Asia-Africa 

Pyrgophorus coronatus 1998 BW? Caribbean 

Vermetus alii


69 

well on the rocky reefs and sea walls.” Between 1927 and 1959, several attempts were made to plant 

California abalone, particularly the black abalone Haliotis cracherodii, on O‘ahu, as summarized 

below. Coles et al. (2002a) note that in 1968–1969 two additional species of California abalone, 

Haliotis fulgens and H. corrugata, were brought to Coconut Island in Kāne‘ohe Bay, but whether 

these were released in open water is not clear. Eldredge (1994) and Olin (1994) also note the occasional 

importation since 1989 of H. fulgens and the Asian abalone “Haliotis diversicolor supertexta” 

for closed-system research. No abalones are known to have survived in the wild in Hawai‘i. 

Releases of the California Abalone Haliotis spp. on O‘ahu 

Date Where Number Reference/Source 

1927 Kāne‘ohe Bay — Edmondson & Wilson, 

1940 (“where they 

seemed to do well at first 

but eventually 

disappeared”) 

1927–1928 Mokapu Point 5, 30 Brock, 1952 (two separate 

releases: Haliotis 

rufescens or H. 

cracherodii ?) 

1958 off Rabbit Island 167 Brock, 1960 (Haliotis 

cracherodii) 

1959 off Pyramid Rock 728 Brock, 1960 (Haliotis 

cracherodii, as two 

separate plantings, of 2.5- 

7.5 cm individuals from 

“southern or Lower 

California”) 

Fissurellidae 

Diodora ruppelli (Sowerby, 1834) 

Introduced 

Kay (1979, page 42) noted that this keyhole limpet was “... first reported on O‘ahu in 1962 from the 

reef near the entrance to the Pearl Harbor channel; specimens have since been recorded at depths to 

15 m ... described from the Red Sea and apparently occurs throughout the Indo-West Pacific.” It was 

recollected in Pearl Harbor in 1996 (Coles et al., 1997). Coles et al. (2002a) report it from Kāne‘ohe 

Bay based upon collections in 2000; Coles et al. (2002b) record it at Waikīkī, also based upon collections 

made in 2000. 

As it is unlikely that this species was previously overlooked in the molluscan fauna, and given 

that the first populations were discovered adjacent to a major harbor, we regard it as introduced, 

either in ship fouling or by ballast water. 

Trochidae 

Trochus niloticus Linnaeus, 1758 

Released; Not Established 

This famous large edible snail, native to Indo-Malayasia, Melanesia, and parts of Micronesia, has 

been widely released in the Pacific Ocean to establish new fisheries resources (Eldredge, 1987; Bour, 

1990). It has been released three times on O‘ahu, twice in 1952 and again in 1963, as shown in the 

following table.

70 


Releases of the Top Shell Trochus niloticus on O‘ahu 


1952 in Coconut Island ponds, 27 Brock, 1952, from Guam, 

Kāne‘ohe Bay 

as T. obelisus 

1952 in Coconut Island ponds, 2 Brock, 1952, from Fiji (12 

Kāne‘ohe Bay 

imported, only 2 alive at 

time of release); Bour 

(1990) reports the 1952 

release as a total of 39 

specimens. 

1963 Kāne‘ohe Bay 750 Bour, 1990; Eldredge, 

1994; from Guam; no 

evidence of subsequent 

reproduction, although 

specimens found over the 

years until 1978. 

Cross (1968) noted that divers had reported seeing T. niloticus in depths up to 27 m in Kāne‘ohe Bay. 

While Kay (1979, p. 20) and Kay & Schoenberg-Dole, (1991, p. 4) indicated that this snail has 

become established, there are no known reproducing populations. Specimens from the 1963 planting 

continued to appear until 1978, in keeping with age distributions for this snail (Bour, 1990, figure 

5, page 12). 

Wes M. Thorsson reports to us (June, 2003) that he saw a “few” specimens in about 20 m of 

water off Makapu‘u (south of Kāne‘ohe Bay), of uncertain dates. One “fresh-dead specimen”, 

encrusted with coralline and brown algae, and approximately 100 mm in diameter, was collected in 

January 1993 off Rabbit Island, Makapu‘u Point, at 8 m by C. Takahashi (C. Takahashi, pers. comm., 

June 2003). The latter specimen would be too young to be linked to the 1963 releases in Kāne‘ohe 

Bay, suggesting that later importations or releases, intentional or accidental, have occurred. 

Order Caenogastropoda 

Thiaridae 

Tarebia granifera (Lamarck, 1816) 

Introduced 

= Melania mauiensis Lea, 1856, described from Maui (Cowie, 1997) 

The natural range of this snail is unclear, but it probably is the Indo-Pacific region (Abbott, 1952). The 

first record in the Islands is based on the description of Melania mauiensis in 1856, although it may 

have been introduced prehistorically (Cowie, 1998), such as in the roots of wetland plants (for example, 

taro) brought with the Polynesians (Englund et al., 2000a). We include T. granifera here because 

of its occurrence in brackish and marine waters in Hawai‘i, such as at Kalauao in 3 ‰ in Pearl Harbor 

(Englund et al., 2000a), at Moanalua Stream in 15 ‰, and in Wai‘alaenui Stream in 30 ‰ (Englund et 

al., 2000b). 

Melanoides tuberculata (Müller, 1774) 

Introduced 

For the first record in the islands of this well-known snail in 1994, the native range is unclear but 

has been suggested to be either Asia, the Middle East, and Africa, but it has “probably been in the 

islands for many years” (Cowie, 1997). Indeed, it may have been introduced prehistorically (Cowie, 

1998) by the Polynesians. Cowie (2007) suggests that “earlier Bishop Museum material will probably 

result in much earlier dates for the presence of this species on these [O‘ahu and Maui] and the 

other main Hawaiian Islands.” Melanoides tuberculata occurs in brackish and marine waters on 

O‘ahu—in 15–23 ‰ in Moanalua Stream and to 31 ‰ in Kapalama Stream and to tidal areas of 

lower Mānoa-Palolo Stream, ranging from 14 to 33 ‰ (Englund et al., 2000b).


71 

Barleeiidae 

Amphithalamus inclusus Carpenter,1864 

Establishment Unknown 

We discuss this tiny Californian barley snail below along with species of pyramidellids. Whether it 

is established is not known. It was found in 1981 in Kawaihae Harbor, on the northwest coast of the 

island of Hawai‘i by B. Draper (in litt., 9 June 1986), and again in 1985 near the Navy Pier on the 

northeast side of Sand Island, Midway Islands, by D. Shasky. 

Hydrobiidae 

Pyrgophorus coronatus (Pfeiffer, 1840) 

Introduced 

Cowie (1999) has provisionally identified a brackish water snail from Pearl Harbor, first collected 

in 1998, as this species, which was previously known from the Caribbean islands and from continental 

areas bordering the Caribbean. Populations were found in the Waiawa Wildlife Refuge, 

Pouhala Marsh, and Waiawa Springs. Englund (2002) notes that “densities of this newly introduced 

species were high with up to several thousand incidentally captured in a single seine haul.” 

Pearl Harbor specimens exhibited morphological variation in the extent of shell spination and 

relative height and width (Cowie, 1999). Although it does not have planktonic larvae, Cowie suggests 

that introduction via ballast water cannot be excluded. Snails entrained on bits of floating 

debris could be taken up in ballast water. 

vermetidae 

Thylaeodus (?) sp. 

Introduced 

Strathmann & Strathmann (2006) report a small (up to 7.7 mm diameter) vermetid snail (“Hadfield’s 

vermetid”, named after Michael G. Hadfield, its discoverer in Hawai‘i) from the Hawaiian Archi - 

pelago and Guam. We hypothesize that it was introduced from the Atlantic Ocean. While best known 

from O‘ahu, specimens are reported from Kure, Midway, Pearl and Hermes, Lisianski, Maro, French 

Frigate Shoals, and Necker, as well as from Luminao Reef in Guam. It appears to be indistinguishable 

(in terms of shell morphology) from a species known from Florida, the Azores, and the Medi - 

terranean, which may be Thylaeodus rugulosus (Monterosato, 1878), described from the latter sea. 

Of particular interest is that specimens from O‘ahu and Florida differed by only 20 of 1,745 base 

pairs (in various sequences of rRNA and tRNA). 

Thylaeodus (?) sp. occurs on O‘ahu in harbor fouling communities, and on intertidal and shallow 

subtidal shells and coral rubble. It was first noted in 1970 on the walls of the outdoor aquaria at 

the Kewalo Marine Laboratory in Honolulu. Its occurrence on oyster valves suggests that examination 

of museum material would be of interest to establish earlier records, if any. In Pearl Harbor, it 

occurs with other vermetids, bryozoans, and sponges, in a classic fouling facies. 

The genetic identity of Hawaiian material with an Atlantic population suggests recent gene 

flow, which while unlikely to be maintained naturally by floating debris between the Atlantic and the 

Hawaiian Islands, would be expected of ship-mediated transport. Its widespread distribution across 

the Atlantic-Mediterranean offers the hypothesis that it may be native to that ocean. If introduced initially 

to O‘ahu by shipping, downstream dispersal [such as by rafting, to which it would be well 

adapted (Strathmann & Strathmann, 2006)] across the Hawaiian Island chain would be expected, 

while ship traffic would take a fouling species to Guam. 

Vermetus alii Hadfield & Kay in Hadfield et al., 1972 Introduced 

Michael G. Hadfield (pers. comm. May 1991 and December 1995) informs us that what he and 

Alison Kay described as V. alii he had also seen in Miami, Florida, and he felt that it was a probable 

introduction to Hawai‘i. We concur, and regard it as a species introduced with ship fouling. 

Evans et al. (1972) reported it (as Vermetus sp. n.) from Pearl Harbor. Hadfield et al. (1972) 

described V. alii with a type locality at Mokuloe (Coconut Is.), Kāne‘ohe Bay, O‘ahu, on subtidal 

coral, noting that it was “the largest species of Hawaiian vermetid, occurring intertidally and subtidally 

cemented to coral heads and concrete sea walls such as those in the docks at Mokuoloe Island” 

(alii is an Hawaiian word for chief or royalty, in reference to the snail’s large size and purple and

72 


yellow color). Kay (1979) noted that it occurs “intertidally and subtidally cemented to coral heads 

and concrete sea walls in bays such as Kaneohe Bay and Pearl Harbor, Oahu.” Grovhoug & Rastetter 

(1980) also reports it from Pearl Harbor and Kāne‘ohe Bay (1976–1977) as does Hen derson (1990; 

Pearl Harbor, 1986). It still occurs in Pearl Harbor (Coles et al., 1999a). Additonal records include 

Honolulu Harbor, and other south and west shore harbors and marinas on O‘ahu; Kāne‘ohe Bay, 

Waikīkī and Koko Marina, and in harbors and on some nearby coral reefs on Kaua‘i, Moloka‘i, 

Maui, and Hawai‘i (Coles et al.,1999b, 2002a, b, 2004, 2006). Carlton (1999) lists it as tentatively 

originating from the Caribbean (the subtropical Western Atlantic Ocean), although alternatively it 

may originate from the subtropical mainland Eastern Pacific. Kay & Switzer (1974) reported it from 

Fanning Island (as Serpulorbis alii) based on 1972 collections. 

Rudiger Bieler informs us (pers. comm., 9 August 1996) that he believes that Vermetus alii is a 

junior synonym of Eualetes tulipa (Chenu, 1843), a species of the western Atlantic and Eastern 

Pacific. Coles & Eldredge (2002) and the various Coles et al. reports used the latter name (Eulates 

tulipa) to refer to this species. 

Hipponicidae 

Hipponix australis Lamarck, 1819 

Cryptogenic 

= Sabia conica of authors 

= Hipponix minor Garrett, 1857 vide Kay, 1979 (as “1853”) 

Kay (1979, as Sabia conica) notes that this Indo-West Pacific hoofsnail, which lives attached to other 

snails such as Turbo, Terebra, and Conus occurs in “all islands in the Hawaiian chain”; she illustrates 

a specimen on Nassarius papillosus. As it is capable of being transported either in ship fouling 

or by ballast water [and has thus been introduced to British Columbia (Carlton, 1992b)], and as 

there appear to be no fossil records of it on the Islands—although snails of similar structure, morphology 

and size are represented in the O‘ahu fossil record, such as Cheilea and Crepidula (Kosuge, 

1969)—we consider it cryptogenic. It was first collected in the 1850s in the Islands. Hoover (1998, 

2006) provides a color illustration of specimens attached to a Terebra shell at Lāna‘i Lookout, 

O‘ahu, in shallow water. 

Calyptraeidae 

Bostrycapulus calyptraeformis (Deshayes, 1830) Introduced 

= Crepidula aculeata of Hawaiian authors, not of Gmelin, 1791 (see Collin, 2005). 

= Crepidula aculeata hawaiiensis Dall, 1922 [a “manuscript” name on specimen labels in the Bishop 

Museum with the note, “identified by author” (William Healey Dall)]. 

Small spinous crepidulas around the world have often been identified as the “cosmopolitan” species 

Crepidula aculeata, departing strikingly from the general observation that shallow-water marine 

gastropod mollusks are not globally distributed. Collin (2005) has identified the Hawaiian stocks of 

this slipper limpet as Bostrycapulus calyptraeformis. 

Bishop Museum collections contain one lot of Bostrycapulus calyptraeformis collected by W.A. 

Bryan at Paumalu, O‘ahu in 1913, and a number of lots from Pearl Harbor and Honolulu Harbor from 

1915 and later. Bryan (1915) listed and illustrated “C. aculeata” from Hawai‘i without a specific location. 

Edmondson (1933) referred to it as “about the shores.” Edmondson & Ingram (1939) reported it 

from fouling communities in 1935 and later years from Kāne‘ohe Bay. Evans et al. (1972) reported it 

from Pearl Harbor (1971), as did Long (1974, who also found it settling on nearshore panels, collections 

of 1968–1972) and Grovhoug & Rastetter (1980, Pearl Harbor and Kāne‘ohe Bay, collections of 

1976–1977). Kay (1979) noted it commonly occurs “... on boat bottoms, pilings, and rocks, especially 

near harbors.” It occurs in harbors, marinas, and some coral reef sites throughout the Islands. 

B. Draper (in litt., 9 June 1986) regarded it as an introduction to the Hawaiian Islands; his 1984 

specimens came from Pohoiki Bay near an old Sugar Cane barge dock. Draper also reported that D. 

Shasky’s collection included specimens from Hilo and Kawaihae, Hawai‘i as well as Midway Island 

near the Navy piers. Burgess (1995) noted specimens collected in 1959 in the Ala Wai Boat Harbor. 

Bostrycapulus calyptraeformis is native to the warmer waters of the Eastern Pacific, from 

Panama to Peru [Collin (2005), who noted that Reeve (1859) listed “Crepidula aculeata“ from the


73 

“Sandwich Islands” (Reeve’s record may, in turn, have been the basis of Sowerby’s (1883) listing it 

from the “Sandwich Islands” as well, rather than the listing by Sowerby representing an independent 

record)]. No further 19th century reports of C. aculeata from Hawai‘i were forthcoming; in the 

nineteenth century many shells were erroneously reported from the “Sandwich Islands”. 

Collin (2005) also noted that there are specimens of an unidentified “Pleistocene” calyptraeid 

from Hawai‘i “possibly attributable to Bostrycapulus” in the Academy of Natural Sciences of Phila - 

delphia collections (ANSP 116536). However, the material is in very poor condition, consisting of 

several broken shells (Collin, 2005, and pers. comm.) and the specimens are not verified as fossil (R. 

Collin, pers. comm.). Spiny crepidulas are otherwise unreported as fossils from the Hawaiian 

Islands. 

Collin (2005) found that Hawaiian populations were a close genetic match to populations in 

Panama. We regard B. calyptraeformis has a ship-fouling introduction in the late 19th or early 20th 

century from Panama to Hawai‘i. 

Crucibulum spinosum (Sowerby, 1824) 

Introduced 

This “spiny cup-and-saucer shell” appeared—as did a number of other species—in the Hawaiian 

fauna during or soon after World War II. Keen (1971) noted an Eastern Pacific distribution from 

“California and southward through the Gulf [of California] to Tomé, Chile, on stones and dead 

shells, intertidally, and offshore in depths to 55 m.” Kay (1979) referred to it as “circumglobal in distribution,” 

perhaps with Crepidula aculeata in mind; C. spinosum is classically restricted to the 

warm Eastern Pacific Ocean and is not listed in molluscan works for Australasia, the Indo-Pacific, 

the Western Pacific, or the Atlantic Ocean. However, Springsteen & Leobrera (1986) report it as 

introduced to Palawan in the Philippines as a new record (although the first date of collection is not 

given); they note that it has been introduced to Hawai‘i and that the Philippine record represents a 

“considerable westward expansion of the known range.” Given the volume of commercial ship traffic 

between Pearl and Honolulu Harbors and the Philippines, as well as the number of ocean-going 

yachts that visit both Hawai‘i and Palawan, the appearance of C. spinosum in the Philippines is not 

a surprise—indeed, it probably now already occurs in other Pacific harbors but goes unreported. 

Edmondson (1946d) was the first to remark on its presence in Hawai‘i, noting that “One interesting 

gastropod has recently appeared in Honolulu Harbor. It apparently is a species of Crucibulum, 

closely related to Cheilea dillwyni, except that the internal cup of the shell is entire instead of crescent 

shaped.” Edmondson (1948) identified the Honolulu Harbor population as Crucibulum spinosum; 

in 1949, he added Pearl Harbor (Edmondson, 1949), noting that “This species described from 

the west American coast, has apparently become widely established in Hawaii within recent years.” 

Edmondson (1952a) referred to some “fine specimens” obtained at Pier 27, Honolulu Harbor. 

Tinker (1958, as Calyptraea spinosum) and Burgess (1959) further referred to its introduction, 

Burgess noting that it was by then widespread around O‘ahu. Ulbrick (1969) reported that it occurred 

on pieces of dead coral or basalt rocks dredged from sand and rock bottom at 5 to 8 m in Kāne‘ohe 

Bay and that it was also found on sea walls and experimental trays put out for oyster spat. Long 

(1974) reported it from fouling panels in Pearl Harbor (1968–1972); Grovhoug & Rastetter (1980) 

also reported it from Pearl Harbor and Kāne‘ohe Bay (1976–1977). Coles et al. (1999a) report it 

from Pearl Harbor in 1996, Honolulu Harbor, Ke‘ehi Lagoon and Barbers Point Deep Draft Harbor 

in 1997 (Coles et al., 1999b) Kāne‘ohe Bay (Coles et al. 2002a) in 2000, and Moloka‘i, Kawaihae 

and Hilo Harbors based upon 2003 collections (Coles et al. 2004, 2006). 

Kay (1979) noted that it “... may have been introduced in Hawaii during World War II.” Burgess 

(1995) remarked that it “became abundant on flat submerged objects in about 1950. Perfect specimens 

were collected [from a refloated boat] in 1959 from Ala Wai Boat Harbor. The probable source 

was some port in Southern California.” 

Cypraeidae 

Tinker (1958) listed three species of Indo-Pacific cowries, Cypraea cylindrica Born, 1778, Cypraea 

cribraria Linnaeus, 1758, and Cypraea staphylaea Linnaeus, 1758, each with the following comment 

but without further explanation: “It has been brought into the Hawaiian Islands on the hulls of

74 


ships.” Burgess (1959) noted that the source of these records were collections made off the USN concrete 

barge YON-146, which had arrived in Pearl Harbor from Guam, by S. Tinker, C.M. Burgess, 

and others in 1950 (although neither Tinker nor Burgess mention the date of sampling). Burgess 

added that four additional species of cowries had also been found on the barge: Cypraea clandestina 

Linnaeus, 1767, Cypraea carneola Linnaeus, 1758, Cypraea helvola Linnaeus, 1758, and Cy - 

praea poraria Linnaeus, 1758. Cypraea carneola, C. helvola, and C. poraria were all already known 

from and are considered native to Hawai‘i, the first two being known from fossils as well. Cypraea 

cylindrica and C. cribraria are not further known from the Hawaiian biota. Cypraea staphylaea is 

known from a few living specimens in Hawai‘i. The discovery of such a diverse array of cowries in 

hull fouling from a vessel that had traversed the ocean is notable and suggests that the original distribution 

of cowries throughout the Indo-Pacific may have been altered by shipping over the past 500 

years, although little to no reference is made to this vector in discussions of cowrie biogeography. 

Eyerdam (1959) had earlier reported on the discovery of Cypraea tigris on an army transport vessel 

travelling from the Marshall Islands to Puget Sound (see Carlton, 1987, Table 1, p. 454). 

The late C.M. Burgess, a well-known cowry expert, in a paper entitled “Strangers in Hawaii” 

(1995), argued that several cowry species were probable introductions to the Hawaiian Islands. That 

cowries can be transported in hull fouling is demonstrated above, and Burgess, a seasoned observer 

of the Hawaiian fauna, took note of unusual records. We tentatively admit one species, otherwise 

known from Japan, as cryptogenic. We note that the islands are naturally bathed with planktotrophic, 

teleplanic larvae from distant locations, and volunteer species in open deeper waters may be 

expected, and, indeed, these would be most likely recognized among the mollusks. Burgess listed 

Cypraea gaspardi Biraghi & Nicolay, 1993 [type locality: Kwajalein; collected in 1993 and 1994 

(alive?) off Pearl Harbor in 12 m of water; “this recent finding proves that introduction of exotic 

mollusks is still occurring, as can be expected,” Cypraea arabica (Linnaeus, 1767); Cypraea depressa 

Grey, 1824; Cypraea pallida Gray, 1824 [“probably”, collected in 1969 (1, alive, 50’, off 

Molokini islet near Maui; previous known distribution was Bay of Bengal and Sri Lanka)]; Cypraea 

labrolineata Gaskoin, 1849; Cypraea hirundo Linnaeus, 1758; Cypraea gracilis Gaskoin, 1849; and 

Cypraea staphylaea Linnaeus, 1758. We do not further treat any of these species here. 

Cypraea kuroharai Habe, 1961 

Cryptogenic 

Of the many cowry species discussed by Burgess (1995), one species, Cypraea kuroharai, is compelling 

as a possible ballast-water or ship fouling introduction. The only records appear to be Japan 

and Hawai‘i. Burgess notes that it was first found in 1971 in O‘ahu in 20 m by John Earle and that 

a total of about 14 collections have been made since then, “spread over most of Leeward O‘ahu.” 

Burgess (1995) felt that the resolution that Cypraea kuroharai as a distinct species “verifies the 

introduction of this species to Hawai‘i from Japan, its type locality.” Burgess further considered this 

the only introduced cowry that could be considered as established. 

Muricidae 

Vitularia miliaris (Gmelin, 1791) 

Native 

= Vitularia sandwicensis Pease, 1861, fide Kay (1979) 

Burgess (1963) offered a case for the introduction of this “murex” in the Hawaiian Islands, stating 

that the first live specimens of this Indo-Pacific species in Hawaiian waters were not collected until 

1962 (or 1959, a date Burgess later gives in the same paper). Burgess (1963) noted that live specimens 

were found on Barge YON-146 in 1950 in Pearl Harbor, thus leading him to suggest that 

“World War II vessels docking in Hawaiian harbors” had introduced it. Burgess’s records included 

collections made in April, 1959 [36 m, off Kewalo Basin, O‘ahu (fide caption)], May 1962 (at 27 m, 

vide text; at 36 m, vide caption, in sand and coral off Kewalo Basin) and February 1963 (Mā‘alaea 

Bay, Maui, dredged). Subsequently, however, Rehder (1964) noted that there were specimens in the 

Smithsonian Institution from Hawai‘i collected in 1902 (Albatross dredging, one specimen off 

O‘ahu, 26 - 51 m), 1915 (Thaanum and Langford collectors, at entrance to Honolulu Harbor), 1916 

(off Waikīkī), and 1918 (off Launiupoko Camp, Maui). Kay (1979) lists Vitularia sandwicensis


75 

Pease, 1861, described from Hawai‘i, as a synonym of V. miliaris. James (1989) notes Pleistocene 

material of Vitularia sandwicensis from the Mokapu Peninsula, O‘ahu. 

While Kay (1979) did not mention Burgess’s 1963 paper, the apparent absence of collections from 

between 1918 and 1959 is curious and it would be of interest to find similar cases of such hiatuses 

among other Hawaiian mollusks. Burgess noted, as did Kay, that the species is found in shallower 

waters elsewhere: “On Guam the shell is found in very shallow water or even exposed on the reef at 

low tide.” Burgess provided a photograph of a specimen collected by Thaanum “at intertidal zone on 

the reef” on Guam. Kay (1979) stated that in Hawai‘i it occurs “at depths of 30 to 60 m; elsewhere in 

the Indo-West Pacific V. miliaris occurs in shallow water or even exposed on reefs at low tide”, quoting 

Cernohorsky (1967). Kay (1967) noted that many prosobranchs “which are widely distributed in 

the Indo-West-Pacific live in deeper water in the Hawaiian Islands than they do elsewhere.” 

Montgomery & Montgomery (1963) reported that in seven years of collecting on Guam, they 

had only encountered (or heard of) Vitularia miliaris on pilings, usually on the oyster Dendostrea 

hyotis on pilings in Apra Harbor (V. miliaris, if it is not in the fossil record in Guam, is thus perhaps 

a candidate for consideration as an introduction to Guam). Burgess (1995), not considering Rehder’s 

(1964) note or Kay’s (1979) synonymy, repeated his view that Vitularia miliaris was introduced, 

linking its occurrence again in O‘ahu to Barge YON-146: “This mollusk has become firmly established 

in deeper (10–20 fathoms) leeward Oahu waters. The Pele dredged many in all stages of 

growth at nearly all stations in this area in 1960.” Burgess continued, “That this mollusk would 

thrive in rather deep Hawaiian waters is an ecological mystery when its ‘normal’ habitat is shallow 

or even intertidal water” (but note Kay, 1967, above). 

While the inevitable conclusion is that this species is native, it may be that it is a waif in the 

islands, going through periods of recruitment from the Indo-Pacific. Finding it in fouling on a barge 

arriving in Hawai‘i, combined with a perhaps coincidental mid-20th century natural colonization or 

a resurgence episode in the decade thereafter, would have lead Burgess to the possible conclusion 

that it was introduced. There remains the possibility that vessel fouling could also have transported 

to the islands novel genotypes that would have lead to its increased abundance by 1960. We consider 

it to be native. 

Conidae 

Conus capitaneus Linnaeus, 1758 


Burgess (1959) proposed that this Indo-Pacific cone snail was introduced to Hawai‘i in World War 

II, based upon collections of unspecified date “by the Harrisons”, who were local shell collectors. 

Kohn (1959) stated that, “This species was not known to occur in the Hawaiian area until the collection 

of a living specimen by Mrs. A.M. Harrison... ,” noting two specimens: one, collected alive, by 

Mrs. Harrison, at a depth of 5.4 m off Nanakuli, O‘ahu (April 1958), and a second, by Mr. A.M. 

Harrison, at a depth of 10.7 m near Ka‘ena Point, O‘ahu (August 1958). Kohn & Weaver (1962), 

however, re-identified the Ka‘ena Point specimen as Conus vexillum Gmelin, 1791. 

Kay (1979) noted it is rare, found at depths of 20 m. Alan J. Kohn (pers. comm., July 1996) felt 

that the records represented the “arrival of larvae spawned elsewhere, and that the species (like several 

others that have been collected once or twice in Hawai‘i), do not maintain breeding populations 

there.” We do not further consider it here. 

Subclass Heterobranchia 

Superorder Pyramidellidacea 

Pyramidellidae 

The late B. Draper (Draper, 1986) records seven species of small California snails in Kawaihae 

Harbor, on the northwest coast of the island of Hawai‘i, which “proved to be identical to species 

described from Southern California.” He suggested that these species were “accidentally imported 

on U.S.Marine Corps landing craft during practice landings at the Kawaihae Harbor”, some of which 

craft came from Long Beach, California. Draper (in litt, 9 June 1986) provides to us the names of 

these species, further detailed information, and photographs. One of these seven species, Amphi -

76 


thalamus inclusus, is a barleeiid, but it is listed here (as well as above) because of the nature of the 

dispersal phenomenon. We treat all of these species as of unknown establishment: 

Amphithalamus inclusus Carpenter, 1864 


Boonea cincta (Carpenter, 1864) 


[identified as Chrysallida vicola (Dall & Bartsch, 1909) by Draper; now regarded as a synonym of B. cincta 

(Turgeon et al., 1998)] 

Ividella navisa (Dall & Bartsch, 1907) 


Peristichia pedroana (Dall & Bartsch, 1909) 


Evalea sp., cf. E. americana (Dall & Bartsch, 1904) Establishment Unknown 

Chrysallida trachis (Dall & Bartsch, 1909) 


Iolaea eucosmia Dall & Bartsch, 1909 


The specimens were collected from screenings from bottom dredgings by a U.S.Government scoop 

dredge, during deepening of the harbor from 12 to 15 m, in October 1981. Kawaihae Harbor served 

ships operating for C&H Sugar Co. and Chevron Oil, and was used by U.S.Marine Corps amphibious 

landing craft for practice. Landing craft were reported to be from Long Beach, California, 

according to a resident of Kawaihae and were brought over two or three times a year. 

Draper’s photograph of one species, Ividella navisa, bears the inscription, “dark material in 

early whorls may be dead body material”, implying that Draper felt it was possible the specimens 

were collected alive. All the photographs uniformly show very fresh, well-sculptured shells, not suggestive 

of worn (dead or beach) material. Draper subsequently sampled a similar habitat (dredgings 

under a docking pier used by naval vessels and oil tankers) on Midway Island and did not find any 

of the above species. He felt that this supported his hypothesis that these southern California species 

were brought over to Hawai‘i by these landing craft. 

It remains to be determined if any of these species are established in Hawaiian waters. Draper 

felt that pyramidellids, in particular, could be introduced to one locality and remain in one site for 

many years because of their habitat specificity. Eldredge (1994, p. 10) noted that an unidentified 

species of pyramidellid was reported to have been on giant clams (Tridacna spp.) imported to 

Hawai‘i in the 1980s. 

Hinemoa indica (Melville, 1896) 

Cryptogenic 

= Odostomia pupu Pilsbry, 1918 fide Kay, 1979 

We regard this species, because of its disjunct distribution and its habitat in Hawai‘i, as cryptogenic. 

According to Kay (1979) this species is “common in tide pools and on fringing reefs, and abundant 

in shallow, nutrient rich waters such as Pearl Harbor and south Kaneohe Bay, Oahu....” It was originally 

described from the Indian Ocean and is also known from Sri Lanka and “Indo-China” (Kay, 

1979). Coles et al. (1999b) record it from Pearl Harbor in 1996 (Coles et al., 1999a), Honolulu 

Harbor in 1997 and Barbers Point Deep Draft Harbor in 1998 Coles et al. 1999b), Kāne‘ohe Bay in 

1999 and 2000 (Coles et al., 2002a) Waikīkī in 2001 and Hawai‘i Kai in 2002 (Coles et al., 2002b), 

and Maui (Coles et al. (2004, 2006), based upon 2003 collections. 

It has been present in the islands for many years, having been described as Odostomia pupu by 

Pilsbry (1918). Pilsbry’s material came from “Waikiki beach, near Honolulu.” Gary Rosenberg of 

the Academy of Natural Sciences of Philadelphia informs us (pers. comm., January 2000) that the 

type lot of Odostomia pupu lists Frederick Stearns as the collector. Pilsbry noted that some of his 

specimens were ”minutiae picked out of shell-sand sent many years ago by Mr. Frederick Stearns.” 

Stearns died in 1907, the same year the material was cataloged at ANSP. 

Pyrgulina oodes (Watson, 1886) 

Cryptogenic 

Reminiscent of Hinemoa indica, this snail, “common in tide pools and on reefs and in sediments to 

depths of 65 meters” is also “the dominant pyramidellid in micromolluscan assemblages from nutrient-rich 

waters such as Pearl Harbor and south Kaneohe Bay, Oahu” (Kay, 1979). Described from 

Flinders Passage, northwestern Australia, it has otherwise been reported from Enewetak (Kay, 1979).


77 

We regard it as cryptogenic. It remains to be determined when specimens were first collected in the 

Islands. 

Superorder Opisthobranchia 

Order Cephalaspidea 

Bullidae 

Bulla adamsi (Menke, 1850) 

Failed 

Tinker (1958) stated that this Eastern Pacific snail had been “recently introduced into the Hawaiian 

Islands.” Burgess (1959) believed it to have been introduced during World War II and noted that it 

was “now” widespread around O‘ahu. Burgess (1995), noting that specimens were first found in 

1946, stated that Bulla adamsi “were abundant in the muddy silt at night across the Ala Wai Channel 

from the Honolulu Yacht Club in 1949.... They continued to be collected until about 1956. The construction 

of Magic Island obliterated this area” (a figured specimen was collected in 1949 at night at 

Ala Moana Reef, opposite the Honolulu Yacht Club). It is not mentioned by Kay (1979). We consider 

it a failed invasion, possibly destroyed by shoreline destruction. 

Bulla vernicosa Gould, 1859 

Native 

= Bulla marmorea Pease, 1861 (type locality Hawai‘i) 

= Bulla peasiana Pilsbry, 1893 (type locality Hawai‘i) 

Burgess (1959) considered this (as Bullaria vermicosa [sic]) to be a World War II-era introduction. 

However, Kay (1979) noted that the species had long been known in Hawai‘i under other names (see 

above) and further notes its sporadic appearances. Bulla vernicosa is “widespread in the Pacific” 

(Kay, 1979). Burgess (1995) noted that this species was “abundant on an exposed (at low tide) sand 

islet in mid Kāne‘ohe Bay in 1954. This large Bulla probably came from Guam. Whether these mollusks 

are still present there is unknown to the author who has not collected the area in over forty 

years. Chris Takahashi collected this species in 100 feet buried in sand at Kōloa, Kaua‘i in 1981. I 

have no record of other findings. This large handsome shell would be difficult to overlook.” 

Burgess’s figured specimen is from Kāne‘ohe Bay collected in 1954. We regard this as a native 

species, with its aperiodic appearances having been mistaken for an invasion. Cephalaspideans are 

particularly well known for their sporadic blooms in shallow water. 

Order Nudibranchia 

Cuthona perca (Marcus, 1958) 

Introduced 

Gosliner (1980) reported this western Atlantic species [Brazil, its type locality, as well as Florida and 

the Caribbean (Jamaica, Barbados)] from the floating docks in association with the sea anemone 

Aiptasia sp. at Coconut Island, Kāne‘ohe Bay, based upon specimens collected in 1972 and 1973. 

The only Aiptasia reported from Hawai‘i is A. pulchella Carlgren, 1943 (Cutress,1977). This nudibranch 

is also known from New Zealand (Gosliner, 1980). Terrence Gosliner (pers. comm. July 

1996) feels that it is ”highly likely to be introduced. [The] only known collections that I have seen 

are from Kaneohe Bay.” 

Caloria indica (Bergh, 1896) 

Introduced 

= Learchis indica 

= Caloria militaris (Alder & Hancock, 1866) 

Gosliner (1980) reported this species, found preying on the hydroid Pennaria tiarella, as common 

in Kāne‘ohe Bay on patch reefs and on floating docks of Coconut Island; single specimens were also 

collected from Ala Moana Beach Park and from 12 m off Pokai Bay, based upon specimens collected 

in 1972–1973. Earlier reports include those of Baba (1969), based upon material collected in 1968 

by G. Robilliard from “Hawaii”; Baba noted that Robilliard reported that “his specimens were found 

feeding exclusively on ... Pennaria growing in shallow water.” Burn & Narayanan (1970) recorded 

specimens also collected in 1968 by Robilliard from Kāne‘ohe Bay, on Pennaria hydranths. Bertsch 

& Johnson (1981) illustrate a 22 mm specimen from Magic Island. Hoover (1998, 2006) provides a 

color photograph of a specimen at Kahe Point, O‘ahu, in 6 m.

78 


Described from India, it is regarded as being widespread throughout the Indo-Pacific, including 

India, Sri Lanka, Australia, New Zealand, Japan, and New Caledonia. Bertsch & Johnson (1981) 

noted it as “a common subtidal species that eats the hydroid Halocordyle [ = Pennaria].” We regard 

both it, and its prey, as introduced. 

Okenia pellucida Burn, 1967 

Introduced 

This nudibranch, first described from Australia, was collected in 1972 on the introduced bryozoan 

Zoobotryon on floating docks on Coconut Island in Kāne‘ohe Bay; it was also found in the Ala Wai 

Canal next to Magic Island, O‘ahu (Gosliner et al., 1986). Gosliner (2004) reports further specimens 

from Mā‘alaea Bay, Maui, in 1991. It is now known from Australia, New Zealand, Japan, Palmyra 

Atoll, and Malaysia (Rudman, 2004; Gosliner, 2004). Outlier popuations are those in Hawai‘i and in 

Dubai, United Arab Emirates (Rudman, 2004). Burn (1967) noted that it occurred in Sydney Harbour 

(specimens collected in 1956) on the bryozoan Zoobotryon (as Z. pellucidus = Z. verticillatum of 

current literature and as cited in this monograph), growing on wharf piles, floating pontoons, and 

small boats. Willan & Coleman (1984) report it from Queensland (Moreton Bay), Western Australia 

(Fremantle Harbour), and New Zealand (Waitemata Harbour), confined to harbors where Zoo - 

botryon occurs in shallow water. Rudman (2004) suggested “that its wide distribution has probably 

been aided by shipping.” Larval dispersal and floating Zoobotryon no doubt play a regional role in 

the dispersal, although we know of no reports of Zoobotryon being dispersed by ocean currents. 

Phyllodesmium poindimieri (Risbec, 1928) 

Introduced 

Wagner et al. (2007) report this Western Pacific and Indo-Pacific nudibranch (Rudman, 1991; S. 

Kahng, pers. comm., September, 2005), a known predator specialist of Carijoa, from Hawai‘i. It was 

first found in October 1995 on Maui (at Mā‘alaea) by C. Pittman and remains present at the site, 

where the nudibranchs usually remain concealed during the day near the base of the Carijoa colonies 

but move to the tips of the colonies at night, “looking like little iridescent pom-poms” (C. Pittman, 

pers. comm., September, 2005). They are also known from O‘ahu as of the late 1990s as well, based 

upon a photograph by J. Hoover (C. Pittman, pers. comm., September 2005) and are still found at 

various sites on O‘ahu (S. Kahng, pers. comm.). 

Class Bivalvia (mussels, oysters, scallops, clams, and shipworms) 

Introduced Bivalvia 


Anomia nobilis


79 



Mytilus galloprovincialis 1998 SF Mediterranean Sea 

Status: Failed 

Pinctada fucata martensi 1956 R Japan 

Tridacna crocea 1951 R American Samoa 

Tridacna squamosa 1983 R Palau 

Meretrix meretrix 1926 R Japan 

Mya arenaria 1923 R California 

Tivela stultorum 1924 R California 

Pustulostrea tuberculata 1956 R Australia 

Ostrea conchaphila 1893 R California 

Mercenaria mercenaria 1930s; 1967–1968 R Connecticut 

Status (above nine species): Released not established 

Mytilidae 

Mytilus galloprovincialis Lamarck, 1819 

Failed 

A naturally settled population of this originally Mediterranean mussel was discovered in the ballast 

tank of a resident submarine in September 1998 in Pearl Harbor, having been derived from larvae 

spawned by adult mussels arriving on the bottom of the USS Missouri in June of that year from Puget 

Sound in the state of Washington (Apte et al., 2000). No further specimens have been found in the 

fouling communities in Pearl Harbor or elsewhere. As Apte et al. (2000) note, M. galloprovincialis 

is established in other subtropical environments, such as Hong Kong, and it thus not impossible that 

a warmer-adapted genetic strain of this mussel (such as those in southern California, or in Hong 

Kong) could become established in Pearl Harbor or elsewhere on the islands. 

Anomiidae 

Anomia nobilis Reeve, 1859 

Introduced 

We suggest that this brackish-water jingle shell, although described from the Hawaiian Islands, is an 

early ship-fouling introduction from the Indo-West Pacific Ocean. Its specific geographic origin 

remains unknown at this time. It has not yet apparently been reported as a fossil in the Islands. It is 

unlikely to have been missed in collections of some two or three decades prior to the 1850s [for 

example, Conrad’s (1837), description of shallow-water endemic Hawaiian brackish water mytilids]. 

It may thus be another example of a species being first described from the site of introduction. 

Genetic studies of this species would be especially compelling. 

Anomia nobilis was described by Reeve (1859) from the “Sandwich Islands” as the “noble 

anomia” from the Cuming Museum collections but with no further data. Bryan (1915) recorded it 

from “Ford’s Island, Pearl City.” Edmondson (1933) noted that it was “under surface of stones in 

shallow water”; Dall et al. (1938) recorded it on waterlogged boards in Honolulu Harbor, from Pearl 

Harbor, and from off Honolulu in 31 to 60 m. Edmondson & Ingram (1939) found a naval vessel at 

anchor for 3 months in Pearl Harbor to be heavily fouled with Anomia nobilis. Edmondson (1944a) 

commented that “Anomia is seen on the hulls of boats more frequently than is Ostrea, and this fact 

may be accounted for by its mode of life and rapid development.” Long (1974), Grovhoug (1979), 

and Henderson (1990) reported it from Pearl Harbor, on settlement panels both inside and outside of 

the Harbor. Kay (1979) commented that it was one of the “characteristic fouling organism in 

Hawaiian waters and numerous specimens may be piled one on top of the other on waterlogged 

boards and pilings in Pearl Harbor and Kāne‘ohe Bay.” Hurlbut (1991c) reported on settlement and 

juvenile survival in Pearl Harbor. It was more recently reported in Pearl Harbor in 1996 (Coles et 

al., 1999a), Honolulu Harbor in 1997 and Barbers Point Deep Draft Harbor in 1998 Coles et al., 

1999b), Kāne‘ohe Bay in 1999 (Coles et al. 2002a), Waikīkī in 2001 and Hawai‘i Kai in 2002 (Coles 

et al. 2002b), Kaua‘i in 2002 and Maui in 2003(Coles et al. (2004, 2006).

80 


Ostreidae 

Crassostrea virginica (Gmelin, 1791) 

Introduced 

The Eastern, Virginia, or Atlantic oyster, Crassostrea virginica remains established in Pearl Harbor 

(Coles et al., 1999a), as a result of what is widely regarded as large plantings made in 1895 by J.F. 

Colburn of Honolulu. Colburn provided a detailed history of his attempts in 1893 and 1895 to D.S. 

Jordan and B.W. Evermann, who published Colburn’s account in 1905. In 1895 Colburn imported 

over 38,000 oysters from San Francisco Bay by means of five voyages of the steamship Australia. 

About two-thirds of the oysters were transported in boxes on the open deck and washed down each 

morning with seawater; the remainder were carried in a refrigerated compartment: “With the former 

way my loss was more in number, but the latter way was the most expensive” Colburn wrote, noting 

that the deck oysters could be transported for $10 per ton, whereas the refrigerated oysters cost 

five cents per pound for freight. 

Pilsbry (1918) noted that, “Professor Bryan [1915] has recorded the unsuccessful attempts to 

introduce eastern and California oysters. It could not reasonably be expected that oysters from waters 

so much colder would thrive, and so far as is known, they have died out completely. There is a large 

valve of the Virginia oyster in the Bryan collection, picked out of material dredged in Honolulu Harbor, 

probably a ballast shell.” This “ballast” shell, dated 1917, is now in the Bishop Museum collections; 

rather than being from ballast, it probably represents a shell from the Pearl Harbor plantings in 1895. 

The history of plantings of this oyster in Hawai‘i is given by Jordan & Evermann (1905), Bryan 

(1915), McClellan (1938), Anonymous (1940), Brock (1952, 1960), and Kay (1979). These are summarized 

in the table below. Carlton & Mann (1996, Table 1) give these Hawaiian records as “1866; 

1883 to 1949 and perhaps later years”; however, their table should read, “1866; 1883 to 1929”, as 

later releases (see below) were only movements within the islands. A transport event of 1871 noted 

by Bryan (1915) and McClellan (1938) refers only to live oysters received for consumption from 

California and not an outplanting into O‘ahu waters (Cobb, 1902). 

Plantings of the Atlantic Oyster Crassostrea virginica on O‘ahu 


1866 Pearl Harbor “cans” McClellan, 1938 (aboard the Comet, Captain John Paty; 

oysters “handed over to J. H. Coney, Honolulu”) 

1877 Honouliuli ? Kay, 1979: 19, by James Campbell 

1883 Kalihi Bay 300 Jordan & Evermann, 1905; Bryan, 1915; McClellan, 1938; 

oysters from San Francisco planted by Allan Herbert; 

heavy freshets from stream covered them with mud. 

1890 Moanalua ca. 100 McClellan, 1938: plantings by S. M. Damon from San 

fishpond 

Francisco; in April 1892 “three young oysters had grown” 

[= settled?] on the coral slab where oysters had been 

planted. 

1893 Manana, Ewa, 1,000 Jordan & Evermann, 1905; Brock, 1952; McClellan, 1938; 

Pearl Harbor 

plantings from California made by J. F. Colburn 

1895 Manana, ‘Ewa, 38,614 Jordan & Evermann, 1905; Bryan, 1915; Brock, 1952; 

Pearl Harbor 

from California made by J. F. Colburn. 

1921 ‘Aiea Bay, 2,000 seed Coleman, 1923 and McClellan, 1938 (5 barrels, from Great 

Pearl Harbor 

South Bay, Long Island, that had been replanted in 

Tomales Bay, California). 

Kāne‘ohe Bay 3,000 seed McClellan, 1938; Brock (1952, as Coral Garden, He‘eia) 

1922 Kāne‘ohe Bay a shipment McClellan, 1938 

1922 ? ? Coleman, 1923 

1923 Kāne‘ohe Bay ? Brock, 1952: 118 

1923 Pearl Harbor ? Anonymous, 1940 (in error?) 

1924 Kāne‘ohe Bay ? Brock, 1952: 118 

1924 Pearl Harbor ? Anonymous, 1940 (in error?) 

1926 Kāne‘ohe Bay “small Anonymous, 1940 

shipment” 

1929 Hanaloa Pond 2 barrels Brock, 1952: 118 

1929 Mokapu 2 barrels Brock, 1952: 118


81 

Edmondson (1933) noted that while the Atlantic oyster was growing in Kāne‘ohe Bay, “there is no 

conclusive proof that it spawns in these waters.” Anonymous (1940), noting the 1920s plantings in 

Kāne‘ohe Bay and Pearl Harbor, reported that “Those planted in Kaneohe Bay did not thrive but 

those sown in Pearl Harbor did well for a time. At present one can find them in scattered places, but 

not enough for commercial purposes.” However, at the same time, Edmondson & Wilson (1940) 

reported that the plantings in Pearl Harbor “were quite successful and there is evidence that spawning 

and setting of spat occurred.” Edmondson (1949a) noted that “Several examples of shells of 

Ostrea virginica were presented by Spencer Tinker. This large oyster has now become quite well 

established in certain areas of Pearl Harbor, from plantings of many years ago. However, there is a 

closed season throughout the year against taking the oyster which will continue until productivity 

reaches a higher level.” Sakuda (1966) studied the reproduction cycle in West Loch, Pearl Harbor. 

Hartman (1966) recorded the spionid worm Polydora websteri and the phyllodocid worm Eumida 

sanguinea from mud blisters in shells of C. virginica collected in June 1966 in Pearl Harbor. Preston 

(1971) used material from the West Loch, Pearl Harbor, in studying chromium-51 accumulation. 

Evans et al. (1972) again report it from Pearl Harbor based upon 1971 specimens. Sparks (1962) 

reported that there were 150,000 square yards (125,415 m 2 ) of C. virginica in Pearl Harbor, representing 

over 35,600,000 oysters ranging in density from 67 to 406 per square yard (56 to 339 per 

square meter), with spawning probably occurring in late spring or summer. 

In July 1996 we examined an exposed bed of Crassostrea virginica at low tide in the upper part 

of the West Loch, Pearl Harbor. A predator (likely a crab) had created a series of distinctively broken 

shells throughout the oyster reef. Outside of Pearl Harbor, Kay (1979) reported C. virginica from 

Kāne‘ohe Bay and from fishponds on Moloka‘i. Bailey-Brock (1990) reported that the spionid worm 

Polydora nuchalis was found in June 1988 at Kahuku, O‘ahu, in a Crassostrea virginica farm. The 

oysters were being grown in the effluent water from a commercial shrimp (Litopenaeus vannamei) 

farm. There is a small record of interisland movements, doubtless more frequent than recorded. 

Brock (1952) noted such movements in 1940 and 1949, and Kay (1979) reported that in 1939, 2.5 

tons of oysters were removed from Kāne‘ohe Bay and distributed to other localities [citing Brock, 

1960; however, that information is not in Brock (1960) nor in Brock (1952)]. In the early 1970s C. 

virginica was recorded as having suffered extensive mortalities in Pearl Harbor. The fact that the 

population survived has not been clear in the literature since that time. In July, 1972 it was reported 

that “some 34 million oysters” were killed in West Loch, Pearl Harbor (Anonymous, 1972). Three 

reasons for this mortality have been suggested: 

(1) Disease: Kern et al. (1973) reported that “dermo”, the lethal, disease-causing dinophyceaen 

Perkinsus marinus, was found in the oysters in Pearl Harbor during and following 

the mass mortality. 

(2) Water quality/trophic conditions: A. Kay (pers. comm., 31 May 1991) noted that “most 

of the oysters died out when the sewage was diverted out of Pearl Harbor”, indicating 

a potential change in trophic conditions. 

(3) Increased sedimentation: Anonymous (1972) reported that while oysters had been 

sent off for pathological testing, in addition “a large number of oysters were buried 

during the past three years as a result of mud runoff from land cleared for development 

but not yet built on. In West Loch, a new storm drain empties silt into the oyster 

beds with every rain.” Increased sedimentation has also been cited as a possible 

cause by M. Hadfield (pers. comm., 1995). 

Crassostrea virginica apparently recovered over the next quarter century and remains in Pearl 

Harbor. Coles et al. (1999a) reported that “West Loch areas receiving highly turbid runoff and sedimentation 

were dominated by Crassostrea virginica wherever hard substrata were available for settlement 

and growth.” 

Crassostrea gigas (Thunberg, 1793) 

Introduced 

The Pacific or Japanese oyster is well established at least in Kāne‘ohe Bay. Records of known plantings 

are as follows:

82 


Plantings of the Pacific Oyster Crassostrea gigas in the Hawaiian Islands 

Date Where Number Reference 

1926 Kalihi, Kāne‘ohe Bay “a small shipment” Anonymous, 1940 (all died within 2 years); 

Brock, 1952 

1935 Mōkapu two boxes McClellan, 1938 

1938 Pearl Harbor 2,000 Brock, 1952 

Mōkapu 500 

1939 Kāne‘ohe Bay: 210 cases Anonymous, 1940 (210 cases containing 

Coconut Island, Mōkapu 

ca. 2.5 million oyster spat); Brock (1952, as 

203 cases with 2,150,000 oysters); see text 

Anonymous (1940) reported that over 2.5 million spat released in 1939 (see table above) thrived and 

did well, until dredging in Kāne‘ohe Bay for the Marine Corps Base commenced, causing the resulting 

sedimentation to smother a large number of the oysters; a flatworm (Stylochus) was said to have 

preyed on the remaining oysters. 

Edmondson & Wilson (1940) noted that in March 1939 more than one million spat were planted 

in Kāne‘ohe Bay: “Exceptionally rapid growth has taken place during the past four months and 

it is hoped that this species may become permanently established in Hawaii.” Brock (1952, 1960) 

thought it had not established, but Kay (1979) noted that it was abundant in Kāne‘ohe Bay, attached 

to pilings, cement walls, and coral heads in the intertidal zone. Bailey-Brock & Ringwood (1982) 

noted “natural populations” in “Kaneohe Bay and Pearl Harbor, Oahu.” It remains abundant in 

Kāne‘ohe Bay, especially along the south shore, in 2003. 

Saccostrea “cucullata” (Born, 1778) 

Introduced 

= Ostrea cucullata 

Early, relatively small, plantings of this species, from Australian sources, on O‘ahu (Kāne‘ohe Bay, 

Kalihi Basin, Mōkapu, West Lock, and Sand Island) in 1928–1929 [Anonymous, 1940; Edmondson 

& Wilson, 1940; Brock, 1952; and perhaps as early as 1925 (Galtsoff, 1964)] failed to result in permanent 

populations at the time. Paul Galtsoff examined specimens in 1930 in Kāne‘ohe Bay and 

found them to be “ripe and spawning” but noted that “During World War II the small population of 

this species was destroyed by dredging operations.” 

However, S. cucullata was detected to have colonized Pearl Harbor in 1996 surveys (Coles et 

al., 1999a). We interpret it as a hull-fouling ship introduction; when this invasion occurred is uncertain 

but is likely to have been in the last quarter-century. Saccostrea cucullata is described as a widespread 

oyster, occurring from West Africa around the Cape of Good Hope into the Indo-West Pacific 

to southern Japan, Australasia, China, and the Philippines (Carriker & Gaffney, 1996). However, 

genetic analyses (Lam & Morton, 2006) reveal a complex of species under the morphoname “cucullata“, 

which we thus place in quotation marks here, pending resolution as to which clade the introduced 

Hawaiian stock represents. 

Gryphaeidae 

Hyotissa hyotis (Linnaeus, 1758) 

Intercept 

Paulay (1996) reported this well-known Indo-Pacific oyster-like bivalve as a “new state record” for 

the Islands. However, the specimens represent shells taken from vessel hulls in Pearl Harbor circa 

1950–1951, both of which ships were almost certainly recently arrived from the South Pacific. We 

do not further consider it here. It is of interest to note that it has recently become established in the 

Florida Keys (Bieler et al., 2004), and it would not be surprising if it does eventually find its way to 

the Islands.


83 

Chamidae 

Paulay (1996) reported Chama lazarus Linnaeus, 1758 and Chama pacifica Broderip 1835 from 

Pearl Harbor as “new state records” for Hawai‘i, based upon specimens collected in 1950–1951. 

U.S.Navy Lieutenant Commander H. Ryder sampled vessel bottoms in Pearl Harbor in 1950 and 

1951 and either presented the material to, or it came into the hands of, the shell collector A.E. 

Mehring. Because these species had not been collected before 1950–1951, because there are references 

to a “South Pacific” ship bottom, and because no other fouling studies (such as those of C.H. 

Edmondson) from the 1930s through the 1970s reported these bivalves, we conclude that Lt. Com - 

mander Ryder sampled vessels that had become fouled in foreign ports, and that these 1950–1951 

specimens do not represent actual Hawaiian populations. 

However, both species were subsequently collected in Pearl Harbor surveys in 1996 (Coles et 

al., 1999a). We suggest that these modern populations are not linked to the intercepts of Lt. Com - 

mander Ryder some 50 years earlier. 

Chama fibula Reeve, 1846 

Introduced 

= Chama hendersoni Dall, Bartsch, & Rehder, 1938 

This small South Pacific bivalve (described from the Philippines) was present by 1915 in the Islands 

(Bryan, 1915, plate 104, figure 21, this species, fide Dall et al., 1938). It was described as a new 

species, Chama hendersoni by Dall et al. (1938) from specimens from Pearl Harbor. Dall et al. 

(1938) noted additional material from Honolulu Harbor, including a valve dredged off the entrance 

of Honolulu Harbor in shallow water. Kay (1979) noted that C. hendersoni was a synonym of C. 

fibula, known from the Philippines and recorded from Sydney Harbour, Australia, and noted that it 

was known only from Pearl Harbor and Honolulu Harbor, although it was “very common in beach 

drift at Midway.” Coles et al. (1999a) record it from Pearl Harbor, Honolulu Harbor, and Barbers 

Point Harbor(1999a); Coles et al. (2000a) report it in Kāne‘ohe Bay based upon collections in 2000, 

at Waikīkī in 2001 (Coles et al. (2002b), at Nawiliwili Harbor in 2002 and in Hilo Harbor in 2003 

(Coles et al. 2004). 

Its absence from the fossil record of the Islands, its restriction to primarily harbor waters in 

O‘ahu, and its natural absence from Micronesia (Paulay et al., 2002) leads us to suggest that this 

species was introduced by ship fouling to O‘ahu, and perhaps from there to Midway. 

Chama macerophylla Gmelin, 1791 

Introduced 

Coles et al. (1999a, as Chama elatensis Delsaerdt, 1986) reported this species as established in Pearl 

Harbor, based on 1996 collections. Coles et al. (2002a) also report it (as C. macerophylla) from 

Kāne‘ohe Bay based upon collections in 1999. DeFelice et al. (2001) note that the Chama under consideration 

has been tentatively identified as the warm-water Western Atlantic species C. macerophylla, 

on the authority of G. Paulay. Chama macerophylla is known from North Carolina to Brazil, 

including the Caribbean and Bermuda (Abbott, 1974; DeFelice et al., 2001; Mikkelsen & Bieler, 

2007). 

Chama lazarus Linnaeus, 1758 

Introduced 

This southern hemisphere chamid was first collected in 1996 in Pearl Harbor (Coles et al., 1999a), 

and was also found in Honolulu Harbor in 1997 (Coles et al., 1999b). Paulay (1996) gives the range 

as East Africa to Tonga, in Western Polynesia; Matsukuma (2000) as Amami Islands to northern 

Australia and the Indian Ocean, living on rocks in the lower intertidal zone to 20 m. As discussed 

above, we take 1996, and not 1950, as the date of first collection. 

Chama pacifica Broderip, 1835 

Introduced 

This southern hemisphere chamid was first collected in 1996 in Pearl Harbor (Coles et al., 1999a). 

Paulay (1996) gives the range as Thailand to the Line Islands. Matsukuma (2000) as the Kii 

Peninsula southward to Southeast Asia and the Indian Ocean, living, as does C. lazarus, on rocks in 

the lower intertidal zone to 20 m. As discussed above, we take 1996, and not 1950, as the date of 

first collection.

84 


Myidae 

Sphenia coreanica Habe, 1951 

Introduced 

Coles et al. (1999a) and Coan (1999) report the establishment of this Japanese species in Pearl 

Harbor. The name is tentative, awaiting further study of more mature material from Asia. It was first 

reported from Pearl Harbor (1968–1972) by Long (1974) as Sphenia cf. fragilis (H. & A. Adams, 

1854) in fouling assemblages at 9 m depth. Sphenia fragilis is a closely related species from the 

warm Eastern Pacific which could easily be confused with S. coreanica. 

Semelidae 

Abra sp. 

Introduced 

Coles et al. (1999a) record this semelid as introduced to the Pearl Harbor, based on 1996 collections. 

veneridae 

Venerupis philippinarum (A. Adams & Reeve, 1850) Introduced 

= Tapes philippinarum 

= Tapes semidecussata Reeve, 1864 

= Tapes japonica Deshayes, 1853 

= Tapes philippinarum okupi Bryan, 1919 (described from Hawai‘i) 

= Paphia philippinarum (as used by Edmondson & Wilson, 1940). 

This species is often placed in the genus or subgenus Ruditapes; see Carlton (1992) and Paulay 

(1996) for taxonomic notes. The Japanese littleneck clam Venerupis philippinarum was apparently 

intentionally introduced at least twice to the Islands. Bryan (1919) reported that populations existed 

at ‘Ewa and Pearl Harbor between the late 1880s and about 1890. They reappeared about 1918–1919 

and became abundant at Kalihi and Moanalua on the mudflats (Bishop Museum collections contain 

a large suite of shells collected by Bryan in 1918 in Kalihi Harbor). Thaanum (1921) noted that, “A 

Japanese now living in Honolulu has twice planted this bivalve on the mud flats at Moanalua on 

Oahu” in apparent reference to the failed first and the successful second plantings. Thaanum noted 

that the clams “are frequently imported from Japan to Honolulu by the barrel for sale among the 

Japanese”; it is likely that some of these were released alive. Edmondson (1933) noted that “since 

its introduction it has multiplied rapidly in shallow bays about Oahu and is now used extensively as 

food.” Dall et al. (1938) noted material collected by Thaanum. They quote Thaanum as follows: 

“Bought in fish markets in Honolulu in 1918, collected by vendor at Maunalua [sic], Oahu. Inquiry 

brought information that a certain Japanese had planted them there. This man was located in Honolulu 

by Langford who was told by him that he had made several attempts at planting shells imported alive 

from Japan as the Japanese are fond of the clam, but only recently plantings had succeeded, and the 

shells seemed to be now well established. Barrels of live shells frequently shipped from Japan are 

inspected by Mr. Langford before being released. [some of these specimens] are samples from these 

importations. Japanese emigrants also frequently bring live shells in their baggage of this species, and 

also of species of Tivela and a large univalve (Rapana thomasi Crosse).” 

Of specimens from Kēōkea, Hilo, Thaanum (1921) wrote, “After visiting this pond at least once a 

month for over twenty years without ever seeing this species, I suddenly found them one day in considerable 

numbers — all dead, but with valves still joined. I must conclude that they are the victims of 

an attempted planting, especially as several Japanese families live close by. No live shells could be 

found.” Edmondson & Wilson (1940) reported 10 barrels from Japan were planted in Kalihi Basin, 

Pearl Harbor, and Kāne‘ohe Bay in 1920, and that it was called the “Damon clam” because a Mr. 

Damon was instrumental in transplantations. They also note that in 1937, 400,000 young specimens 

were transplanted from Kalihi Basin and Pearl Harbor to localities on Kaua‘i, Hawai‘i, Lāna‘i, Maui, 

and O‘ahu [this appears to be the record noted in Kay (1979) as 400,000 pounds transported in 1957]. 

Brock (1952) summarized the history of Venerupis in Hawai‘i and gave details of records from 

1920 to 1937. Brock (1960) noted it was well established on O‘ahu, Moloka‘i, and Hawai‘i, and possibly 

on Maui and Kaua‘i, and Withington (1966) reported that it had been planted again in Kāne‘ohe 

Bay. Kay (1979, pp. 20, 570) and Yap (1978) discuss the fishery in the 1960s. Withington (1966), in


85 

a long story in the Honolulu Star-Bulletin, noted that 10,000 clam diggers turned up on September 

1, 1966, the opening day of clamming season, to search for clams in Kāne‘ohe Bay. Titcomb (1979) 

reproduced a photograph of “opening season” for taking Venerupis philippinarum in Kāne‘ohe Bay 

in September, 1968, showing hundreds of people on the tidal flats. She states that “Following the 

1968 season, however, soil erosion, associated with heavy rain, contributed to the demise of the 

Kaneohe Bay population.” The population in Kāne‘ohe Bay has decreased considerably since 

sewage release ceased in the Bay in 1977–1978 (R. Kinzie, pers. comm., 2001; see also Hunter & 

Evans, 1995). It was reported at 12 of 15 sites in Pearl Harbor in 1996 and at one site in Kāne‘ohe 

Bay in 1999 (Coles et al., 1997, 1999a). Yap (1978) studied aspects of the biology of Venerupis in 

Kāne‘ohe Bay. Peak spawning occurs in January and February; new recruits appear from April to 

June. The native crabs Thalamita and Calappa and the native gastropod Natica gualteriana are predators 

of Venerupis [Higgins, 1969 (Natica), Yap, 1978 (crabs)]. Cheng & Rifkin (1968) reported cestode 

larvae from Venerupis in Kāne‘ohe Bay. 

Venerupis philippinarum is known in Hawai‘i as okupi (see Bryan, 1919 for origin of the name) 

or the Damon clam. It also goes by the name of pupu ‘olepe (Titcomb, 1979). 

Lioconcha fastigiata (Sowerby, 1851) 

Introduced 

Goodwin (2003) reported (as Lioconcha fastigata) this relatively small south and western Pacific 

venerid from 0.3 to 30 m of water near “the Pearl Harbor entrance.” Based upon the photographs 

given by Goodwin, Gustav Paulay agrees with this identification and its prior absence from the 

Hawaiian biota (G. Paulay, pers. comm., June 2003). Goodwin (2003) reported that most specimens 

were “buried in silty sand only a few inches from the surface.” He suggested that L. fastigiata may 

have been introduced in ballast water. The first specimens were collected about December 1985 (D. 

Goodwin, pers. comm., July 2003). 

Mercenaria mercenaria (Linnaeus, 1758) 


The Annual Report of the Hawaii Department of Land and Natural Resources for 1967–1968 

(Hawaii Department of Land and Natural Resources, 1968) noted that two large specimens of this 

well-known North American Atlantic clam, the quahog, were collected in 1967 in Kāne‘ohe Bay. 

These were believed to be linked to an authorized private introduction in the 1930s and would thus 

have to be clams about 30 years old, within the known age of M. mercenaria. Also in 1967–1968, 

the State imported 985 Mercenaria from Connecticut that were planted in Kāne‘ohe Bay and in the 

Nomilo fishpond on Kaua‘i. The next year’s report (Hawaii Department of Land and Natural 

Resources, 1969) recorded that had grown slightly and largely survived. Kay (1979, p. 20) stated that 

Mercenaria had “apparently settled and are reproducing.” 

No populations of Mercenaria mercenaria are known to have survived in Hawai‘i. It is listed 

at: http://www.hawaiiaquaculture.org/products.html [accessed February 2007] as a species currently 

under commercial aquaculture production in Hawai‘i. 

Hiatellidae 

Hiatella arctica (Linnaeus, 1767) 

Introduced 

= Saxicava hawaiensis Dall, Bartsch, & Rehder, 1938 fide Kay, 1979 

One or more species of fouling and nestling bivalves bearing this name now occur in temperate and 

subtropical harbors worldwide. Dall et al. (1938) described Hawaiian populations as a new species, 

Saxicava hawaiensis. The type material of S. hawaiensis was dredged by Thaanum on a coral reef 

at Fort Armstrong and off Honolulu Harbor, both in shallow water. These collections were likely 

made before 1920 (Dall et al., 1938, p. 5). 

Evans et al. (1972; 1971 collections), Kay (1979), Grovhoug (1979), and Grovhoug & Rastetter 

(1980) all reported if from Pearl Harbor; Grovhoug & Rastetter (1980) add Kāne‘ohe Bay, based on 

1976–1977 collections. Coles et al. (1999b) record it from Honolulu Harbor, Ke‘ehi Lagoon, and 

Barbers Point Harbor. Coles et al. (2002a) record it at Mokumanu Island off Kāne‘ohe Bay in 2000, 

at Waikīkī in 2001 and Hawai‘i Kai in 2002 (Coles et al. 2000b). Coles et al. (2004, 2006) report it

86 


in 2002 at Kaua‘i, and 2003 Moloka‘i, Maui, and the island of Hawai‘i. We regard it as an early 

introduction with ship fouling. 

Pholadidae 

Martesia striata (Linnaeus, 1758) 

Introduced 

= Martesia hawaiensis Dall, Bartsch & Rehder, 1938 [type locality Pearl Harbor] vide Kay, 1979 

Martesia was first reported from Pearl Harbor by Gordon (1916) and Bartsch (1916). We take 1915 

to be the first year of record, based on Gordon’s statements; no earlier material is in the Bishop Mu - 

seum collections. Clapp & Kenk (1963) stated that Gordon reported that a “pile-driver scow, constructed 

of Douglas Fir covered with ships’ felt and sheathed with 1-inch Redwood planks, was 

inspected after 18 months of service at the Naval Station at Pearl Harbor, Hawaii. The outer sheathing 

showed practically no damage from teredinids, but was attacked by a borer which produced a 

hole ⅛ inch wide and increasing to ¾ inch as it penetrated deeper. Asphalt and felt did not impede 

the progress of the [Martesia]. The author recommends that the scow be sheathed with copper rather 

than Redwood.” 

Clapp & Kenk (1963) noted that Bartsch (1916) commented on Gordon’s paper and photograph, 

stating that the “photographs show a Martesia and shipworms (Xylotrya and Teredo) in the 

timber... Martesia may bore into rocks and wood, but is confined to the outer two inches.” 

Anonymous (1917), at about the same time, indicated that Martesia”... was reported by the Bureau 

of Yards and Docks to cause considerable trouble in timber structures around the Hawaiian Islands. 

Specimens were taken from timber near the Pearl Harbor dry dock. In addition to wood, this mollusk 

is reported to bore through tar paper and asphalt coverings, and even rock ... .” Miller (1924a), 

reporting upon formal studies initiated in 1922, also found M. striata in Pearl Harbor, and noted 

“burrows attributed to this species” in a wood test block in Nawiliwili Bay, Kaua‘i. Dall et al. (1938) 

described it as a new species (above) from Pearl Harbor, and noted additional material from a 

beached drift log at Ka‘a‘awa, O‘ahu. Henderson (1990) reported specimens collected in Pearl 

Harbor in 1986. It remains common in fouling in Pearl Harbor (Coles et al., 1999a). 

Martesia is now recorded from circumtropical and circumsubtropical seas globally (Turner & 

Santhakumaran, 1989). Its origin remains unknown, but it is likely to be the southern hemisphere, 

and perhaps the southwestern Pacific Ocean. It may have commenced its global voyages as early as 

the 1500s in wooden ships. 

We regard it as introduced with ship fouling and boring communities. No fossil Martesia are 

known from the Islands. As with another introduced clam, Venerupis philippinarum, Martesia has a 

Hawaiian name: Olepe-naka-loa, meaning “long face” (Titcomb, 1979). 

Teredinidae 

Bennett (1840) may have been the first to inadvertently record the earliest marine introduction in the 

Hawaiian Islands during his visit there in 1835. Relative to shipworm activity, he noted “The Teredo 

navalis called, par excellance, ‘the worm’ by sailors, infests the coast of Oahu in sufficient numbers 

to do serious injury to a boat left but for a single night exposed to their attacks.” As we regard no 

shallow-water shipworm as native to the Islands, this statement likely represents a ship-inoculated 

invasion of one of the many introduced shipworms. 

Much later, Edmondson (1962b) wrote “... the appearance of exotic species of shipworms in 

Honolulu and Pearl Harbors during World War II is substantial evidence that surface craft are still a 

factor in the dispersal of marine wood borers.” Unfortunately Edmondson never specified which 

species he had in mind of shipworms appearing in World War II. Edmondson (1940) noted that “In 

the earlier surveys no species of Bankia were observed in Hawaii ... . The recent appearances of ... 

one or more species of Bankia introduces additional and perhaps more destructive members to the 

catalogue of shipworms invading Hawaiian waters.” It may be that by the 1960s Edmondson had in 

mind this presumably 1920s–1930s episode, albeit pre-war. Edmondson (1940) further noted that 

“Boats of many descriptions, including wooden-bottomed fishing sampans of long cruising ability, 

offer easy transportation among the islands of the Hawaiian archipelago for marine boring organisms.”


87 

We regard the shallow water, fixed-wood dwelling shipworms of the Hawaiian Islands as shipintroduced 

species. These introductions may have commenced with the arrival of the first European 

wooden vessels in the Islands. Edmondson (1940, 1946b, 1946c, 1946d, 1962b), and other workers, 

have often referred to the role of ships in dispersing teredinids globally and specifically to the 

Islands. However, the global shipworm literature in general rarely identifies, or speculates, which of 

two processes—natural dispersal in drifting wood, or human-mediated dispersal in ships—may play 

the more important role for which species, or, indeed, when it may play the only role (Carlton, 1999). 

Rather, both processes are presumed to be amalgamated with, by implication, natural drift playing 

the more fundamental and important role. 

As a result, introduced shipworms are rarely identified as such for many ports of the world, and 

the post-Edmondson era literature on Hawaiian teredinids makes no reference to their potential nonindigenous 

status. An examination of the detailed studies of Edmondson reveals, however, that while 

there is an oceanic (high seas) drift teredinid fauna that comes ashore on the islands, the harbordwelling 

species are not found in such drifting materials. The native (naturally occurring) pelagic 

drifting shipworms of the Central and North Pacific Ocean include Lyrodus medilobatus (Edmond - 

son, 1942), Teredo princesae Slvickis, 1928 (= Teredo gregoryi Dall, Bartsch, & Rehder, 1938), and 

Teredo triangularis Edmondson, 1942. Edmondson (1962b) further considered Teredo clava and 

Teredo palauensis as species that have adopted a “typically oceanic existence.” 

Paulay (1996) commented on Roch’s (1976) revisions of the names of Hawaiian teredinids; we 

follow Paulay’s conclusion here and retain the nomenclature and taxonomy of Turner (1966). 

A modern survey of the shipworms of the Hawaiian Islands has not been undertaken. It may 

well be that the six introduced species treated here have been joined in recent decades by additional 

nonnative shipworms. A modern-day assessment of the economic damage of shipworms in the 

Islands is also lacking. Earlier assessments are largely anecdotal: 

“A few years ago a long wooden trestle was built off Sand Island opposite Honolulu Harbor to carry 

the equipment used in laying a concrete sewer outfall. Since the trestle was needed only temporarily, 

it was not treated for protection against marine borers. But the shipworms won the race; in less than 

70 days large sections collapsed, dropping heavy machinery into the sea.” Smith (1956). 

We take this damage to have been caused by one or more of the shipworms treated below. 

Bankia bipalmulata (Lamarck, 1801) 

Introduced 

= Bankia hawaiiensis Edmondson 1942 (from Honolulu Harbor, 1939) 

= Bankia konaensis Edmondson 1942 (from submerged algaroba branch in Kealakekua Bay, Kona, Ha - 

wai‘i). 

This Southern Hemisphere shipworm [the type locality is questionably India (Turner, 1966)] was 

first collected in 1939 (Edmondson, 1942, p. 136) and described as two different new species. As B. 

hawaiiensis, Edmondson (1942) reported it from O‘ahu (Pearl Harbor, Honolulu Harbor, Hanauma 

bay, and in drift timbers at Kalihi Entrance), Kaua‘i (drift materials at Wailua), and Hawai‘i (Kona 

coast, in Kealakekua Bay and at Kahalu‘u). Edmondson (1944c) recorded Bankia hawaiiensis in test 

blocks to about 23 m in O‘ahu. Long (1974) also reported it off O‘ahu. 

Lyrodus affinis (Deshayes, 1863) 

Introduced 

= Teredo milleri Dall, Bartsch & Rehder 1938, nomen novum for T. affinis, but an unnecessary nomenclatural 

change (see Turner, 1966, pp. 86 and 111; name change also in and rejected by Edmondson, 1940, 

p. 245). 

Miller (1924a) recorded L. affinis from Nawiliwilli, Kaua‘i, in 1923, and also noted its presence in 

Honolulu Harbor. Edmondson (1940) noted that on O‘ahu it “dominates in Hanauma Bay, on 

Waikiki Reef, and in the Ala Wai Canal, and also occurs in Kaneohe Bay.” Edmondson (1940, p. 

248) noted that this shipworm was “a constant menace to small craft occupying the anchorage at the 

mouth of the Ala Wai Canal.” Edmondson (1942) noted that it was “almost wholly absent from test 

blocks in Pearl Harbor”, and added additional records from Kaua‘i, Maui, Moloka‘i, and Hawai‘i. 

Edmondson (1944c) found it at 23 m in test blocks off O‘ahu, and Long (1974) also recorded it from 

off O‘ahu. We regard this shipworm as probably native to the Southern Hemisphere.

88 


Lyrodus pedicellatus (Quatrefages, 1849) 

Introduced 

= Teredo hawaiensis Dall, Bartsch & Rehder 1938, from 386 to 463 m (Albatross station 3810 [March 27, 

1902], as 211–253 fathoms, off the south coast of O‘ahu) from a dredged palm log (fide Turner, 1966, 

plate 1F caption), the name applied as a “new species” for the material identified by Miller (1924a) 

as Teredo bartschi. 

= Teredo diegensis Bartsch, 1916 (from San Diego) 

= Teredo honoluluensis Edmondson 1946b (from test block, Honolulu Harbor, O‘ahu) 

= Teredo kauaiensis Dall, Bartsch & Rehder,1938 (from Nawiliwili, Kaua‘i, as a new species, for material 

originally identified by Miller as Teredo diegensis [= Lyrodus pedicellatus]; Edmondson (1942) 

favored retaining diegensis, and placed T. kauaiensis in synonymy. Teredo kauaiensis was proposed 

in part because Dall, Bartsch & Rehder noted, “From what we know of the distribution of shipworms, 

we are disinclined to believe that this is T. diegensis.” 

= Teredo midwayensis Edmondson 1946 (from Midway Island) 

This “blacktip shipworm,” with a complicated global taxonomic history, has been spread for centuries 

by wooden sailing vessels. Based upon the number, frequency, and location of most records 

(including synonyms), it appears to be a species of southern hemisphere origin, although typical of 

an exotic, ship-dispersed species, the type locality is Spain. Dall et al.’s (1938) comment, relative to 

proposing the name T. kauaiensis, is a classic example of overlooking the probable role of ships in 

creating the modern distribution of this shipworm. 

Curiously, it was first recorded in 1902 from Hawai‘i as a new species, Teredo hawaiensis, from 

a deep water log off O‘ahu. Lyrodus pedicellatus is not a deep water species; the log was doubtless 

derived from shallow water. It has been recorded from the Hawaiian Islands by Miller [(1924a, as 

Teredo bartschi.; Dall et al. (1938); Edmondson (1942, as Teredo hawaiiensis); and Edmondson 

(1946b)]. Edmondson (1942) noted its presence at Midway Island. 

Teredo bartschi Clapp, 1923 

Introduced 

=Teredo hiloensis Edmondson, 1942 (from Hilo Harbor) 

= not Teredo bartschi of Miller, 1924a (see Lyrodus pedicellatus) 

This shipworm, possibly also of southern hemisphere origin, was recorded by Edmondson (1942) 

from Hilo Harbor (from between 1935 and 1941). Edmondson (1946b, p. 221) confirmed that he 

concurred with R. Miller’s identification of this species from O‘ahu. It was described from Port 

Tampa, Florida, and Coan et al. (2000) refer to it as a Caribbean species. However, we regard it as 

introduced to the Caribbean (see also Carlton & Ruckelshaus, 1997). 

Teredo clappi Bartsch, 1923 

Introduced 

= Teredo trulliformis Miller 1924 (from Honolulu Harbor) 

This shipworm was first described from a ship’s keel at Key West, Florida, but we regard it as introduced 

to the western Atlantic, including Florida and the Caribbean (see also Carlton & Ruckelshaus, 

1997). It was recorded from Honolulu Harbor, Pearl Harbor, and Nawiliwili, Kaua‘i, as a new 

species by Miller (1924a; 1922–1923 collections). Further records are those of Edmondson (1940, 

from Waikīkī Bay, Kāne‘ohe Bay, and Ala Wai Canal), Edmondson (1942, submerged algaroba timber 

in Kealakekua Bay and at Pearl Harbor, Honolulu Harbor, Waikīkī, Ala Wai Canal, Kāne‘ohe 

Bay), Edmondson (1944c, in test blocks to 23 m off O‘ahu) and Kay (1979, common in Kāne‘ohe 

Bay). The introduced flatworm Taenioplana teredini is found in the burrows of this species. 

Teredo fulleri Clapp, 1924 

Introduced 

Edmondson (1942) records this shipworm from the Islands, although it is omitted in Kay (1979). We 

take the records to date between 1935 and 1941. Edmondson notes that it “was first recovered locally 

from submerged algaroba branches in Kalihi Entrance, O‘ahu, and later was taken from the same 

kind of wood in Kealakekua Bay, Hawai‘i, at Olowalu, Maui, and on the shore of Moloka‘i.” He also 

noted its occurrence in Apia, the seaport capital of Western Samoa, in the keel of a trading vessel 

seeing service between Samoan islands. Edmondson hypothesized that T. fulleri was native to the 

Central Pacific Ocean and not to the West Indies, from where it was described.


89 

Teredo furcifera von Martens, 1894 

Introduced 

=Teredo parksi Bartsch, 1921 (young T. furcifera; from Pearl Harbor) 

=Teredo bensoni Edmondson 1946 (see discussion, below) 

=Teredo furcillatus Miller of Dall, Bartsch, and Rehder 1938 (from Samoa) 

This shipworm was first described from the Molucca Islands in Indonesia (Turner, 1966). It was 

described again from the Hawaiian Islands in Pearl Harbor as T. parksi by Bartsch (1921), although 

no date of collection is again. Records from Hawai‘i include Miller (1924a, as Teredo furcillatus and 

T. parksi), Dall et al. (1938, as Teredo furcillatus and Teredo parksi Bartsch, from Pearl Harbor; also 

in block of wood at Waikīkī, O‘ahu), Edmondson & Ingram (1939, as T. parksi, the common shipworm 

destroying wood structures in Kāne‘ohe Bay and Pearl Harbor and infesting test panels), 

Edmondson (1942, as both T. parksi and T. furcillatus, with details of distribution on Kaua‘i and 

O‘ahu), and Edmondson (1946b, as Teredo bensoni, new species): “live mature specimens were 

recovered from the wooden guard rail of the dredger Benson on its return to Honolulu after completing 

operations at Canton Island. Four months after [its] return from Canton Island, a single immature 

shipworm closely resembling the type specimen was recovered from a test block in Honolulu 

Harbor near the dry dock where the Benson had been reconditioned.” 

Additional Records: Imported Nonnative Bivalves 

Over the decades a number of nonnative species of clams and other bivalves have been intentionally 

brought to Hawai‘i for mariculture or experimental purposes. Imported species are listed below. 

Details of importation are provided by Coleman (1923), Edmondson & Wilson (1940), Brock (1952, 

1960), Hanna (1966), Lachner et al. (1970), Kay (1979), Heslinga & Perron (1983), and Eldredge 

(1994). 

From the Indo-Pacific: 

Pinctada fucata martensi (Dunker, 1872) 


Hayami (2000) treats P. martensii and P. fucata (Gould, 1850) as separate taxa. 

Tridacna crocea Lamarck, 1819 


Tridacna squamosa Lamarck, 1819 


Meretrix meretrix Linnaeus, 1758 


Of these, we comment briefly on the last, Meretrix meretrix, as apparently adult shells were at one 

time characteristic of the Kāne‘ohe Bay shores, and it is possible that old shells could still be found. 

Edmondson (1933) noted that “A species introduced into Hawai‘i from Japan and occasionally seen 

in Hawaiian waters is Cytherea (Meretrix) meretrix Linnaeus. ... The species apparently finds a habitat 

in muddy bottoms. Many empty shells are to be found in certain localities along the south shore 

of Kaneohe Bay.” These were apparently the results of plantings made in 1926 (Brock, 1952). 

Plantings were made again in 1939 in Kāne‘ohe Bay (Brock, 1952), Edmondson & Wilson (1940) 

noting that 20,000 clams from Japan had been released and that “rapid growth has taken place.” 

Apparently the population survived robustly, moving Brock (1952) to categorize it as a species that 

was “either firmly established or appear(s) to be becoming established”, with Brock (1960) listing it 

under “Established Introduced Species”, and further commenting that it was “well established on 

Oahu and possibly on other islands as well.” After that, no further mention of its status is found in 

the original literature; it is not listed by Kay (1979). We presume that the population died out. We do 

not list it as “failed”, as there is no actual evidence of reproduction. 

From North America: 

Mya arenaria Linnaeus, 1758 

Apparently Released; Not Established 

Clinocardium nuttallii (Conrad, 1837) 

Released? 

or Protothaca staminea (Conrad, 1837) 

Tivela stultorum (Mawe, 1823) 


In addition, several nonnative oysters have been imported and planted in Hawai‘i. Details are provided 

by Anonymous (1940), Edmondson & Wilson (1940), Brock (1952, 1960), Galtsoff (1964), 

Hanna (1966), and Kay (1979). The two oyster species that did not survive are as follows:

90 


From the Indo-Pacific: 

Pustulostrea tuberculata (Lamarck, 1804) 


[in Hawaiian literature previously as Crassostrea amasa (Iredale, 1939)] 

From North America: 

Ostrea conchaphila (Carpenter, 1857) 


[in the Hawaiian literature previously as Ostrea lurida (Carpenter, 1857)] 

CRUSTACEA 

Copepoda (copepods) 

Introduced and Cryptogenic Copepoda 



Pseudodiaptomus marinus 1964 BW Japan 

Psammopsyllus stri 1979 SBA Panama (Caribbean) 

Teredicola typica 1939 SF Southern hemisphere? 


Haplostomides hawaiiensis 1985 SF 

Pseudodiaptomidae 

Pseudodiaptomus marinus Sato, 1913 

Introduced 

Jones (1966) reported this Japanese copepod as abundant in 1964 and 1965 in the Ala Wai Canal, 

O‘ahu, in water of 18 ‰. Maximum numbers were found near the inland end of the canal, about 2.4 

km from the mouth. Walter (1986, p. 147), in reviewing the complex taxonomic history of P. marinus, 

examined Jones’ material and concluded they were the same as Japanese populations. Fleminger 

& Kramer (1988) subsequently reported P. marinus from southern California embayments. We 

regard it as introduced by ballast water. It should be expected in other Hawaiian localities, such as 

Pearl Harbor. 

Leptopontiidae 

Psammopsyllus stri Mielke, 1983 

Introduced 

Kunz (1993) reported this interstitial harpacticoid copepod in coarse and medium sand, from Kaua‘i 

[Lumahai Beach, Wahina Bay (lagoon near the mouth), Anahola Beach] and from O‘ahu (Pūpūkea 

Beach), all collected in 1979. It was previously known only from the Atlantic Ocean, on the 

Caribbean coast of Panama. Noting that similar distributions were known for other interstitial copepods 

where the same species was believed to be on either coast of North or Central America, Kunz 

offered a variety of speculations, including geological processes and aquatic waterfowl, as to the origin 

of such broadly disjunct distributions, but only one that seems applicable to the occurrence of an 

Atlantic interstitial sand species on islands in the mid-Pacific Ocean. Kunz (1993) remarked that, 

“Eine Verfrachtung von Bestandteilen der Sandfauna durch Ballastsand von Segelschiffen ist nicht 

auszuschließen” [Transport as a member of the sand fauna in ballast sands of sailing ships is not 

impossible]. We consider Psammopsyllus stri to be introduced. It may further prove to be one of 

many examples of sand biota that must have been moved globally by sailing vessels up to the early 

20th century. 

Claussidae 

Teredicola typica Wilson, 1942 

Introduced 

Charles H. Edmondson found this parasitic copepod, the female of which clings tightly to the lining 

of the infrabranchial cavity of the host, in five species of the shipworm genus Teredo and one species


91 

of the shipworm genus Bankia in Honolulu Harbor (the first record was 1939), as well as at additional 

stations on O‘ahu, on Hawai‘i (Hilo Harbor), and on Maui (Kahului). It was described by 

Wilson (1942) as a new species with a type locality of Honolulu Harbor. Edmondson (1942, as 

Teredo milleri) noted it was in 75 percent of specimens of Lyrodus affinis in Honolulu Harbor; additional 

records were Kalihi Entrance, Waikīkī, and Hanauma Bay, all on O‘ahu. Edmondson (1942 

and 1945b) provided further notes on its biology and natural history. Wilson (1957) redescribed this 

species based upon specimens from Honolulu Harbor. Humes & Turner (1972) and McKoy (1975) 

reported it from Australia, New Zealand, and Japan. As this species is associated only with what we 

regard as introduced shipworms, we consider this copepod to be introduced as well. 

Ascidicolidae 

Haplostomides hawaiiensis Ooishi, 1994 

Cryptogenic 

Ooishi (1994) described this symbiotic copepod, the first copepod associate of an ascidian reported 

from Hawai‘i, from the introduced ascidian Polyclinum constellatum. The material was collected in 

1985 and again in 1993 on O‘ahu from Ke‘ehi Lagoon (Honolulu), in ascidians “attached to ropes, 

floats and small ships’ bottoms.” While ascidicolid copepods are not necessarily host specific 

(Ooishi & Illg, 1977; Marchenkov & Boxshall, 2003), we hypothesize that this copepod is also not 

native. Nevertheless, we treat it as cryptogenic, pending further information about its host specificity 

and distribution outside of the islands. 

Amphipoda (amphipods) 

Introduced and Cryptogenic Amphipoda 



Gammaridea: Caprellidae 

Caprella scaura 1929 SF, BW Unknown 

Caprella penantis 1921 SF, BW Unknown 

Caprella equilibra 1944 SF, BW Unknown 

Caprella danilevskii 1921 SF, BW Unknown 

Paracaprella pusilla 1937 SF, BW Unknown 

Gammaridea: other families 

Incisocalliope derzhavini 1967 SF, BW Japan 

Jassa falcata 1997 SF, BW North Atlantic 

Ericthonius brasiliensis 1935 SF, BW North Atlantic 

Monocorophium acherusicum 1943 SF, BW Northwest Atlantic 

Monocorophium insidiosum 1959 SF, BW Northwest Atlantic 

Laticorophium baconi 1967 SF, BW Northeast Pacific 

Grandidierella bispinosa 1996 SF, BW Indo-Pacific 

Grandidierella japonica 1992 SF, BW Japan 

Paraleucothoe sp. 1996 SF, BW Unknown 

Leucothoe micronesiae 1997 SF, BW Indo-Pacific 

Elasmopus “rapax” 1937 SF, BW North Atlantic 

Podocerus brasiliensis 1935 SF, BW North Atlantic 

Stenothoe gallensis 1935 SF, BW Unknown 

Stenothoe valida

92 


Gammaridea: Caprellidae (skeleton shrimp) 

“The wide dispersal of caprellids may be accounted for, without doubt, by their clinging habits and 

their customary association with fouling organisms, which are frequently transported on the bottoms 

of ships. Caprellids have been taken from fouled buoys and the bottoms of boats in Hawaii and elsewhere” 

(Edmondson & Mansfield, 1948). Caprellids are another group of crustaceans that achieved 

a wide global distribution via ship hull fouling centuries before systematic collections commenced. 

This, combined with unresolved species complexes in so-called “cosmopolitan” taxa, has prevented 

a clear resolution of the history and biogeography of potential invaders in many areas of the world. 

We consider at least four species of common Hawaiian caprellids to be nonnative species. 

Guerra-García & García-Gómez (2001) found that Caprella danilevskii and Caprella penantis 

inhabit areas of high exposure or strong currents, suggesting an ability to do well on ship bottoms 

on the high seas. 

Caprella scaura Templeton, 1836 

Introduced 

A now-global caprellid of unknown origin, described originally from “among marine plants at 

Riviere Noir, Mauritius” (Templeton, 1836) in the Indian Ocean, centuries after the first Portuguese 

vessels touched at those islands in the 1500s. It is now said to occur on most coasts of the world 

(McCain, 1968; Arimoto, 1976), with many purported synonyms (McCain & Steinberg, 1970; Ar - 

imoto, 1976). Bishop Museum material is from 1929 (Pearl Harbor), 1943 (Waikīkī, boat bottom), 

and later years. Edmondson & Mansfield (1948) illustrated specimens from Pearl Harbor and Wai - 

kīkī. Evans et al. (1972) also record it from Pearl Harbor (1971). Coles et al. (1999b) record it from 

Ke‘ehi Lagoon, Ala Wai Harbor, and Kewalo Basin, based upon 1997–1998 collections; it was also 

found in 2003 at Mā‘laea Harbor, Maui (Coles et al., 2004). We regard this species as a ship introduction. 

Caprella penantis Leach, 1814 

Introduced 

= Caprella acutifrons Latreille, as used by Edmondson, 1946a: 241; Edmondson & Mansfield, 1948 

This is another cosmopolitan species (Arimoto, 1976) with centuries of ship-borne dispersal obscuring 

any original geographic patterns. Of its occurrence in Hawai‘i, Edmondson & Mansfield (1948) wrote: 

“This species ... seems to be the most abundant of the Caprellidae to be found in the shoal waters of 

Hawaii. It is typically associated with the hydroid Pennaria ... Large numbers ... have been taken in 

Kaneohe Bay, Kahana Bay, and Honolulu Harbor, where colonies of the hydroid grow luxuriantly. This 

caprellid is a voracious species, a dozen specimens being capable of completely stripping a large 

colony of Pennaria of its polyps in a few hours.” MacKay (1945) reported large numbers in Honolulu 

Harbor also eating the polyps of Pennaria, based on observations made in 1941–1942. 

Bishop Museum material dates from 1921 and later years. Schellenberg (1938) reported it from 

Kāne‘ohe in 1927 as “Seichtwaßer, sehr zahlreich” (shallow water, very numerous). We regard this 

species as a ship introduction. 

Caprella equilibra Say, 1818 

Introduced 

This now-cosmopolitan caprellid was first recorded from Hawai‘i by Edmondson & Mansfield (1948) 

based upon material (in the Bishop Museum collections) collected in 1944 by Edmondson from algae 

on a submerged buoy 1.6 km off the west coast of O‘ahu in 4.6 m of water. It remains present in Ala 

Wai Harbor where specimens (identified by I. Takeuchi, pers. comm. to R.C. DeFelice, May 1999) 

were found to be common in 1998. We regard this species as a ship introduction. 

Caprella danilevskii Czerniavskii, 1868 

Introduced 

Caprella danilevskii is a fourth species of caprellid that has apparently been widely dispersed from 

its original home. Described from the Black Sea (McCain & Steinberg, 1970) it too is now said to 

occur on most coasts of the world. Edmondson & Mansfield (1948) noted it from many shoal-water 

stations around O‘ahu “typically associated with the brown seaweeds Sargassum species. It has also 

been recovered from the tufted colonies of the bryozoan Bugula neritina.” It may have spread from


93 

an initial inoculation in ports and harbors to more open systems, as have other invasions in Hawai‘i. 

Bishop Museum collections contain specimens from O‘ahu collected in 1921 and later years. We 

regard this species as a ship introduction. 

Paracaprella pusilla Mayer, 1890 

Introduced 

This Atlantic caprellid (McCain, 1968) has been reported from several locations in the Pacific theater, 

including Amoy, China (McCain, 1968) and the Hawaiian Islands. It has now recently been 

reported from Chile, a first record for the Pacific coast of South America (Guerra-Garcia & Thiel, 

2001). Edmondson & Mansfield (1948) noted it from Honolulu Harbor in 1937 from the screen of a 

water intake pipe at an electrical company. Additional material came from a hydroid (Pennaria) 

colony in 1941 in Honolulu Harbor. Edmondson & Mansfield (1948) note that it had been previously 

known from the West Indies “and South America.” McCain (1968) notes a wide distribution in the 

western North Atlantic and western South Atlantic Oceans (type locality Rio de Janeiro, Brazil), with 

outlier records in the eastern South Atlantic (tropical West Africa, Congo), South Africa (Durban), 

the Indian Ocean (Tanzania), and the Suez Canal (as well as the Pacific stations noted above). It 

appears to be a strongly Caribbean species. We also regard this species as a ship introduction. 

Gammaridea (gammarid amphipods) 

Pleustidae 

Incisocalliope derzhavini (Gurjanova, 1938) 

Introduced 

= Parapleustes derzhavini 

= (?) Parapleustes derzhavini makiki Barnard, 1970 

This Asian amphipod was first collected in 1967 in Kāne‘ohe Bay in 3 to 4 m of water from washings 

from corals and coralline and “fleshy” algae (Barnard, 1970, 1971). Ishimaru (1984) synony - 

mized Barnard’s subspecies into the stem species. Chapman (1988) concluded that P. derzhavini was 

“likely to have been introduced to Hawaii from Asian and North American coasts with the fouling 

on the hulls of ships and with discharged ballast water... or on logs towed to Hawaii from North 

America.” An alternative interpretation is that of Bousfield & Hendrycks (1995), who elevate 

Barnard’s subspecies to full species level, as Incisocalliope makiki. We tentatively retain Ishimaru’s 

interpretation here, as a worker who was intimately familiar with within-species variation in 

Japanese populations of I. derzhavini. 

Ischyroceridae 

Jassa falcata (Montagu, 1808) 

Introduced 

While Barnard (1970), based upon surveys in 1967, was unable to find Jassa falcata in Hawai‘i (and 

took pains to point out its absence), Coles et al. (1999b) collected this species at Sea Land Pier in 

Honolulu Harbor in 1997. We regard this North Atlantic species as a ship fouling or ballast water 

introduction. 

Ericthonius brasiliensis (Dana, 1853) 

Introduced 

= Ericthonius disjunctus Stout as reported by Edmondson & Ingram (1939) 

This widely-distributed amphipod was first collected in Kāne‘ohe Bay fouling studies in 1935 and 

later years (Edmondson & Ingram, 1939, as E. disjunctus, as identified by Clarence Shoemaker; see 

Barnard, 1955 for synonymy). Barnard (1955) reported it from Pearl Harbor (1938), Waikīkī Beach 

(1943, on the bottom of a small boat), off of Barbers Point (one mile offshore, on iron buoy suspended 

to depth of 9 m), and from Honolulu Harbor (1951). Barnard (1970) noted that it was a strong 

candidate as an introduced species. Barnard (1971) reported that it formed “great masses of silty 

tubes attached to piles and docks in harbors in Oahu”, but that it also occurred “sparingly in the open 

sea to depths of at least 30m” (an error apparently for “30 feet” [9 m] as reported in Barnard, 1955). 

Grovhoug (1979) reported it from Pearl Harbor (based on 1978 collection), where it still occurs 

(Coles et al., 1999a; 1996 collections). The ”great masses of silty tubes” noted by Barnard (1971) 

were not observed in 1996, but there may be seasonal aspects to this species’ abundance. Coles et

94 


al. (1999b) also report it from Honolulu Harbor, Ke‘ehi Lagoon, Ala Wai Harbor, Barbers Point 

Harbor, and Kewalo Basin (1997–1998 collections), and Coles (2002b) report it at Waikīkī (2001). 

Coles et al. (2006) report it from Kaua‘i, Moloka‘i, and the island of Hawai‘i, based upon 2003 collections. 

We tentatively treat it here as a North Atlantic species, where most of the historical records are 

centered. It was introduced in ship fouling or ballast water. 

Ruffojassa ventosa (Barnard, 1962) 

Cryptogenic 

= Eurystheus ventosa, = Ventojassa ventosa 

Barnard (1970, 1971) reports this warm-temperate Californian species from collections made in 

O‘ahu in 1967. Locations included (Barnard, 1970) the “open sea off the west end of Pearl Harbor, 

4–5 m, sand bottom, wash of giant encrustation mass from hard sand reef;” off “Ewa Beach, in 18 

and 30 m, wash of Pocillopora, other corals, and bryozoans”; “off the west end of Pearl Harbor, 

25–30 m, several giant Pocillopora heads and large masses of short-tufted red algae, calcareous 

encrustations”, and “seaward of Moku Manu, windward Oahu, 33 m, stem of black coral Antipathes 

irregularis Verrill, including bivalve community attached to coral.” Coles et al. (2002b) report it at 

Waikīkī based on 2002 collections. While these habitats are not typically those of ship-mediated 

invaders, the disjunct distribution, and the frequency of other known invasions in open marine 

waters in Hawai‘i, suggests that this species is cryptogenic in the Hawaiian fauna and may have been 

transported by ships to the islands. 

Ruffojassa angularis (Shoemaker, 1942) 

Cryptogenic 

= Parajassa angularis 

Barnard (1970, pp. 202–204) noted that “This species and Ventojassa ventosa were not suspected to 

be present in the Hawaiian fauna prior to this study. They are enigmatic ... it is of considerable interest 

that together they have crossed the wide ocean gap between North America and Hawai‘i.” It is, 

like R. ventosa, otherwise known from warm-temperate waters of the Pacific coast (California and 

Mexico). Specimens were collected in 1967 at two of the same sites as noted above for R. ventosa, 

off the west end of Pearl Harbor. 

Isaeidae 

Photis hawaiensis Barnard, 1955 

Cryptogenic 

This mysterious amphipod has been known only from O‘ahu since its first collections in 1936 in 

Kāne‘ohe Bay (Barnard, 1955). It was reported from Pearl Harbor based on 1978 collections by 

Grovhoug (1979). Barnard (1970, 1971) added no further new data, and the species remains 

unknown from elsewhere. Coles et al. (1997) report it again from Pearl Harbor (1996 collections). 

It remains present in Kāne‘ohe Bay (2000 collections; Coles et al., 2002a) and occurs in other O‘ahu 

harbors (1997 and 1998 collections: Honolulu Harbor, Ke‘ehi Lagoon, Ala Wai Harbor, Barbers 

Point Harbor, and Kewalo Basin). Coles et al. (2006) report it from Maui (2003 collections); additional 

records are from harbors at Nawiliwili and Port Allen, Kaua‘i; Kahului and Mā‘laea on Maui; 

and Hilo Harbor, also based on 2003 samples (Coles et al., 2004). 

Photis hawaiensis may represent an introduction first described from the site of invasion. We 

predict that it will be found to be native elsewhere, perhaps having been described under another, 

earlier name. 

Corophiidae 

Monocorophium acherusicum (Costa, 1857) 

Introduced 

= Corophium acherusicum 

This North Atlantic amphipod was mentioned by Shoemaker (1947) from “Oahu” (based on 

Smithsonian Institution specimens, but without any further data). Barnard (1955, 1971) recorded it 

from the bottom of a small boat at Waikīkī Beach (1943 record). It was collected in Pearl Harbor in 

1996 (Coles et al., 1997, 1999a), and in Ke‘ehi Lagoon, Ala Wai Harbor, and Kewalo Basin in 

1997–1998 (Coles et al., 1999b). We consider it introduced (as did Barnard) in ship fouling or ballast 

water.


95 

Monocorophium insidiosum Crawford, 1937 

Introduced 

= Corophium insidiosum 

This North Atlantic amphipod was first reported from the Hawaiian Islands by Barnard (1970), who 

noted specimens from Hilo collected in 1959. Barnard (1971) noted that Hilo specimens were “highly 

aberrant” morphologically, suggesting that the differences in Hawaiian material “may represent 

the loss of phenotypic stability as the result of a ‘founder effect’.” It was collected in Pearl Harbor 

in 1996 (Coles et al., 1997, 1999a). Janet Lambertson (pers. comm., 1999) reports that in 1992 C. 

insidiosum (identified by Faith Cole) was abundant on O‘ahu in Ka‘elepulu Stream, Kailua, and also 

occurred at Kāne‘ohe Beach Park, Kāne‘ohe. We consider it introduced (as did Barnard) in ship fouling 

or ballast water. 

Laticorophium baconi (Shoemaker, 1934) 

Introduced 

= Corophium baconi 

Barnard (1970, 1971) reported this Pacific American species questionably from Kāne‘ohe Bay based 

upon collections in 1967. Grovhoug & Rastetter (1980) recorded it from Pearl Harbor and Kāne‘ohe 

Bay (1976–1977 collections) and Grovhoug (1979) reported it from Pearl Harbor (1978 collections), 

where it still occurs (1996 collections; Coles et al., 1997, 1999a). Coles et al. (1999b) report it from 

Honolulu Harbor, Ke‘ehi Lagoon, Ala Wai Harbor, and Kewalo Basin based upon 1997–1998 collections. 

It was found in 2002 collections at Port Allen, Kaua‘i (Coles et al., 2004). Barnard discussed 

and listed (1970, Table 4, column 1) this species as a probable introduction, with which we 

concur. It was introduced in ship fouling or ballast water. 

Aoridae 

Grandidierella bispinosa Schellenberg, 1938 

Introduced 

Muir (1997) reported this Indo-Pacific species from Pearl Harbor, based on 1996 collections (Coles 

et al., 1997). It was found at a brackish water location on the north shores of the East Loch, Pearl 

Harbor. It was previously known from the Bismarck Archipelago, Fiji, and the Marianas Islands. 

Myers (1985) described its habitat in Fiji as being “in mangrove litter and among phanerograms 

(Syringodium, Halophila).” Coles et al. (1999a) listed it as cryptogenic. Given its absence in earlier 

collections from estuaries and harbors on O‘ahu and its location in Pearl Harbor, we regard it as 

introduced, in ship fouling or ballast water. 

Grandidierella japonica Stephensen, 1938 

Introduced 

= (?) Neomicrodeutopes makena Barnard, 1979 (see below) 

Muir (1997) reported this brackish water Japanese species as a “new state record” from brackish 

water sites from throughout Pearl Harbor based on 1996 collections. Janet Lambertson (pers. comm., 

1999) reports that in 1992 Grandidierella japonica (identified by Faith Cole) was collected on O‘ahu 

in abundance from Kāne‘ohe Beach Park, Kāne‘ohe (near Waikalua fishpond, at the mouth of 

Kāne‘ohe Stream) and from Ka‘elepulu Stream, Kailua. The population was still “thriving” in 1993 

and 1994. It has also been introduced to the west coast of North America (Chapman & Dorman, 

1975). Chapman & Dorman (1975) suggested that Grandidierella makena (Barnard, 1970) (= Neo - 

microdeutopes makena), described from a high energy, high surf location at Makapu‘u Point, on 

O‘ahu (based on 1967 collections) might be a junior synonym of G. japonica. Muir (1997) felt that 

the two species were distinct; the matter requires reexamination (J. Chapman, pers. comm., 1999). 

It was introduced in ship fouling or ballast water. 

Leucothoidae 

Paraleucothoe sp. 

Introduced 

Specimens of this sponge- or ascidian-associated gammarid were collected in 1996 in Pearl Harbor 

(Muir, 1997) in fouling communities (Coles et al., 1997). Coles et al. (2006) report it from Kaua‘i 

and Moloka‘i, based upon 2003 collections. It is similar but not identical to various Paraleucothoe 

species, and in particular to P. fllindersi Stebbing, 1888 (Muir, 1997), known only from a single

96 


specimen collected in Flinder’s Passage, Australia. The Paraleucothoe in Pearl Harbor thus remains 

unidentified and perhaps undescribed. It was listed as P. ?flindersi in Coles et al. (1999a) as a cryptogenic 

species, and as P. findersi in Coles et al. (2004), where it was reported in every harbor sampled 

on Kaua‘i, Moloka‘i, Maui, and Hawai‘i. We concur with Muir (1997) that it is introduced. We 

tentatively treat it here as an Indo-Pacific species, arriving in ship fouling or ballast water. 

Leucothoe micronesiae Barnard, 1965 

Introduced 

Coles et al. (1999b) record this species from Honolulu Harbor based upon 1997 collections. The type 

locality of this species is Ifaluk Atoll in the Caroline Islands (Barnard, 1965). Ledoyer (1986) subsequently 

recorded it from Madagascar. We regard it as introduced. 

Melitidae 

Elasmopus “rapax” Costa, 1853 

Introduced 

Barnard (1970) discussed this species as “… undoubtedly a strong candidate... as an import by artificial 

means.” Barnard (1955, 1970) reported material from Kāne‘ohe Bay (1937), Pearl Harbor 

[1948 (C.H. Edmondson) and 1951], and the Wai‘anae shore (1952). Barnard’s (1955) record of it 

from the Honolulu Aquarium is Elasmopus calliactis Edmondson, 1951; see Barnard, 1970, p. 131). 

Grovhoug (1979) also reported it from Pearl Harbor. Coles et al. (1997, 1999a) further record it from 

Pearl Harbor (1996 collections); Coles et al. (1999b), based upon 1998 material, report it from Ala 

Wai Harbor and Kewalo Basin. Coles et al. (2006) report it from reefs and Coles et al. (2004) from 

harbors on Kaua‘i, Moloka‘i, Maui, and the island of Hawai‘i based upon 2002–2003 collections. 

Elasmopus “rapax” was introduced in ship fouling or ballast water. We place the species name 

in quotation marks as this taxon may represent a worldwide complex of closely related species. We 

tentatively treat it here as a North Atlantic species, where most of the historical records are centered. 

Elasmopus pectenicrus (Bate, 1862) 

Cryptogenic 

Barnard (1955) first reported this “tropicopolitan” (Barnard, 1970) species from Hawai‘i, in and near 

Pearl Harbor in fouling communities, based upon material collected between 1937 and 1950, to 

depths of 18 m. It had been reported from the Caribbean Sea, Red Sea, Indian Ocean, and tropical 

Pacific Ocean (Barnard, 1955). Coles et al. (1999b) report it from Honolulu Harbor, Ala Wai Harbor, 

and Kewalo Basin. Its apparent restriction to harbor locations on O‘ahu (Barnard, 1970, 1971), combined 

with the probability that a global species complex is involved, lead us to consider it cryptogenic. 

Podoceridae 

Podocerus brasiliensis (Dana,1853) 

Introduced 

Barnard (1955) first reported P. brasiliensis from Kāne‘ohe Bay (1935), Pearl Harbor (1937) and 

Hanauma Bay (1937); Barnard (1970) suggested that it was a “strong candidate to be an introduced 

species”; Barnard (1971) noted that it occurred in Hawai‘i “mainly as fouling organism in harbors.” 

Grovhoug (1979, 1978 collections) reported it from Pearl Harbor as well, where it still occurs (Coles 

et al., 1997, 1999a; 1996 collections). Coles et al. (1999b) report it from Ala Wai Harbor and Kewalo 

Basin based on 1998 material; Coles et al. (2002b) report it at Waikīkī (2001specimens), and Coles 

et al. (2004) report it in Kahului Harbor, Maui (2003 collections). 

We tentatively treat it here as a North Atlantic species, where most of the historical records are 

centered. It was introduced in ship fouling or ballast water. 

Stenothoidae 

Stenothoe gallensis Walker, 1904 

Introduced 

= Stenothoe valida of Schellenberg, 1938 

Schellenberg (1938) first reported this species from Kāne‘ohe Bay, O‘ahu (“Seichtwaßer, mehrere 

females ovig. 4 mm. O‘ahu, Riff, Mitte Kāne‘ohe Bay...”, [Shallow water, several females, oviger-


97 

ous, 4 mm, O‘ahu, Reef, Middle Kāne‘ohe Bay]). Barnard (1955) listed it from Kāne‘ohe Bay 

(1935–1937 material) and Pearl Harbor (buoy, 1944; dry dock, 1948, 1950). Barnard (1970) discussed 

the possibility of its introduction, and taxonomic challenges. Barnard (1971) noted that in 

O‘ahu it was primarily a “fouling organism in harbors probably associated with hydroids.” Coles et 

al. (1997, 1999a) report it from Pearl Harbor based on 1996 collections. Coles et al. (2002a) report 

it from Kāne‘ohe Bay (2000 collections) and from Hale O Lono and Kaunakakai Harbors on Molo - 

ka‘i. 

Stenothoe gallensis is known from warm waters worldwide [for example, Caribbean, Medi - 

terranean, Sri Lanka, and in French Polynesia in the Gambier Archipelago on the island of Man - 

gareva (Barnard, 1955)]. Its origin is unknown. We regard it as a ship fouling or ballast water introduction. 

Stenothoe valida Dana, 1853 

Introduced 

Barnard (1970) reported this species as a questionable identification, noting its possible introduction 

as a typical harbor species. However, the specimens Barnard had in hand were collected in 1967 by 

D.M. Devaney from seaward of Moku Manu, O‘ahu, on the stems of black coral (and possibly from 

the bivalve community attached to the coral), causing Barnard to make the following remarks: “Socalled 

S. valida has been found in Hawai‘i only in association with the black-coral Antipathes irregularis 

off the east coast of Oahu in 33 m of water depth. Under normal circumstances that unusual 

habitat would suggest a misidentification of S. valida, for the species normally inhabits harbors... the 

Oahuan population of Antipathes is composed of colonies moved by biologists to that locale from 

other parts of Hawaii. One might thus expect that a eurytopic harbor fouling organism could find an 

open niche in such an artificial situation. But S. valida has not as yet been collected in Hawaiian harbors.” 

Barnard (1971) repeated this record. Chilton (1924) earlier noted that he had “recently 

received specimens from the Hawaiian Islands which appear to belong to this species.” 

We tentatively admit Chilton’s record as the first for the islands, but note that the next record 

(other than the 1967 material, above) appears to be that of collections made in 1996 in Pearl Harbor 

(Coles et al., 1999a). The 1978 record for Pearl Harbor noted by Coles et al. (1999a) is based on a 

report by Grovhoug (1979) who questionably identified his material as either S. gallensis or S. valida. 

Coles et al. (1997, 1999a) report it from Pearl Harbor based on 1996 collections; Coles et al. 

(1999b) report it from Honolulu Harbor, Ala Wai Harbor, Barbers Point Harbor, and Kewalo Basin 

(1997–1998 collections); Coles et al. (2002a) further report it in Kāne‘ohe Bay (2000). Coles et al. 

(2006) report it from Kaua‘i, based upon 2002 collections; Coles et al. (2004) further record in harbors 

on Moloka‘i and Maui, from 2003 collections. 

Stenothoe valida is now reported from many areas of the world; we tentatively regard it as a 

North Atlantic species. 

Cheluridae 

Tropichelura insulae (Calman, 1910) 

Introduced 

Miller (1924b) reported this wood-boring amphipod, originally described from Christmas Island in 

the Indian Ocean, from wooden test blocks in Samoa and in Honolulu Harbor, based upon collections 

made in 1922. Barnard (1955) recovered specimens in 1949 from Honolulu Harbor as well, and 

noted further records from the Caroline and Mariana Islands. Cookson (1991) reported the first 

record from Australia (Green Island, associated with Limnoria). While Barnard (1970, p. 17) stated 

that “The question as to whether this species has been distributed around the world by modern shipping 

is entirely open to question,” he listed it in his Table 4 as a possible candidate for human importation. 

Barnard (1971) noted it again from Honolulu Harbor, on pilings and floating docks. 

We believe this species is a classic case of an introduction in the days of wooden-hulled ships. 

We regard it as introduced from the Indo-West Pacific.

98 


Talitridae 

Platorchestia platensis (Kroyer, 1845) 

Cryptogenic 

= Orchestia platensis 

This now globally occurring sand hopper or beach hopper has been reported from the Hawaiian 

Islands for many years over a wide variety of habitats and elevations. Chilton (1922) stated that 

“... in the Hawaiian islands Orchestia platensis is found from the seashore up to very considerable 

heights, and numerous other examples of the same kind could be given.” Subsequent reports are 

given at http://www2.bishopmuseum.org/HBS/checklist/citation.asp?grp=Arthropod, including 

records from the islands of Kaua‘i, O‘ahu, Lāna‘i, Maui, and Hawai‘i. While Fee (1967) studied 

beach specimens from Kāne‘ohe Bay, the identity of his material was uncertain according to that 

author. Alicata (1936) reported P. platensis (as O. platensis) from inland poultry farms, “especially 

in damp and wet areas around water fountains”, based upon material identified at the time by 

M.A. Miller. Earlier records of P. platensis from high elevations in Hawai‘i (from the islands of 

Hawai‘i, O‘ahu, and Lāna‘i) by Stebbing (1900) have been assigned to Platorchestia pickeringi 

by Barnard, 1955 (as Orchestia pickeringi). Barnard (1971, p. 131) referred to the “Hawaiian 

beach-hopper Orchestia platensis” as making excellent introductory laboratory specimens for a 

study of amphipods. 

If Hawaiian populations include true P. platensis, we admit here as cryptogenic populations the 

strictly littoral, marine, beach-wrack associated populations that may have been introduced with 

early ship’s ballast. 

Tanaidacea (tanaids) and Isopoda (isopods) 

Introduced and Cryptogenic Tanaidacea and Isopoda 



Tanaidacea 

Parapseudes pedispinis 1996 SF,BW Northeast Pacific 

Apseudes sp. 1996 SF,BW Japan? 

Isopoda 

Gnorimosphaeroma rayi 1972 SF,BW Japan 

Sphaeroma walkeri 1943 SF,BW Indian Ocean 

Paradella dianae 2002 SF, BW Eastern Pacific 

Pistorius bidens 2002 SF, BW Australia 

Paracerceis sculpta 1943 SF,BW Northeast Pacific 

Exosphaeroma sp. 1996 SF,BW Unknown 

Mesanthura sp. 1996 SF,BW Unknown 

Limnoria tripunctata 1922 SB Southern hemisphere? 

Paralimnoria andrewsi 1922 SB Indo-West Pacific 

Caecijaera horvathi 1975 SB Southern hemisphere? 

Littorophiloscia culebrae 1984 SBA North Atlantic? 

Halophiloscia couchii 1997 SBA Northeast Atlantic 

Armadilloniscus ellipticus 1985 SBA Unknown 

Alloniscus oahuensis 1879 SBA Indo-Pacific 

Porcellio lamellatus 1973 SBA Northeast Atlantic 

Buchnerillo sp. 1985 SBA Atlantic? 

Olibrinus truncatus 1985 SBA Southern hemisphere? 

Ligia exotica 1996 SF, SBA Unknown 


Tanaidacea 

Leptochelia “dubia” 1932 SF,BW


99 



Isopoda 

Sphaeroma quoianum ca. 1920 SF/BW New Zealand 


Sphaeroma terebrans 

Status: Erroneous record; see discussion 

Tanaidacea (tanaids) 

Parapseudididae 

Parapseudes pedispinis Boone, 1923 

Introduced 

This Eastern Pacific tanaid, previously known from California to Ecuador, was reported by Muir 

(1997) from Pearl Harbor, based on 1996 collections (Coles et al., 1997). Coles et al. (1999b) report 

it also from Honolulu Harbor (1997 collections). Coles et al. (2006) report it from Moloka‘i, based 

upon 2003 collections. Muir regarded it as an introduced species, and we concur. It was introduced 

in ship fouling or with ballast water. 

Apseudididae 

Apseudes sp. 

Introduced 

This tanaid is widely distributed in Kāne‘ohe Bay (on the Kāne‘ohe Bay Sand Bar) and in Pearl 

Harbor (1996 collections) (Muir, 1997; Coles et al., 1997). Muir (1997) found it to be most closely 

related to the Japanese Apseudes nipponicus Shiino, 1937, but felt that differences between Ha - 

waiian material and A. nipponicus warrant the description of the Kāne‘ohe Bay—Pearl Harbor specimens 

as a new species. Given its broad distribution on both the north and south sides of O‘ahu, it 

is not likely to have been missed in earlier surveys, including the World War II buoy surveys (Miller, 

1940, 1968). We regard it as introduced, regardless of whether it is described or not. 

Anthruridae 

Leptochelia “dubia” (Krøyer, 1842) 

Cryptogenic 

Miller (1940) reported this tanaid, which has a purported global distribution, from various stations 

on O‘ahu and Maui, with the earliest collection being at Waikīkī by C.H. Edmondson in 

1932. Additional collections included Kawela Bay, Kāne‘ohe Bay, Hanauma Bay, and Hālona 

on O‘ahu, and Hawea Point, Hāna, and Hanamanioa on Maui. Grovhoug (1979) reported it (as 

a questionable identification) from Pearl Harbor in 1978. Bailey-Brock (1984) found it in densities 

up to 32,800 per square meter in the sand beds of the onuphid worm Diopatra leuckarti 

at Niu Valley on the south shore of O‘ahu. Bishop Museum collections also include specimens 

collected in 1938 at Black Point, O‘ahu. 

Coles et al. (1997, 1999a) record it from Pearl Harbor (1996 collections); Coles et al. 

(1999b, based on 1997–1998 collections) report it from Honolulu Harbor, Ke‘ehi Lagoon, Ala 

Wai Harbor, Barbers Point Harbor, and Kewalo Basin; Coles et al. (2002a) report it from 

Kāne‘ohe Bay (2000), and Coles et al. (2002b) report it from Waikīkī (2001). Coles et al. (2004, 

2006) report it from reefs and harbors on Kaua‘i, Moloka‘i, Maui, and the island of Hawai‘i, 

based upon 2002–2003 collections. 

The systematics of this species remain to be elucidated; compounding the problem is that it is 

probable that both a species complex is involved and that one or more members of this species group 

have been transported in ship fouling, with oysters and by other means. We regard it as cryptogenic 

until Hawaiian material has been reexamined and compared with other subtropical populations. 

Isopoda (isopods) 

Sphaeromatidae 

Gnorimosphaeroma rayi Hoestlandt, 1969 

Introduced 

Hoestlandt (1973, 1975) reported this Japanese isopod from the Wai‘anapanapa Caves on Maui 

Island; the specimens were collected in 1972 (NMNH collections). Subsequently, Hoestlandt identified 

additional specimens of G. rayi from Waiahuakua Stream (Kaua‘i) and from Lua o Palahemo

100 


pond and Waiahukini (Hawai‘i) (H. Hoestlandt to D.M. Devaney, in litt., August 3, 1976). Collection 

dates are 1974–1975 for the last three sites (BPBM collections). Holthuis (1973, p. 11) noted this 

species as unidentified isopods from Lua o Palahemo pond, with a salinity of 22 ‰, an usually low 

salinity for this more marine isopod. 

Although described originally from California, Hoestlandt later found it in the Sea of Japan and 

believed it be introduced to California from there with oyster culture. We regard it as introduced to 

Hawai‘i as well, perhaps in ship fouling or in ballast water. 

Paradella dianae (Menzies, 1962) 

Introduced 

Coles et al. (2004) report this well known and widespread, often harbor-dwelling isopod from fouling 

communities from Kaua‘i in 2002 and from Maui in 2003. Now reported from harbors around 

the world, it is native to the warm waters of the Eastern Pacific, where it is a member of a small 

group of Paradella with distinctively sculptured pleons and pleotelsons (Wetzer & Bruce, 2007) 

Pistorius bidens Harrison & Holdich, 1982 

Introduced 

Coles et al. (2004) report this Australian isopod from fouling communities in Kaua‘i in 2002, 

Moloka‘i in 2003, and Maui in 2003. 

Sphaeroma walkeri (Stebbing, 1905) 

Introduced 

This well-known ship-dispersed isopod (Carlton & Iverson, 1981) was first collected in the Ha - 

waiian Islands in 1943 in Hilo (Miller, 1968) and again in 1961 at Hanamā‘ulu Bay, Kaua‘i. Bishop 

Museum material also includes collections from Pearl Harbor in 1976. Carlton & Iverson (1981) 

review its global dispersal history. We regard it as introduced. That it is uncommon is revealed by 

the remarks of Edmondson (1955) who, unaware of the 1943 collections [which were not to be published 

until 1968 (Miller, 1968)], and a long observer of the boring and fouling fauna of the Islands, 

noted that “[Sphaeroma], in so far as I know, has not been observed in Hawaiian waters.” It was also 

noted by Grovhoug & Rastetter (1980) for Kāne‘ohe Bay and Pearl Harbor (1976–1977 collections). 

Sphaeroma quoianum Milne-Edwards, 1840 

Deleted 

= Sphaeroma quoyanum, incorrect subsequent spelling 

= Sphaeroma pentodon Richardson, 1904 

A specimen of this widespread isopod is in the Smithsonian Institution collections, collected by Paul 

Bartsch in Pearl Harbor, probably in 1920 (see Callinectes sapidus). There are no further records. 

Sphaeroma terebrans Bate, 1866 


O’Neill (1983) and Pendleton & O’Neill (1986) state that “The principal borers recorded from Pearl 

Harbor are the crustaceans, Limnoria tripunctata and Sphaeroma terebransi [sic], and the mollusks...” 

While the Indian and Atlantic Ocean isopod S. terebrans would not be unexpected in the Hawaiian 

Islands, there are no other records of this species from Hawai‘i, and no indication of the source of these 

statements. In their own studies, O’Neill & Pendleton (1968) do not record any species of Sphaeroma 

from Pearl Harbor. We consider this report a lapsus and do not consider it further here. 

Paracerceis sculpta (Holmes, 1909) 

Introduced 

Miller (1968) reported this eastern Pacific species from Pearl Harbor and Hilo Harbor (Hawai‘i) 

based upon material collected during World War II (1943), noting that “probably it was transported 

on the hulls of naval ships plying between San Diego and Hilo and Pearl Harbor.” Grovhoug (1979) 

reported it again from Pearl Harbor based upon specimens collected in 1978. It was also noted by 

Grovhoug & Rastetter (1980) for Kāne‘ohe Bay and Pearl Harbor (1976–1977 collections), by Coles 

et al. (2002a) in Kāne‘ohe Bay, and by Coles et al. (2004) in Port Allen Harbor, Kaua‘i.


101 

Exosphaeroma sp. 

Introduced 

Coles et al. (1997, 1999a) report an Exosphaeroma sp. from Pearl Harbor, based on 1996 collections. 

Although not yet identified to species, we regard this isopod as introduced, as no other sphaeromatids 

of this or any related genera (such as Paracerceis or Dynamenella) are known to exist naturally 

in the Islands. We regard it as introduced with ship hull fouling or in ballast water. 

Anthuridae 

Mesanthura sp. 

Introduced 

This isopod, previously unknown from the islands, was collected in Pearl Harbor surveys of 1996 

(Coles et al., 1999a). Coles et al. (1999b) record it from Honolulu Harbor, Ke‘ehi Lagoon, Barbers 

Point Harbor, and Kewalo Basin; further stations include Kāne‘ohe Bay (Coles et al., 2002a) and 

Waikīkī (Coles et al., 2002b). As with the introduced tanaid Apseudes sp., given the broad distribution 

of this isopod, it is not likely to have been missed in earlier surveys, including the World War II 

buoy surveys (Miller, 1940, 1968). 

Limnoriidae 

Limnoria tripunctata Menzies, 1957 

Introduced 

= Limnoria lignorum of pre-1957 Hawaiian authors 

This wood-boring gribble was collected at least as early as 1922 in test blocks in Pearl and Honolulu 

Harbors and at Nawiliwili Bay on Kaua‘i (Miller, 1924b, as Limnoria lignorum). Numerous specimens 

are in Bishop Museum collections from Honolulu Harbor collected between 1945 and 1949 

and identified by R.J. Menzies. While it is curiously absent from Menzies’ 1957 monograph, L. 

tripunctata is marked on map 3 of Menzies’ 1959 paper (p. 33c) as being on the Hawaiian islands. 

Cooke (1977b) found it in 1975 at Makai Range, Makapu‘u, O‘ahu heavily infesting Douglas fir test 

blocks at a depth of 1 m after a submergence period of 6 months. Coles et al. (1999a) continue to 

report its presence in Pearl Harbor (1996 collections). 

Paralimnoria andrewsi (Calman, 1910) 

Introduced 

Miller (1924b, as Limnoria andrewsi) reported this wood-boring gribble, originally described from 

Christmas Island, Indian Ocean, from Honolulu Harbor and Nawiliwili (Kaua‘i), as well as Samoa, 

based on test blocks suspended in 1922. Paralimnoria andrewsi occurred in Honolulu Harbor with 

Limnoria tripunctata, but greatly outnumbered this latter species; in turn, Limnoria tripunctata “predominates 

to such an extent [in Pearl Harbor] that if [P.] andrewsi occurs [there] at all its presence 

has been overlooked.” This interesting observation in two harbors only a few kilometers apart may 

indicate competition between the two species. Bishop Museum collections include specimens collected 

in Honolulu Harbor in 1949. 

Paralimnoria andrewsi occurs in the Indian Ocean (Christmas Island, its type locality, above, 

and the Cocos Islands), in the South Pacific Ocean (Samoa, Philippines, Papua New Guinea), in the 

North Pacific Ocean (Japan and Hawaiian Islands), and in the Atlantic Ocean (Florida, Caribbean 

[Puerto Rico], Ghana) (Cookson, 1991). We regard it as introduced to the Atlantic Ocean, to Hawai‘i 

and perhaps to Japan as well. 

Janiridae 

Caecijaera horvathi Menzies, 1951 

Introduced 

Cooke (1977b) reported this tiny, eyeless asellote isopod from the tunnels of the wood boring gribble 

Limnoria tripunctata in test blocks of Douglas fir below the pier at a depth of 1 m at Makai Range, 

Makapu‘u, O‘ahu, in March 1975. Carlton (1979a) considered it introduced with Limnoria to Los 

Angeles—Long Beach Harbors (its type locality) as well. Cooke (1977b) suggested that it could be 

introduced. We consider it introduced, with ship boring communities, possibly in the hulls of yachts, 

to the Hawaiian Islands. Its origin remains unknown, but in concert with the general nature of the 

wood-boring warm-water biota, it may be native to the southern hemisphere.

102 


Oniscidea 

We include in our treatment here eight species of maritime isopods that are intimately associated 

with salt water habitats. 

Philosciidae 

Littorophiloscia culebrae (Moore, 1901) 

Introduced 

= Philoscia culebrae; = Vandeloscia culebrae 

Taiti & Howarth (1995) describe this species as, “A strictly halophilic littoral species found under 

debris, logs, and stones on both rocky and sandy shores.” It was first found (Taiti & Ferrara, 1986) 

in O‘ahu in 1984 (Ala Moana Park beach, under coconuts, S. Taiti collector). It was previously 

known from Florida, Puerto Rico, Cuba, Virgin Islands, Angola, and Madagascar (Taiti & Ferrara, 

1986; Taiti & Howarth, 1996), and thus appears to be a species of largely Atlantic origin. Taiti & 

Howarth, (1996) added new records from Pearl and Hermes Atoll (1983) and Lisianski (under rocks 

on beach, 1983). Stefano Taiti regards this species as introduced (pers. comm., August 1996), possibly 

with early ship ballast. Taiti (1999) extended the range to Midway Atoll (1997 collections, under 

splash zone rocks on jetty). 

Halophilosciidae 

Halophiloscia couchii (Kinahan, 1858) 

Introduced 

Taiti (1999) reported the first record of this maritime isopod from the Hawaiian archipelago, based 

upon specimens collected in Midway in 1997. He notes that “this halophilic species occurs on coasts 

of marine and brackish waters.” Its native range appears to be the warm eastern Atlantic Ocean; Taiti 

(1999) notes that “it has also been introduced to Virginia, Bermuda, Argentina, and Western Aus - 

tralia.” 

Scyphacidae 

Armadilloniscus ellipticus (Harger, 1878) 

Introduced 

= Armadilloniscus litoralis Budde-Lund, 1885 (fide Garthwaite et al., 1992) 

Taiti & Ferrara (1989, as A. litoralis) record this species from Coconut Island, O‘ahu, based upon 

specimens collected by S. Taiti in 1985; they note that “... this species has a very wide distribution, 

which is certainly due to the ease with which this halophilic form is passively transported.” Its origin 

is not clear, as it is known from the Atlantic coast of North America (Massachusetts to Florida; 

the type locality of A. ellipticus is in Connecticut), Bermuda, the Azores, Madeira, and Medi ter - 

ranean (the type locality of A. litoralis is the Venice Lagoon, Italy), in the Indian Ocean from 

Madagascar, and in the Pacific Ocean from Tahiti, Malaysia (Pulau Pinang), North Korea, Hong 

Kong, Japan, and Hawai‘i. Genetic analyses should aid in determining its geographic origin. Schultz 

(1972) provided a detailed habitat description for Bermuda; he notes that this species “at times of 

high tide (is) many times completely under water.” Stefano Taiti regards this species as introduced 

(pers. comm., August 1996), possibly with early ship ballast. Taiti (1999) extends the range of this 

isopod in the Archipelago to Midway Atoll (1997 collections, under splash zone rocks on jetty). 

Scyphacidae 

Alloniscus oahuensis Budde-Lund, 1885 

Introduced 

This species was recorded from O‘ahu by Budde-Lund in 1879 and then formally described in 1885. 

Taiti & Howarth (1996) note that A. oahuensis “is a strictly littoral species that occurs under logs or 

litter in the upper part of sandy shores.” It is now recognized as widely distributed in the Indian and 

Pacific Oceans; S. Taiti regards this as an introduced species in the Hawaiian Islands (pers. comm. 

August 1996). Schultz (1984) provided a detailed redescription of the species based upon material 

from Hālawa Valley, Moloka‘i Island, noting that it is a species found “on and near marine beaches


103 

and along the margin of rivers which empty into the sea.” Schultz also records specimens from 

Kaua‘i. Taiti & Ferrara (1991) record it from a number of stations in 1984–1985 around O‘ahu. Taiti 

& Howarth (1996) add further Hawaiian records from O‘ahu (Kāne‘ohe Bay, near Kahalu‘u mud 

flats above high water (1973); Kawainui Marsh, litter adjacent to marsh (1980); Popoi‘a Islet, debris 

(1973); Moloka‘i: 1.6 km west of Kamehameha, Coconut Grove, near mauka edge of mangrove 

(1974). Taiti et al. (1992) also record it from the Togian Islands, Indonesia (1987). Taiti & Ferrara 

(1991) discuss the status of an older record from an anomalous habitat from Lāna‘i noted by Schultz 

(1984) and conclude that this material would require reexamination. It was likely carried to the 

Islands with ship ballast. 

Porcellionidae 

Porcellio lamellatus lamellatus Budde-Lund, 1885 Introduced 

Schultz (1972) noted that this species in Bermuda “lives in sandy soil and is also present on the upper 

parts of the beach in the transition region to the large vegetation zone .... It was especially abundant 

in grass patches on the side of a steep cliff-like slope ...where it was living with Armadillidium vulgare.” 

Taiti & Ferrara (1991) recorded it from Turtle Bay, O‘ahu, based upon specimens collected in 

1984. Taiti & Howarth (1996) further recorded it from French Frigate Shoals (1984, under debris), 

Kaua‘i (1973, leaf litter on beach), and Moloka‘i (1974, near mauka edge of mangrove). This species 

is native to eastern and southern Europe and northern Africa, and is also now known from Bermuda, 

Cuba, Argentina, western Australia (Rottnest Island) (Schultz, 1972; Taiti & Ferrara, 1991, 1996). 

Stefano Taiti regards this species as introduced (pers. comm., August 1996). 

Family Uncertain [fide Taiti & Howarth, 1996 and S. Taiti (pers. comm., May 2001)]. 

Buchnerillo sp. 

Introduced 

Taiti & Ferrara (1991) reported this littoral species from Coconut Island, O‘ahu, based upon specimens 

collected in 1985. They noted that it appeared similar to B. oceanicus Ferrara from Somalia. 

Only one other species is known, B. litoralis Verhoeff, from the Mediterranean, Madeira, and Flor - 

ida. Stefano Taiti regards this species as introduced (S. Taiti, pers. comm., August 1996). 

Olibrinidae 

Olibrinus truncatus Taiti & Ferrara, 1991 

Introduced 

Taiti & Ferrara (1991) described this maritime isopod as a new species from Coconut Island, O‘ahu 

(collected in 1985). Stefano Taiti regards it as introduced to the Hawaiian Islands (S. Taiti, pers. 

comm., August 1996). Taiti et al. (1992) recorded it from Pulau in the Togian Islands, Sulawesi, 

Indonesia, noting that it “is strictly halophilic and most probably amphibious, since it is common in 

and around mangrove forests under stones and logs which are submerged during the high tide.” They 

further comment that “Its apparently disjunct range is certainly due to the lack of investigations in 

most parts of the Pacific area.” 

Ligiidae 

Ligia exotica Roux, 1828 

Introduced 

A recent (1996) collection of this species at Hilo by R.C. DeFelice represents the first valid record 

of L. exotica from the Hawaiian Islands. The specimens were identified by S. Taiti. Previous records 

of L. exotica from Hawai‘i by Robertson, Edmondson, Van Name, and others, are all based on Ligia 

hawaiiensis, an endemic species (S. Taiti, pers. comm., 1996).

104 


Cumacea (cumaceans) and Mysidacea (mysids) 

Introduced Cumacea and Mysidacea 


Cumacea 

Nannastacus sp. 1996 BW Unknown 

Scherocumella sp.


105 

Edmondson (1948) reported briefly upon his observations of fouling on the hull of the Swedish scientific 

research vessel Albatross, while it was drydocked in Honolulu Harbor on 1 December 1947. 

Edmondson noted that the hull was “neatly and evenly covered by a species of rock barnacle de - 

scribed many years ago from the peninsula of Lower [Baja] California. The nearest the Albatross 

came to the type locality of the barnacle was the Pacific end of the Panama Canal. As large quantities 

of the barnacle were deposited in Honolulu Harbor from the dry dock, it is anticipated that this 

unique form may make its appearance in local waters before very long.” In the Bishop Museum collections 

there are three lots of barnacles collected from the Albatross in December 1947 at Honolulu, 

identified as Balanus tintinnabulum peninsularis?, Lepas anatifera, and Conchoderma auritum. The 

first barnacle has not been found in Hawai‘i. The last two species occur widely on floating materials, 

whales, and boat bottoms around the world. In a letter in Bishop Museum files dated 8 April 

1960, Edmondson indicates that I.E. Cornwall identified the material and that the Albatross had 

come through the Panama Canal on the way to Hawai‘i. Edmondson further notes in this letter that 

“B. tintinnabulum peninsularis Pilsbry... was the principal fouling on the hull of the boat, although 

large numbers of Conchoderma auritum were also present and also what appeared to be Lepas anatifera.” 

Balanus tintinnabulum peninsularis is now considered a full species, Megabalanus peninsularis 

Pilsbry, 1916 (Newman & Ross, 1976; Henry & McLaughlin, 1986). 

Gordon (1970) reviewed the Hawaiian barnacles, reporting Amphibalanus amphitrite and 

Amphibalanus eburneus from wooden floats and styrofoam docks on Coconut Island in Kāne‘ohe 

Bay. Newman (1986) reviewed the history and biogeography of Hawaiian barnacles, including the 

Amphibalanus species discussed here. 

Balanidae 

Amphibalanus amphitrite (Darwin, 1854) 

Introduced 

= Balanus amphitrite; = Balanus amphitrite hawaiiensis Broch, 1922 

This now widespread barnacle of southern hemisphere origin was first collected in 1902 in Honolulu 

Harbor (Henry & McLaughlin, 1975). This material was collected by the R/V Albatross on July 3, 

1902 (USNM catalogue number 32517, http://goode.si.edu/mcs/iz/Query.php [accessed July 2003]). 

The first published record of these specimens is that of Pilsbry (1907, p. 190), who noted that “Some 

small specimens [of Balanus amphitrite] taken from the bottom of a tug at Honolulu are similar 

externally to figure 2e of plate 5 of Darwin’s Monograph, except that the compartments diverge less 

above, the aperture being somewhat smaller and hardly dentate.” 

Bryan (1915, material identified by H.A. Pilsbry) and Broch (1922, as the new subspecies 

hawaiiensis, specimens collected “on broken china” in 1915 in Pearl Harbor) are additional early 

published records. Pilsbry (1928) repeated the W.A. Bryan record (1915, off Honolulu Harbor) and 

further reported material from Pearl City, from the West Lock of Pearl Harbor between Waipahu and 

Ho‘ae‘ae (‘Ewa), very abundant between tides on an Anomia reef in 1913, and from Kualoa, 

Kāne‘ohe Bay, on volcanic rocks in 1920. 

Edmondson (1933, as B. a. hawaiiensis) noted that it was very common in Pearl Harbor on piling 

and shore rocks. Edmondson & Ingram (1939) provide growth data on fouling panels in Kāne‘ohe Bay 

in 1935 and subsequent years. Evans et al. (1972) and Long (1974) recorded it from Pearl Harbor (and 

from nearshore panels off Pearl Harbor). Henry & McLaughlin (1975) note additional specimens collected 

from Pearl Harbor (1931 and later), Honolulu Harbor, and Kāne‘ohe Bay (the latter, 1959). It 

was also noted by Grovhoug & Rastetter (1980) for Kāne‘ohe Bay and Pearl Harbor (1976–1977 collections). 

It is widespread throughout the main Hawaiian Islands, with records from Kaua‘i, O‘ahu, 

Moloka‘i, Maui, Lāna‘i, and Hawai‘i (Coles et al., 1999a, 1999b, 2002a, 2002b, 2004, 2006). 

Utinomi (1960) reviewed some of the early global records of this species. Hoover (1998, 2006) 

presents a color photograph from Coconut Island, O‘ahu. We regard it as introduced by ship fouling.

106 


Amphibalanus eburneus (Gould, 1841) 

Introduced 

= Balanus eburneus 

This western Atlantic Ocean “ivory barnacle” was first collected in 1929 in Pearl Harbor (Henry & 

McLaughlin,1975). Edmondson (1931) wrote that “the barnacle (Balanus eburneus), among which 

the crab [Panopeus pacificus] is found, is a typical species of the east coast of the United States, 

which is suggestive of the view that both the crabs and barnacles may have been transported to 

Hawai‘i through shipments of oysters or on the bottoms of ships.” Edmondson (1933) noted that it 

was in Pearl Harbor, on buoys, floats, and oyster shells. Hutchins (1949, page 67) reported it in fouling 

on five buoys set in Hawaiian waters in 1943, noting that “the presence of B. eburneus is evidently 

the result of its introduction to the islands on ships, since the form is native to the Atlantic 

coast of North America.” Matsui et al. (1964) reported specimens from McCully Bridge, Ala Wai 

Canal, and Mā‘alaea Bay Harbor, Maui, all collected in 1962. Evans et al. (1972) and Long (1974) 

found it on fouling panels in Pearl Harbor and in nearshore waters off Pearl Harbor. It was also noted 

by Grovhoug & Rastetter (1980) for Kāne‘ohe Bay and Pearl Harbor (1976–1977 collections). 

Ball (1950) found the endoparasitic gregarine protozoan Cephaloidophora communis in this 

barnacle, presumably from O‘ahu, in 1949. It was found only in Amphibalanus eburneus and not in 

two other barnacle species (Amphibalanus amphitrite and Balanus sp.) nor in 30 species of 

decapods. 

Amphibalanus reticulatus (Utinomi, 1967) 

Introduced 

= Balanus reticulatus 

This ship-dispersed barnacle, likely originating from the Indo-Pacific, was collected as early as 1929 

in O‘ahu on rocks (Henry & McLaughlin (1975). Henry & McLaughlin listed additional material 

from “off Oahu” on buoys (1944), Pearl Harbor (1931, 1948), Honolulu Harbor (1954), and 

Kāne‘ohe Bay (1959). Southward et al. (1998) report it in 1956 in the Ala Wai Canal. It was also 

noted by Grovhoug and Rastetter (1980) for Kāne‘ohe Bay and Pearl Harbor (1976–1977 collections). 

It is now widespread around O‘ahu (Coles et al., 1999a, 1999b), and is also reported from 

Nawiliwili and Port Allen Harbors, Kaua‘i, and Hilo Harbor (Coles et al., 2004). DeFelice et al. 

(2002) report it from French Frigate Shoals. We regard it as introduced via ship fouling. 

Balanus crenatus Bruguière, 1789 


Long (1974) recorded this North American and European barnacle from fouling panels off Pearl 

Harbor. There are no records before or after Long’s report. The record is considered dubious 

(DeFelice & Godwin, 1999). We do not further consider it. 

Chthamalidae 

Chthamalus proteus Dando & Southward, 1980 Introduced 

John Hoover of Honolulu was the first to formally document the existence of this barnacle in the 

Hawaiian Islands when he photographed it in Kāne‘ohe Bay in March 1995 (the date imprinted on 

the first photographic slide of this barnacle; J. Hoover, pers. comm., June, 1999). When the first surveys 

were undertaken in 1996 to establish the extent of this species, it was already found to be widespread 

around O‘ahu, including Pearl Harbor (Southward et al., 1998; Coles et al., 1999a), and by 

1996–1998 it had been found on Kaua‘i, Maui, Midway Island, and Guam (Southward et al., 1998). 

It now occurs in the South Pacific as well (Zabin et al., 2007). 

The first known specimens are from Pearl Harbor, collected in 1993 by Julie Bailey-Brock and 

first thought to be a native species (Zabin et al., 2007). 

John Hoover submitted his photographs to W.A. Newman for identification, who, familiar with 

the barnacle fauna of the Hawaiian Islands, recognized it as new to the region. Morphological and 

genetic evidence revealed that it was the tropical Western Atlantic barnacle Chthamalus proteus, 

known from the Caribbean and the Gulf of Mexico at least as far west as Louisiana (Southward et 

al., 1998). Southward et al. noted that the “date of introduction was after 1973 [the last thorough bar-


107 

nacle surveys of O‘ahu] and it could have been as recently as 1994 or 1995... However, considering 

the present distribution of C. proteus in the islands and the usual lag time between an introduction 

and notable abundance, it was possibly earlier... .” The dates of first records of C. proteus in Hawai‘i 

are not given in Southward et al. (1998); these are 1993 and 1995, as noted above. Its abundance and 

widespread distribution by 1995–1996 suggest an earlier inoculation. 

Introduction could have been either on ships’ hulls [in waterline fouling for this high intertidal 

barnacle (Southward et al., 1998)] or as larvae in ballast water. Southward et al. suggest that ballast 

water “is less likely than transport of adults, since a dense settlement is needed to establish a breeding 

population of such obligate cross-fertilizing sessile animals.” We note, however, that the discharge 

of nauplii or cyprids within an embayment or lagoonal environment (such as Pearl Harbor) 

would provide such opportunities, as discussed below for the crab Scylla serrata. This said, the probable 

mechanism was hull fouling, as the survival of viable larvae in ballast water from the Atlantic 

to Hawai‘i would appear to be less likely than survival on a hull. 

In what may be a unique documentation of the same adult hull fouling organisms making round - 

trips between ports, Godwin (2003b) noted that Chthamalus proteus was on barges that had left 

Hawai‘i, gone to California and the Pacific Northwest, and were still alive on the barges when they 

returned to Honolulu Harbor. As C. proteus is not yet established in the ports-of-call where these 

barges visited on the American Pacific coast, there is no question that the same individuals are 

involved. Godwin noted that these barges are on regularly scheduled routes between the Islands and 

the mainland. Zabin et al. (2007) comment on the potential for C. proteus to invade the west coast, 

from approximately San Diego and south. 

Chthamalus proteus settles on many substrates in the intertidal zone (Zabin et al., 2007) and 

can form very dense populations on mangroves on O‘ahu (J.T. Carlton, personal observations, 1999). 

Primavera (1999) has observed that Rhizophora mangrove seedlings in the Philippines may be vulnerable 

to barnacle infestation and thus lower survival. 

Zardus & Hadfield (2005) found that C. proteus in the Pacific represent multiple invasions from 

the Atlantic, including both the Caribbean and Brazil. Zabin et al. (2007) compare aspects of the distributional 

ecology and life history of C. proteus in the Caribbean, Atlantic, and Pacific Oceans. 

Zabin & Altieri (2007) demonstrated that C. proteus preferentially settles where the native pulmonate 

limpet Siphonaria normalis occurs but that limpets depart areas of high barnacle densities. 

Orders Decapoda (crabs, shrimps, and lobsters) and Stomatopoda (mantis shrimps) 

Introduced Decapoda and Stomatopoda 


Decapoda 

Macrobrachium lar 1956 R Guam & Tahiti 

Scylla serrata 1926 R Samoa 

Panopeus lacustris 1947 SF,BW Western Atlantic 

Acantholobulus pacificus 1929 SF,BW Eastern Tropical Pacific 

Pilumnus oahuensis 1929 SF,BW Unknown 

Glabropilumnus seminudus 2003 SF, BW Indo-Pacific 

Nanosesarma minutum 1996 SF,BW Japan, Indo-Pacific 

Pachygrapsus fakaravensis 1996 SF,BW Japan, French Polynesia 

Metopograpsus oceanicus 2006 SF,BW Indo-Pacific 

Hyastenus spinosus

108 




Decapoda 

Upogebia pugettensis 1916 R, BW Northeast Pacific 


Hemigrapsus penicillatus 

Status: Erroneous record?; see discussion 

Macrobrachium rosenbergii 1965 R Indo-Pacific 

Status: Released; not established 

Carcinus maenas 1873 SF Northwest Atlantic 


Callinectes sapidus 1985–1992 R Northwest Atlantic 

Status: Released; not established 

Charybdis helleri 


Atergatopsis immigrans 


Schizophrys aspera 


Eriocheir sinensis 

Status: Erroneous record; see discussion 

Litopenaeus stylirostris 

Litopenaeus vannamei 

Marsupenaeus japonicus 

Penaeus monodon 

Status (above four species: ) Escaped; Not Established 

Fenneropenaeus chinensis 

Fenneropenaeus indicus 

Status: Transported to island; see text 

Stomatopoda 

Gonodactylellus hendersoni 

Status: Native 

Order Decapoda 

Thalassinidea (mud shrimps) 

Upogebiidae 

Upogebia pugettensis (Dana, 1852) 

Deleted 

A single specimen of a Upogebia was collected in Hawai‘i, probably on O‘ahu, in 1916 by W.A. 

Bryan (Edmondson, 1933, 1944b, 1946a; K. Sakai, 1982). The specimen was identified in 1977 by 

K. Sakai as Upogebia pugettensis, a well known mud shrimp from the North American Pacific coast. 

Sakai (1982) referred to this Hawaiian specimen as Upogebia littoralis; this is a lapsus, however, as 

U. littoralis is an older name for two different Atlantic and Mediterranean species. Sakai (in litt., 

February 2000) indicated to us that it was likely U. pugettensis, and this is the name that appears on 

the specimen label as identified by Sakai. The specimen was missing for many years from the Bishop 

Museum collections but was found in August 2004 in the laboratory of Professor de Saint Laurent 

in the Muséum National d’Histoire Naturelle in Paris by Alain Crosnier. 

Edmondson and Sakai both suggested that it may have been an accidental introduction from 

North America. The planting of oysters from the California coast in the islands does not correlate in 

time with the discovery of this bay-dwelling thalassinid in Hawai‘i. It may be that it arrived with 

unreported shellfish shipments from the mainland, or as a young stage in ballast water. This odd and 

isolated record is thus reminiscent of a single specimen of the western Atlantic Upogebia affinis that 

was found in 1912 in San Francisco Bay (Williams, 1986).


109 

Caridea (shrimp) 

Penaeidae 

Litopenaeus stylirostris (Stimpson, 1874) 

Escaped; Not Established 

= Penaeus stylirostris 

(native to Baja California to Peru) 

Litopenaeus vannamei (Boone, 1931) 


= Penaeus vannamei 

(native to Gulf of California to Peru) 

Marsupenaeus japonicus (Bate, 1888) 


= Penaeus japonicus 

(native to Eurasia, Africa, South Pacific) 

Penaeus monodon Fabricius, 1798 


(native to Eurasia, Africa, South Pacific) 

Penaeid shrimp culture commenced in the Hawaiian Islands in the 1970s. The above four species 

(whose range data are from Perez Farfante & Kensley, 1997), as well as Fenneropenaeus chinensis 

(Osbeck, 1765) (= Penaeus chinensis) and Fenneropenaeus indicus (H. Milne Edwards, 1837) (= 

Penaeus indicus) were transported to the islands for aquaculture and research (Eldredge, 1994). 

Primarily during pond flooding, the above four species are known to have escaped, but none became 

established (Brock, 1992a, 1992b; Davidson et al., 1992; Eldredge, 1994). 

Palaemonidae 

Macrobrachium lar (Fabricius, 1798) 

Introduced 

Brock (1960) and Eldredge (1994) reviewed the history of the prawn Macrobrachium lar in Hawai‘i; 

individuals were brought from Guam in 1956 and from Tahiti in 1961. They are now established in 

streams on all main islands. We include Macrobrachium here because its life history includes a marine 

phase, with larval stages spent in brackish and marine waters. Adult prawns may also occur in 2–3 ‰ 

water (Englund et al., 2000a). Maciolek & Timbol (1981) reported it from the freshwater portions of 

the Kahana Estuary, O‘ahu, based on collections made from 1969 to 1971. It was first intentionally 

imported in 1956 from Guam (released in Moloka‘i) and in 1957 (released in Nu‘uanu Stream, O‘ahu). 

The native range is the Indo-Pacific, from East Africa to the Ryukyu Islands and the Marquesas. 

In Hawai‘i, M. lar is said to be “in direct competition with the only native prawn, M. grandimanus” 

(Eldredge, 1994). Macrobrachium lar and M. grandimanus overlap in relatively small 

regions of brackish water, with M. lar being found upriver in freshwater and M. grandimanus being 

found downriver in brackish water; M. lar may be more likely to interact with the native freshwater 

shrimp (R. Englund, pers. comm., February 2000). Macrobrachium lar preys on native freshwater 

snails and gobies in Hawai‘i (Englund et al., 2000a). 

Macrobrachium rosenbergii (deMan, 1879) 


Eldredge (1994) reviews the history of the Indo-Pacific giant freshwater prawn Macrobrachium 

rosenbergii in the Hawaiian Islands and noted that there is no evidence that is became established. 

It was first imported intentionally in 1965, but no reproduction was observed in the wild, and no subsequent 

populations are known. 

Brachyura (crabs) 

A persistent mystery in the Hawaiian exotic crab fauna is the xanthid crab Atergatopsis immigrans. 

Edmondson (1962a) described this as a new species, Neoliomera immigrans, based on “two specimens... 

taken from fouling on the bottom of a barge [from Guam] in a Pearl Harbor dry dock in 

1950.” One female and one male crab were collected. This crab has not been recorded from Hawai‘i 

since—nor, curiously, apparently from anywhere else in the world. It is not a failed invasion, in the 

sense that established reproducing populations were never recorded in Hawai‘i. One specimen is at 

the Smithsonian Institution; the other is at the Bishop Museum. Guinot (1969) transferred the species 

into the genus Atergatopsis, commenting that “Au prealable, il conviendrait de verifier si immigrans 

n’est pas synonyme d’une Atergatopsis deja connue” (“To begin with, it is advisable to verify if

110 


immigrans is not a synonym of an Atergatopsis already known”). She noted that A. immigrans was 

similar to both A. granulata Milne Edwards, 1865 and A. lucasi Montrouzier, 1865, species not mentioned 

by Edmondson in his comparison to other species of Neoliomera. Guinot (1971) indicated that 

the assignment to the genus Atergatopsis should be verified by direct examination of the original 

specimens, although later authors (such as Serene, 1984) have followed her generic placement. 

Portunidae 

Scylla serrata (Forsskål, 1775) 

Introduced 

The Samoan crab (also known as the mangrove or red crab) was first introduced into the Hawaiian 

Islands in order to start a fishery in Kāne‘ohe Bay in 1926. Between 1926 and 1935, 98 crabs were 

released on O‘ahu, Hawai‘i, and Moloka‘i, all from Samoa (Brock, 1952, 1960). Given this small 

inoculation size, and the probability that the current populations arose from only a fraction of these 

crabs, genetic studies on the Hawaiian populations would be of interest. By 1940 it had “already 

become thoroughly established about our shores, entering estuaries of streams and ascending far up 

some of the larger rivers” (Edmondson & Wilson, 1940). Edmondson (1954) noted that large specimens 

may exceed 20 cm in breadth and weigh “several pounds” and that it is valued as food “and 

brings a fancy retail price at the Honolulu fish markets”; Brock (1960) made similar comments. The 

Honolulu Advertiser of 13 March 1955 carried a photograph of a specimen from Hawai‘i measuring 

21.6 cm across and weighing 6.4 kg. Maciolek & Timbol (1981) reported it from the Kahana Estuary, 

O‘ahu, based on collections made from 1969 to 1971. Eldredge (1994) noted that as of 1992 it was 

one of the “major species collected in certain areas of the island of Hawai‘i.” Scylla serrata was 

found in Pearl Harbor in 1996 (Coles et al., 1999a). 

Brock (1960) attributed part of the success of the crab, in light of the relatively few individuals 

released, as being due to the fact that “some of the estuarine areas where this species was released 

have a low rate of tidal flushing, a situation which may be conductive [conducive] to the rapid 

growth of a population within the estuarine area.” Cohen et al. (1995) came to a similar and independent 

conclusion relative to the establishment of the European crab Carcinus maenas in San 

Francisco Bay, California: “[lagoons] may nurture new inoculations of nonindigenous species. The 

shallow lagoons are typically a few degrees warmer than the Bay in the spring to fall months, when 

they are characterized by high primary productivity.... The lagoons are also retentive environments, 

which may be crucial to maintaining the critical densities of adult organisms needed for sexual reproduction.” 

Brock (1960) notes a broad distribution from Africa and India through tropical Asia and 

into Polynesia. 

Carcinus maenas (Linnaeus, 1758) 

Deleted 

The U.S.S. Portsmouth, under the command of J.S. Skerrett, visited the Hawaiian and Fanning 

Islands in 1873–1874, while engaged in a survey of the North Pacific Ocean islands. Aboard was 

Assistant Surgeon T.H. Streets of the U.S. Navy (Jordan & Evermann, 1905). On his visit he collected 

one large male specimen of this North Atlantic crab (Streets, 1877, as Carcinides maenas) probably 

in Honolulu Harbor. Edmondson (1954) and subsequent authors doubted the record, Edmond - 

son noting that “So far as I can determine, there is no other record of the species from Hawaii, and 

its presence here now is very doubtful. I have not seen living examples of this species.” The specimen 

in question is at the Smithsonian Institution. We have examined it, and it is Carcinus maenas, 

as also reported in Carlton & Cohen (2003). 

Callinectes sapidus Rathbun, 1896 


Six female specimens of this well-known Atlantic blue crab were trapped between 1985 and 1992 in 

Kāne‘ohe Bay near the He‘eia State Park and Kāne‘ohe Marine Corps base (Eldredge, 1995). Eldredge 

notes that it has been commonly imported into Hawai‘i since 1967, mostly from Louisiana. Font & Tate 

(1994) note that C. sapidus have been observed being packed in Louisiana for air delivery to Hawai‘i, 

and that in Hawai‘i, Hilo fishmarkets often receive shipments of crabs from the Gulf of Mexico. Thus 

it is likely that the crabs found in Kāne‘ohe Bay are intentional private releases.


111 

However, it is of interest that Stephenson (1976) noted a juvenile male Callinectes collected by 

P. Bartsch in September 1920 in Pearl Harbor that was not further identifiable since it lacked the 

abdomen and pleopods, although the specimen resembled C. sapidus. The identity of this specimen 

(in the Smithsonian Institution) may be determinable using molecular techniques. 

Charybdis helleri (Milne Edwards, 1867) 

Intercept 

This crab is consistently cited in the literature as occurring in the Hawaiian Islands. The record is 

that of Edmondson (1954), who, however, only records a single male specimen “taken from among 

the fouling on the hull of a ship in dry dock in Pearl Harbor Navy Yard. The ship had seen service 

in Guam sometime previous to its defouling and may have transported the portunid to Pearl Harbor 

as a juvenile.” This specimen is in the Bishop Museum collections, collected 5 April 1950. Despite 

Edmondson stating that “There is no evidence that this species is established in Hawaii,” subsequent 

authors have recorded the Hawaiian Islands as within the crab’s range. We regard this record as an 

interception and do not further consider it here; it is not a member of the Hawaiian decapod biota. 

Charybdis helleri has since been introduced to eastern North and South America (Tavares & de 

Mendonca, 1996). 

Panopeidae 

Panopeus lacustris Desbonne, 1867 

Introduced 

= Panopeus herbstii of authors, not of H. Milne Edwards, 1834 

This Atlantic mudcrab was first collected in December, 1947 “from the fouling of a boat in Pearl 

Harbor” (Edmondson, 1962a). In 1953, numerous specimens were collected in Maunalua Bay, 

O‘ahu, “where the species appears to be well established. Its introduction into Hawaiian waters 

doubtless came about through transportation on the bottom of a ship in very recent times” 

(Edmondson, 1962a). It remains present in Pearl Harbor (Coles et al., 1999a). 

This species was originally reported from Hawai‘i as Panopeus herbstii, but Williams (1983) 

split this taxon into six species. Williams’ reexamination of O‘ahu material at the Smithsonian 

Institution revealed which of the six species the Hawaiian population represented. While Panopeus 

herbstii sensu lato was a crab ranging from cool temperate New England waters to tropical Carib - 

bean waters (and thus the appearance of this crab in Pearl Harbor could also have been linked to 

importations of Atlantic oysters from more northern waters), the resolution of the Hawaiian populations 

as P. lacustris provides finer scale geographic and transport mechanism resolution. Panopeus 

lacustris is a subtropical to tropical crab, ranging from Bermuda and southern Florida through the 

Caribbean and south to Brazil. It occurs over a wide range of shallow-water habitats in the western 

Atlantic Ocean, including algal mats, sabellariid reefs, mudflats in seagrass beds, under coral debris, 

and on mangroves. It is thus a member of the “Caribbean element” now found in the Hawaiian 

Islands. It likely arrived in ship’s fouling or in ballast water, as commercial oysters are not known to 

have been transported to Hawai‘i from regions where P. lacustris is found. 

Acantholobulus pacificus (Edmondson, 1931) 

Introduced 

= Panopeus pacificus Edmondson, 1931 

= Neopanope sp. Edmondson, 1931 (1929, Pearl Harbor), fide Forest & Guinot (1961) and Felder & Martin 

(2003) 

= (?) Acantholobulus mirafloresensis (Abele & Kim, 1989, new combination in Felder & Martin, 2003, for 

Panopeus mirafloresensis). 

Charles H. Edmondson (1931) described this crab as a new species from Pearl Harbor. While 

Edmondson did not provide a collection date, the type material in the Bishop Museum bears the date 

December 1929. Edmondson (1931) noted that it was “associated with sponges, barnacles and tunicates 

attached to buoys and floats ... . The barnacle (Balanus eburneus Gould), among which the crab 

is found, is a typical species of the east coast of the United States, which is suggestive of the view 

that both crabs and barnacles may have been transported to Hawaii through shipments of oysters or 

on the bottoms of ships.” Edmondson (1933) notes that it also occurred in Pearl Harbor associated 

with the crab Pilumnus oahuensis. Many years later, Edmondson (1962a) repeated his thought that

112 


“As P. pacificus has been taken only in Pearl Harbor among fouling complexes on buoys, it may 

have been transported to Hawaii on the bottoms of boats.” It remains common today in Pearl Harbor 

(Coles et al., 1997, 1999a) and also occurs in Kāne‘ohe Bay (Coles et al., 2002a; 2000 collections), 

and in Ke‘ehi Lagoon, Ala Wai Harbor, and Kewalo Basin (Coles et al., 1999b, 1997–1998 collections), 

all on O‘ahu. Coles et al. (2006) report it from Kaua‘i, based upon 2002 collections; it was 

found in 2003 at Kaunakakai Harbor, Moloka‘i, and Hilo Harbor (Coles et al., 2004). 

In an earlier draft of this monograph in 2001 we wrote: “We regard it as introduced in ship fouling 

or ballast water. As colder water northern hemisphere estuarine panopeids are relatively well 

known and have not been matched with P. pacificus, we predict that it will be found to be a junior 

synonym of an earlier-named warmer-water if not outhern hemisphere panopeid.” 

Based upon the revisionary work of Felder & Martin (2003), it may be native to either the tropical 

eastern Pacific Ocean or to the tropical western Atlantic Ocean. In the tropical eastern Pacific it 

may have gone under the names of P. mirafloresensis or P. bermudensis of authors (Felder & Martin, 

2003, place eastern Pacific records of P. bermudensis Benedict & Rathbun, 1891 into P. mirafloresensis). 

Felder & Martin (2003) note that “except for the apparent larger size evident in most specimens 

of A. pacificus, there are ... no obvious morphological characters to distinguish it from A. 

mirafloresensis.” We note that introduced populations of nonnative species often tend to a larger 

body size than their conspecifics in their native regions. 

Acantholobulus mirafloresensis occurs “throughout coastal waters of the tropical eastern Pacific 

region” (Felder & Martin, 2003). If the Hawaiian populations of A. pacificus should morphologically 

and genetically match eastern tropical Pacific populations of A. mirafloresensis, we conclude that this 

crab, having attached to ship bottoms possibly in the Panama area, was introduced in hull fouling to 

Pearl Harbor from the tropical eastern Pacific in the early decades of the twentieth century. 

However, Felder & Martin (2003) also note that Acantholobulus bermudensis (Benedict & 

Rathbun, 1891) cannot be morphologically distinguished from Acantholobulus mirafloresensis. As is 

common in transisthmanian systematics and biogeography, these auhors retain the two species as western 

Atlantic and eastern Pacific cognates, respectively. Should these two taxa prove identical genetically, 

A. mirafloresensis will become a junior synonym of A. bermudensis. If A. pacificus and A. mirafloresensis 

prove to be genetically identical, then both will fall to the synonymy of the older A. bermudensis. 

Forest & Guinot (1961) reported Panopeus pacificus from Tahiti based upon two specimens 

collected in 1901. No further material has apparently been collected in Tahiti over a century. We suggest 

that A. pacificus no longer occurs in Tahiti, and that the specimens represent a failed introduction 

from ship hull fouling, hailing either from the eastern Pacific or from the western Atlantic. 

We conclude that Acantholobus pacificus is unquestionably introduced to Hawai‘i. Its exact 

biogeographic origin and systematic affinities await further work on related taxa in the Americas. 

Pilumnidae 

Glabropilumnus seminudus (Miers, 1884) 

Introduced 

Coles et al. (2004, 2006) found this crab in 2003 on two islands, at Moloka‘i (Hale o Lono Reef) 

and at Maui (Kahului Harbor, Pier 1). It occurs broadly through the Indo-West Pacific (Davie, 2002). 

Davie (2002) lists Hawai‘i in its distribution, a record based upon the report of Edmondson (1952b). 

Edmondson’s record, however, was based upon a single specimen found on the same Guam barge in 

1950 as noted for Schizophrys aspera; it was not known to be established in Hawai‘i at the time of 

Davie’s listing. 

Pilumnus oahuensis Edmondson, 1931 

Introduced 

Edmondson described this species based upon specimens collected in 1929 (Bishop Museum type 

specimen) from Pearl Harbor. Edmondson found it “associated with masses of sponges, tunicates 

and barnacles which are attached to buoys, floats, and piling.” Edmondson (1962a) added Honolulu 

Harbor (BPBM material from Honolulu Harbor commences in 1947). It was also noted by Grovhoug 

& Rastetter (1980) for Kāne‘ohe Bay and Pearl Harbor (1976–1977 collections). 

It is still found in Pearl Harbor in fouling (Coles et al., 1999a) as well as in other harbors and


113 

bays around O‘ahu (Honolulu Harbor, Ke‘ehi Lagoon, Ala Wai Harbor, Barbers Point Harbor, 

Kewalo Basin [Coles et al., 1999b]; Kāne‘ohe Bay [Coles et al., 2002a] and Waikīkī [Coles et al., 

2002b]). Coles et al. (2004, 2006) report it from Kaua‘i, Moloka‘i, Maui, and the island of Hawai‘i, 

based upon 2003 collections; these are the first reports outside of O‘ahu. It has not been recognized 

elsewhere in the Pacific. Because of its harbor and habitat restriction we regard this species as 

unquestionably introduced through ship fouling or ballast water. It may eventually be recognized as 

a previously described xanthid from elsewhere in the Pacific. 

Xanthidae 

Atergatopsis immigrans (Edmondson, 1962) 

Intercept 

This crab is discussed in the introduction to Brachyura, above. 

Grapsidae 

Nanosesarma minutum (De Man, 1887) 

Introduced 

This tiny widespread Pacific and Indian Ocean crab was first collected in 1996 in fouling communities 

in Middle Loch, Pearl Harbor (Davie, 1998, Coles et al., 1999a) where it was found to be common. 

We regard it as introduced in ship fouling or in ballast water. Vannini & Valmori (1981) report 

it from Somalia to Tanzania, Madagascar, Singapore, Thailand, and Java as “common on rotten 

wood half buried in the mud, in mangroves.” Dai & Yang (1984) report it from China, Japan, 

Indonesia, Singapore, Thailand, India, and Madagascar, from intertidal “muddy flats and under 

stones.” Davie (1998), who adds the Hong Kong-Japanese species Nanosesarma gordoni (Shen, 

1935) to the synonymy of this species, notes it is found “in fouling and amongst oysters ... in the 

intertidal and shallow subtidal zones of sheltered shores” and that “it is common around major shipping 

ports such as Hong Kong and Singapore.” Under the name N. gordoni, Sakai (1976) noted it is 

found intertidally “among the oysters or sponges growing on rocky shore(s).” 

Pachygrapsus fakaravensis Rathbun, 1907 

Introduced 

This rocky intertidal crab, previously known from French Polynesia (the type locality is the 

Tuamotus) and Japan, was collected in 1996 by Darryl Takaoka from Barbers Point Harbor, O‘ahu 

(Davie, 1998), based upon two specimens. Peter Ng (pers. comm., February 2000) found it to be common 

at K‘ewalo, O‘ahu, in February 2000. Sakai (1976) notes the habitat as “coral reef, at high tidal 

mark.” Barbers Point is the major container ship terminal on O‘ahu. We regard it as introduced by ballast 

water, as modern container ships are unlikely to support extensive external fouling communities, 

nor would a ship’s seachest appear to be a conducive environment for this rocky shore crab. 

Metopograpsus oceanicus (Jacquinot, 1852) 

Introduced 

This crab was previously known from East Africa to the Philippines and Palau, and was first collected 

in Guam in 1997 (Paulay, 2007). In February 2006, Gustav Paulay found this crab in the supratidal 

of Coconut Island, Kāne‘ohe Bay, where it co-occurs with the native Metopograpsus thukuhar 

(Paulay, 2007). As Paulay (2007) notes, there has been little documentation of the intertidal crab 

fauna of the Hawaiian Islands since the 1960s, so it is difficult to judge how long this crab has been 

in the region. That said, it would not have been overlooked by C.H. Edmondson in the 1950s had it 

been present. Metopograpsus collections from O‘ahu made by Darryl Takaoka in the 1990s, and now 

in the Bishop Museum collections, await re-examination. 

varunidae 

Hemigrapsus penicillatus (de Haan, 1835) 


Asakura & Watanabe (2005) note that Hemigrapsus penicillatus has been reported from Hawai‘i. 

The record is based upon six specimens collected by W.A. Bryan in 1903 “at Laysan Island;” these 

were deposited in Bishop Museum Collections as catalog number 236 (Edmondson, 1951, pp. 

236–237; 1959, pp. 180–181). However, the specimens are lost. An error in initial reporting may

114 


have been involved; the Bishop Museum ledger catalog number 236 refers to six specimens of this 

crab, but from Japan (where the species is native and common). 

Eriocheir sinensis Milne Edwards, 1854 


Reports in the literature imply that this well-known Chinese mitten crab was or is present in the 

Hawaiian Islands. For example, Rudnick et al. (2000) note that it “was also found in Hawaii in the 

1950s, with no subsequent reports of occurrences,” citing Edmondson (1959). There are however no 

records, past or present, of which we are aware of Eriocheir sinensis in the Hawaiian Islands. 

Edmondson (1959, p. 155) makes a passing reference to this crab being used as food in Asia. 

Majidae 

Hyastenus spinosus A. Milne-Edwards, 1872 

Introduced 

Tinker (1965) noted that this “spiny spider crab”, ranging from “Hawaii southward and westward 

across the tropical Pacific and Indian Oceans” was “found in shallow water where the marine growth 

was abundant. It is known to dwell on piers and pilings which are covered with a variety of algae 

and invertebrate animals.” It is not mentioned by Sakai (1965, 1976) nor Dai & Yang (1984) for 

Japan and China, nor by Poupin (1994) relative to the common crabs of French Polynesia. However, 

Griffin (1974, 1976) recorded it from the Indian Ocean, detailing its distribution as Indo-West 

Pacific, from the Red Sea and Gulf of Aden, eastern Africa from Durban to Cape Guardafui, Sri 

Lanka, the coasts of India, Singapore, Gulf of Siam, northern Australia, Philippine Islands, and up 

to Fiji. However, its habitat outside of Hawai‘i is generally reported as in somewhat deeper waters 

from 12–67 fathoms, on coarse sand and broken shells, although reported in Torres Strait, Australia, 

from 5–7 fathoms (Griffin, 1966, 1976). 

Curiously, the first Hawaiian record appears to be the mention by Tinker (1965) in his popular 

book. Not a tiny crab, it was not mentioned in any of C.H. Edmondson’s earlier books and papers 

on the Hawaiian fauna in general or the decapod fauna in particular, and it thus may be a fairly recent 

(post-World War II) arrival. No early Hawaiian specimens exist in Bishop Museum collections, 

although two specimens were found in 1996 in Pearl Harbor (BPBM collections; Coles et al., 1997). 

No similar crab has been encountered elsewhere in the Islands. 

Based upon its localization in Pearl Harbor, association with fouling communities, and its relatively 

recent reporting from the Islands, we consider it introduced. 

Schizophrys aspera (H. Milne Edwards, 1834) 

Intercept 

Edmondson (1951) thought this majid might have been established on O‘ahu, but the specimens he 

had in hand were from a barge newly arrived in 1950 from Guam into Pearl Harbor. Davie (2002) 

includes Hawai‘i in the range of this species, but S. aspera does not occur here. 

Stomatopoda (mantis shrimps) 

Gonodactylidae 

Gonodactylaceus falcatus (Forsskål, 1775) 

Introduced 

= Gonodactylaceus mutatus (Lanchester, 1903), as used by Barber & Erdmann (2000) for Hawaiian populations 

of G. falcatus 

= Gonodactylus aloha Manning & Reaka, 1981 

In a well-known paper on invasion processes, Kinzie (1968) argued that an Indo-Pacific species of 

mantis shrimp (which he identified as Gonodactylus falcatus) was introduced to the Hawaiian 

Islands. Manning & Reaka (1981) subsequently described the same Hawaiian population as a new 

species, Gonodactylus aloha, and considered it endemic. Kinzie (1984), in a paper titled, “Aloha also 

means goodbye,” examined their arguments in detail and concluded that at the least the species was 

cryptogenic. Carlton (1987) also referred to it as cryptogenic, noting that “G. aloha may be native 

to, but undiscovered in, the southwestern Pacific and/or [there may be] morphological distinctions 

in Hawaiian populations due to founder effect.” Carlton (1996) later treated it as clearly introduced, 

closing a nearly 30-year circle in this species’ Hawaiian history.


115 

Manning & Lewinsohn (1986) restricted Gonodactylaceus falcatus to the Red Sea, and provided 

comparisons of G. aloha and G. falcatus. Manning (1995) expanded the view of G. falcatus to 

include populations both in the Red Sea and Indian Ocean to at least Mombasa, Kenya and 

Mauritius, and possibly to New Caledonia and Japan, although bringing the last two locations into 

question. According to the data of Manning and Reaka, the two species are identical or overlap in 

all characters with the exception that G. aloha lacks and G. falcatus has a median caruncle on the 

sixth abdominal segment. However, in the original description of G. aloha, this character is said to 

be variable (“median caruncle occasionally present on sixth somite, rarely well-developed”). The 

color characteristics chosen by Manning & Lewinsohn (1986) show overlap in color between the 

two species, although Manning & Reaka (1981) argue for other color differences (but see Kinzie, 

1984). Manning & Reaka’s color and morphological distinctions between G. aloha and G. falcatus 

were based on Red Sea specimens of G. falcatus. 

Gonodactylaceus aloha has thus been on uncertain grounds for some time. Barber & Erdmann 

(2000) and Ahyong (2001, 2002) concluded that G. aloha was a synonym of G. mutatus, a species 

previously known from New Caledonia, southern China, and Vietnam to the western Indian Ocean. 

Ahyong (2001) then concluded that G. mutatus was a junior synonym of G. falcatus (and concurred 

that G. aloha is indeed a synonym of G. mutatus), and thus the name returns to the same one used 

by Kinzie originally, closing another loop of more than 30 years duration. 

The first specimens of G. falcatus were observed in Hawai‘i 1954 in dead coral heads in 

Kāne‘ohe Bay [Kinzie, 1968, who also noted that another species of Gonodactylus was observed at 

the same time on the Waikīkī reef, but this may have been Gonodactylellus hendersoni (see Kinzie, 

1968, p. 469, and discussion, below)]. Kinzie suggested that it was introduced onto O‘ahu with concrete 

barges towed to the islands at the end of World War II, particularly from the area of the 

Philippines and the South China Sea. It was also noted by Grovhoug & Rastetter (1980) for 

Kāne‘ohe Bay and Pearl Harbor (1976–1977 collections). Brock (1995) found it in Pearl Harbor 

(1993–1994), where it remains (Coles et al., 1999a). It is now widely distributed throughout the 

main islands, including O‘ahu, Moloka‘i, Maui, Kaua‘i, and Lāna‘i. Kinzie (1968) demonstrated 

experimentally that the more aggressive G. falcatus had displaced the native stomatopod 

Pseudosquilla ciliata from coral head habitats in Kāne‘ohe Bay. Caldwell & Dingle (1975) reviewed 

Kinzie’s study. Reaka (1975, 1976, 1979a, 1979b) reported upon molting and life history of G. falcatus 

from Kāne‘ohe Bay. 

Given the vast movement around the Pacific Basin during and after World War II of landing 

craft, barges, and other vessels with inhabitable holes and crevices, it is remarkable that this is the 

only recognized stomatopod introduction. It seems probable that given the assumption that most 

species of Pacific stomatopods are widespread across the Pacific Ocean, new discoveries were attributed 

to natural larval dispersal, when in fact many populations of stomatopods discovered in the last 

half of the twentieth century may represent cryptogenic if not introduced records. 

Gonodactylellus hendersoni (Manning, 1967) 

Native 

Kinzie (1968) speculated that this stomatopod was introduced to O‘ahu. It is a widely distributed 

species across the central and Indo-Pacific Oceans. Roy Caldwell (pers. comm., February 2000) 

reports that he has examined specimens of this species collected in Hawai‘i many years before the 

first record of 1958 reported by Kinzie (1968) and repeated by Manning & Reaka (1981). Caldwell 

believes that it is a naturally widespread species throughout the Pacific and may have been overlooked 

earlier in Hawai‘i because it is relatively small, generally found in subtidal rubble, and not 

as easy to collect as larger stomatopods. More broadly, the taxonomic status of the Hawaiian populations 

remain uncertain, pending additional examination of more specimens (S. Ahyong, pers. 

comm., 2001).

116 


INSECTA 

Introduced and Cryptogenic Insecta 



Odonata 

Enallagma civile 1936 R, O Western North America 

Ischnura ramburii 1973 R, O North/South America 

Crocothemis servilia 1994 R, O Middle East, Asia, 

Australia 

Tramea lacerata 1873 R, O Eastern North America 

Hemiptera 

Trichocorixa reticulata 1877 R, O North America 

Mesovelia amoena 1971 R, O North/South America 

Mesovelia mulsanti 1933 R, O North/South America 

Micracanthia humilis 1988 R, O North America 

Hymenoptera 

Kleidotoma bryani 1923 R, O Guam, Palmyra Island 

Coleoptera 

Enochrus sayi 1931 O Western North Atlantic 

Tropisternus salsamentus 1968 BW California 

Cercyon fimbriatus 2001 R, O Eastern Pacific 

Parathroscinus murphyi 1996 BW Southeast Asia 

Diptera 

Canaceoides angulatus 1922 R, O Tropical E Pacific 

Procanace williamsi 1944 R, O Asia (Japan?) 

Tethina willistoni 1919 R, O Western Atlantic 

Atrichopogon jacobsoni 1958 R, O Western Pacific 

Atrichopogon sp. 1998 R, O Unknown 

Cricotopus bincinctus 1955 R, O Unknown 

Goeldichironomus holoprasinus 1969 R, O North/South America 

Telmatogeton japonicus 1946 R, O Japan 

Dolichopus exsul 1930 R, O West Indian (Caribbean) 

Medetera grisescens 1914 R, O Indo-Pacific 

Syntormon flexible 1917 R, O Indo-West Pacific 

Psychoda salicornia 1945 R, O California 

Brachydeutera ibari 1980 R, O Asia 

Ceropsilopa coquilletti 1946 R, O North America 

Clasiopella uncinata 1946 R, O Taiwan, Australasia 

Discocerina mera 1948 R, O Western/South Pacific 

Donaceus nigronotatus 1958 R, O Asia 

Ephydra gracilis 1946 R, O Eastern Pacific 

Ephydra milbrae 1950 R, O Eastern Pacific 

Hecamede granifera 1923 R, O Western/South Pacific 

Mosillus tibialis 1944 R, O North America 

Paratissa pollinosa 1945 R, O Caribbean, S. America 

Placopsidella marquesana 1951 R, O Indo-Pacific 

Psilopa girschneri 1952 R, O Unknown 

Scatella stagnalis 1967 R, O Unknown 

Hostis guamensis 1946 R, O Australia 


Collembola 

Oudemansia esakii 1939 R, O 

Coleoptera 

Thinophilus hardyi 1996 R, O 

Dermaptera 

Anisolabis maritima 1912 SBA 

Orthoptera 

Thetella tarnis


117 

Englund (2002) referred to all of the species treated below as introduced to Hawai‘i, but also indicates 

in a column labeled “mode of introduction” that many of these species are cryptogenic as well. 

However, Englund used “cryptogenic” to mean an unknown mechanism of introduction, rather than 

(by definition) an unknown biogeographic origin of the species. Englund’s cryptogenic corresponds 

to the term polyvectic (two or more possible mechanisms of introduction) as used by Carlton & Ruiz 

(2005). 

We treat 43 species of brackish-water and maritime insects here, largely as examples of this 

introduced facies. We are not aware of any review of the invasive marine and maritime insects of the 

Hawaiian Islands: a thorough study of the shoreline maritime arthropods of the Islands may well 

reveal dozens, if not scores, of additional nonnative species. 

Odonata (damselflies and dragonflies) 

Coenagrionidae 

Enallagma civile (Hagen, 1862) 

Introduced 

The “familiar bluet” damselfly, introduced from western North America and first found in 1936 on 

O‘ahu, a species also living in brackish waters in Pearl Harbor in Honouliuli Stream (16 ‰ salinity), 

and on the O‘ahu south shore at Ordy’s Pond, Barbers Point (24 ‰) (Englund et al., 2000a, 

2000b). Polhemus & Asquith (1996) provide a brief summary. 

Ischnura ramburii (Sélys-Longchamps, 1850) 

Introduced 

The “Rambur’s forktail” damselfly, native to North and South America, and found at Hilea, Hawai‘i 

in 1973 (Englund et al., 2000a, 2000b). “Ischnura ramburii has the distinction for being the most 

saline tolerant introduced damselfly, with adults observed around water up to 32 ‰ salinity” 

(Englund, 2000a). Although I. ramburii reproduces only in freshwater, the larvae and juveniles occur 

in 10 ‰, and adults are common around salt water environments (R. Englund, pers. comm., 2000); 

it is thus present in an ecological sense in the estuarine-marine facies. Polhemus & Asquith (1996) 

provide a brief summary. Based upon a preponderance of records, we treat this species biogeographically 

as eastern American. 

Libellulidae 

Crocothemis servilia (Drury, 1770) 

Introduced 

The “scarlet skimmer” dragonfly, native to the Middle East, Asia, and Australia, first collected in 

taro fields in Waiahole Stream, O‘ahu, in 1994; it is found as well in estuarine areas on O‘ahu, 

including Pearl Harbor (Englund et al., 2000a, 2000b). Polhemus & Asquith (1996, p. 49) give a 

color photograph. 

Tramea lacerata Hagen, 1862 

Introduced 

The “black-mantled glider” dragonfly, native to eastern North America, was first recorded in O‘ahu 

in 1873 (Englund et al., 2000b). It is present at Ordy’s Point, Barbers Point, and is saline tolerant, 

with females observed ovipositing in 24 ‰ water (Englund et al., 2000b). 

Dermaptera (earwigs) 

Anisolabis maritima (Bonelli, 1832) 

Cryptogenic 

While there are many early literature records of this widespread littoral earwig on the islands, many 

of these are now assigned to a number of endemic species (Brindle, 1979). There appear to be few 

valid records of the “true” A. maritima; we regard these as potentially ship-introduced populations, 

perhaps with early ballast dumpings. Brindle (1979) admitted only two valid records: specimens 

recorded on Laysan Island by Fullaway (1914), and material collected at Ka‘alaea Beach, O‘ahu 

(1953) and in beach litter in the west Loch of Pearl Harbor (1974). It also occurs on Midway Island, 

where it has been since at least 1960 (Nishida & Beardsley, 2002). The Laysan Island material would 

have been collected between 1905 and 1912 (F. Howarth, pers. comm., 2003). Nishida (2002) listed 

it from many locations from Midway Island down through the Archipelago.

118 


Orthoptera (crickets, grasshoppers) 

Thetella tarnis Otte & Alexander, 1983 

Cryptogenic 

We thank F.G. Howarth for pointing out to us this maritime cricket. It was described from Australia 

(at Portland Roads, Cape York) as “abundant in piles of stone around pier.” It was subsequently 

reported from Hawai‘i (without collection date) by Otte (1994) with a habitat noted as “Rocky 

coastal areas. Lives among wet boulders and in cracks and crevices in the splash zone of all islands. 

In places it is found among roots and in trash along the beach.” It is a widespread species across the 

Pacific and Indian Oceans (D. Otte, pers. comm., September 2005). Modes of transport include rafting 

(these crickets lay their eggs on shore vegetation and sticks that could thus be transported by 

ocean currents; D. Otte, pers. comm.) or historically with shore ballast. The survival of eggs on vegetation, 

for the length of time required to drift to the Hawaiian Islands, has not been demonstrated. 

We thus consider it cryptogenic, pending genetic analysis of these widespread populations. 

Hemiptera (water bugs) 

Corixidae 

Trichocorixa reticulata (Guérin-Méneville, 1857) Introduced 

= Corixa blackburni White, 1877 (O‘ahu?); = Arctocorixa blackburni 

This well-known North American water boatman was described as Corixa blackburni from the 

Hawaiian Islands by White (1877), based upon specimens collected by the Reverend T. Blackburn. 

Zimmerman (1948b) noted that “it is common in brackish pools in the lowlands” and recorded specimens 

from O‘ahu, Moloka‘i, and Maui. Blackburn’s field notes were published by White (1878) and 

include the following observations of this aquatic insect in Hawai‘i of the 1870s: “Very common in 

salt-water pools on the sea-shore. These pools are formed artificially for the manufacture of salt. As 

the liquid becomes more dense by evaporation, the Corixae migrate to pools more recently filled. 

Some would appear, however, to remain too long, as in the last stage of evaporation, the pools generally 

contain a few dead Corixae.” 

Material in the Bishop Museum collections includes some of the specimens found by the 

Reverend T. Blackburn as well as additional collections from 1904 and later years. 

Williams (1944) provided detailed habitat data for this species (under the name Arctocorixa 

blackburni) in Hawai‘i, including the interesting note that he observed “a single individual at 

Waikiki swimming and diving in shallow water about 130 feet out from the shore.” Englund et al. 

(2000a) report this boatman in 1998 in Pearl Harbor at Pouhala Marsh wetlands near Kapakahi 

stream on a pond near the salt flats with 8–9 ‰ salinity. 

Mesoveliidae 

Mesovelia amoena Uhler, 1894 

Introduced 

This water treader, of North and South American origin (Englund, 2002), was first collected in 1971 

on the island of Kaua‘i at Waikanaloa Cave, Hā‘ena, and on O‘ahu in 1971 along the Waimea River 

(Englund et al., 2000a). Englund et al. (2000a) report the species from brackish streams (to 9 ‰) in 

Pearl Harbor. Based on Moreira et al. (2008), we treat this species and the next as biogeographically 

part of the eastern North American. 

Mesovelia mulsanti White, 1879 

Introduced 

A water treader of North and South American origin first collected in 1933 in a reservoir at Waipi‘o, 

O‘ahu (Williams, 1944; Englund et al., 2000a). Zimmerman (1948b) reported it from Kaua‘i, O‘ahu, 

and Moloka‘i. Englund et al. (2000a) report it from fresh and brackish (to 4.5 ‰) streams in Pearl 

Harbor. 

Saldidae 

Micracanthia humilis (Say, 1832) 

Introduced 

This North American shore bug was first collected in Kawainui Marsh, O‘ahu, in 1988 (Englund et 

al., 2000b). Along the south shore of O‘ahu, Englund et al. (2000b) found it along the lower Ala Wai 

Canal in 28 % and Mākua Stream (15 ‰), as well as at fresh water sites.


119 

Hymenoptera (wasps) 

Eucoilidae 

Kleidotoma bryani Yoshimoto, 1962 

Introduced 

This maritime wasp was first collected in 1923 at Pearl City Peninsula, O‘ahu (Beardsley, 1990). 

Yoshimoto (1962) described this species from O‘ahu, based upon collections in 1937 (Maunalua) 

and 1927 (Koko Head). Beardsley (1990) noted a further specimen from 1965 collected at the 

Waipi‘o Peninsula on O‘ahu. It is otherwise known from Guam and Palmyra Island. Beardsley 

(1990) notes that while the host is unknown, “the collection records suggest that this is a parasite of 

some fly which breeds in a littoral environment.” 


Hydrophilidae 

Enochrus sayi Gundersen, 1977 

Introduced 

This Eastern North American water scavenger beetle, which occurs in salinities of up to 16 ‰, was 

reported by Englund et al. (2000a) from Pearl Harbor fresh and brackish streams based upon 1998 

collections. The earliest O‘ahu records are from 1931 from Hale‘iwa (Hansen, 1995). Other records, 

from 1937 to 1982, are from Barbers Point, ‘Ewa, Hickham Air Force base, Honolulu, Kāne‘ohe, 

Kawainui Marsh, Maunalua, and Mt. Tantalus. Englund et al. (2000a) note that the proclivity of this 

species to be attracted to light may have caused colonists to enter ships or planes as they were being 

loaded, and thus carried to the islands. Englund et al. (2000a) provide additional habitat data in Pearl 

Harbor. Howarth & Preston (2002) report Maui as a new island record. 

Tropisternus salsamentus Fall, 1901 

Introduced 

This Californian water scavenger beetle was first collected on Maui in 1968, on O‘ahu in 1970, and 

on Hawai‘i in 1975; it occurs from sea level to 4000 feet (Hansen, 1995). As with Enochrus sayi, it 

is attracted to light. It occurs in fresh and brackish water (up to 16 ‰) in puddles in the Waiawa 

Springs complex in Pearl Harbor (Englund et al., 2000a). Englund et al. (2000b) report it as abundant 

at Ordy’s Pond, Barbers Point, in 24 ‰. Englund (2002) suggests ballast water as the transport 

vector. 

Cercyon fimbriatus Mannerheim, 1852 

Introduced 

Howarth & Preston (2002) report this Pacific American beach beetle from Maui (at the Kahului 

Airport, on the beach and in the strand line) based upon specimens collected in 2001. 

Limnichidae 

Parathroscinus murphyi Woolridge, 1990 

Introduced 

= Parathroscinus sp. Samuelson, 1998 

= Parathroscinus cf. murphyi “Wooldridge”[sic] 1990 of Englund et al., 2000a 

Samuelson (1998) reported this introduced brackish water limnichid beetle as established in Pearl 

Harbor, noting that it “is rather common on mudflats and near streams associated with the harbor.” 

He considered the origin as “probably from Southeast Asia.” Records from Pearl Harbor include 

Pouhala Marsh between Waikele Stream and Kapakahi Stream (1996); mudflats at Waimalu at 

Blaisdell Park (1997) and Honouliuli Stream-estuary area, based on sweepings over mud and saltbush 

(1997). Englund et al. (2000a) note that in Pearl Harbor Parathroscinus “sometimes formed 

dense clouds as these small beetles flew just above the exposed tidal mudflats” and that it “has 

become one of the most common insects on mudflats where mangrove grows.” Englund et al. 

(2000b) note that it is present in “virtually every stream and estuary containing tidal mudflats” on 

the south and west shores of O‘ahu, including the Ala Wai Canal. Englund (2002) remarked that 

“populations have now exploded, and they were found in extremely high densities throughout Pearl 

Harbor.” 

Englund et al. (2000a) note that the Hawaiian material keys to P. murphyi from Singapore. 

Englund (2002) suggests ballast water as the mechanism of introduction.

120 


Anthicidae 

Cyclodinus mundulus (Sharp, 1885) 


Sharp (1895) described this maritime beetle from Hawai‘i as Anthicus mundulus; it was collected 

sometime between 1877 and 1884 (vide the introduction to Blackburn & Sharp, 1895), and was 

found on both O‘ahu and Kaua‘i (vide the table on page 286 in Sharp, 1895: neither the dates nor 

location appear with the species description). The O‘ahu material was collected in Honolulu, according 

to the label with the specimen (Chandler, 2005, page 13). This species is found under seaweeds 

and other drift on beaches, in salt marshes, and occasionally inland but rarely far from the coast. 

Introduction with solid ballast seems probable. 

Werner (1966) mistakenly synonymized Anthicus mundulus with the California beetle then 

known as Thicanus annectens. Chandler (2005) resurrected Cyclodinus mundulus, a Californian- 

Mexican species, noting that “it has been introduced to Hawai‘i [its type locality] and the Dominican 

Republic.” 

Although not verified from the Islands since its first collection (Werner, 1966), two matters 

move us to register this species as Establishment Unknown versus Deleted (the normal assignment, 

based upon no individuals collected for 50 or more years). Young (1979) noted that a data card on 

file at the Bishop Museum listed this species from Kaua‘i, “although [the] record could not be confirmed 

by specimens” (and as of 2007, no specimens of this species were in the Bishop Museum collections). 

The data card noted that the specimens were from “salt marshes near sea level.” The date 

of this material is also not known, but would presumably have been subsequent to the original collections 

(which do not make mention of a habitat on Kaua‘i). In addition, there have been no specific 

searches, as far as is known, for this beetle (D.S. Chandler, pers. comm., February 2007). 

If this beetle has become extinct on the Islands, it would be a rare example of a name being 

based upon an introduced population that no longer exists. 

Diptera (flies) 

Canacidae 

Canaceoides angulatus Wirth, 1969 

Introduced 

This subtropical and tropical Eastern Pacific beach fly was first described from Hawai‘i; the earliest 

material is from Wāwāmulu Beach near Koko Crater in 1922 (Englund et al., 2000a). It is recognized, 

however, as being native to North and South America. It occurs in Pearl Harbor streams in 

salinities of up to 35 ‰ and in a variety of shoreline habitats, such as the splash zone on rocky shorelines, 

on wet sand or mud, and among mangroves (Englund et al., 2000a). Hardy & Delfinado (1980) 

record it from O‘ahu, Kaua‘i, Moloka‘i, Maui, Hawai‘i, Laysan Island, Lisianski Island, Wake 

Island, and Midway Island. Englund et al. (2000b) report numerous recent collections on O‘ahu, 

including the Ala Wai Canal. 

Procanace williamsi Wirth, 1951 

Introduced 

The first record of this beach fly is from 1944 on O‘ahu, from where it was described as a new 

species, although it is now recognized as native to Asia, and perhaps specifically to Japan (Englund 

et al., 2000a). It is one of the most common beach flies on O‘ahu (Englund et al., 2000b). It is common 

in Pearl Harbor streams in salinities of up to 30 ‰ (Englund et al., 2000a) and in many shoreline 

locations on the south shore of O‘ahu, including the muddy banks of the Ala Wai Canal 

(Englund et al., 2000b). 

Tethina willistoni (Melander, 1913) 

Introduced 

=Tethina variseta (Melander, 1952) 

A fly collected in 1919 at Barbers Point (O‘ahu) and later at Wai‘anae, O‘ahu in 1946 (Hardy & 

Delfinado, 1980). It is associated with maritime habitats, including rocky shores and mangroves of 

Pearl Harbor, and in O‘ahu streams along the south shore in salinities up to 42 ‰ (Englund et al., 

2000a, 2000b, as T. variseta). Mathis & Foster (2007) note that it is a Western Atlantic species.


121 

Ceratopogonidae 

Atrichopogon jacobsoni (de Meijere, 1907) 

Introduced 

This biting midge was taken in estuarine waters in Pearl Harbor streams (Englund et al., 2000a), and 

R. Englund advises (pers. comm., 2000) that it be included in our treatment of estuarine and marine 

introduced insects. It was first collected on O‘ahu in 1958, and has “Oriental and Pacific regions” 

origins (Englund et al., 2000a). 

Atrichopogon sp. 

Introduced 

A second introduced species of Atrichopogon occurs both in Pearl Harbor and in saline stream environments 

on O‘ahu’s south shore (for example, in Mākua Stream at 15–43 ‰ and in Niu Stream at 

34 ‰) (Englund et al., 2000a, 2000b). Its origin is unknown; the first date of collection is 1998. 

Chironomidae 

Cricotopus bincinctus (Meigen, 1818) 

Introduced 

This is a widespread (Afrotropical and Holarctic) midge first collected in 1955 in Pearl Harbor; it is 

“now one of the most ubiquitous introduced aquatic insects in the Hawaiian archipelago,” being 

found from low to “nearly pristine” high elevations (>1,220 m) (Englund, 2000a). It occurs in brackish 

waters in Pearl Harbor streams up to 9 ‰ (Englund et al., 2000b). 

Goeldichironomus holoprasinus (Goeldi, 1905) Introduced 

Beardsley (1970) reported this fly from a wide variety of aquatic situations, including “moats, ponds, 

(and) temporary water receptacles.” Beardsley (1970) noted that it was “common in Honolulu and 

has been reared from fish ponds on windward O‘ahu.” The first specimens are from 1969. Beardsley 

et al. (1999) add a new record from Pālā‘au State Park, at 1500 feet, on Moloka‘i, collected in 1994. 

It was described from Brazil and known from the United States. It is also found in brackish ponds 

(F.G. Howarth, pers. comm., 2005). For biogeographic analysis purposes, based upon its overall distribution, 

we treat this species as eastern American. 

Telmatogeton japonicus Tokunaga, 1933 

Introduced 

This Japanese marine fly was listed by Hardy (1960) as an immigrant. Wirth (1947b) reported it from 

Hilo in 1946 “scampering over wave-drenched boulders on bay-front at park” and that it occurred 

on “boulders on the bay-front in a limited area near the outlet of a large storm-sewer, with a heavy 

growth of the algae Ulva sp. and Enteromorpha sp. indicating that the water was of considerably 

lower salinity than pure sea water.” Wirth noted that Tokunaga also found it associated with these 

same algae in Japan. Whereas the other marine midge in Hawai‘i, Telmatogeton pacificus Tokunaga, 

1935, is widespread through the Islands (Nishida, 2002), T. japonicus appears to be restricted to the 

island of Hawai‘i, suggesting a human-mediated introduction. 

Dolichopodidae 

Dolichopus exsul Aldrich, 1922 

Introduced 

A West Indian (Caribbean) fly first found in 1930 in O‘ahu, and found by Englund et al. (2000b) in 

Pearl Harbor in hypersaline environments of Mākaha Stream (38 ‰) and along the concrete channel 

at the mouth of Wai‘alaenui Stream (32 ‰). 

Medetera grisescens Meijere, 1916 

Introduced 

This fly, thought to be native to Asia, the South Pacific (including Australasia), and the Indian Ocean, 

was first collected in Honolulu in 1914 (Englund et al., 2000a). It was collected in 1998 in Hālawa 

Stream flowing into Pearl Harbor in 32–37 ‰. 

Syntormon flexible Becker, 1922 

Introduced 

A long-legged fly native to the western and southern Pacific, it was first collected in 1917 in Mānā, 

Kaua‘i (Englund et al., 2000a). It occurs in brackish waters in streams flowing into Pearl Harbor

122 


(Englund et al., 2000a), and in other streams on the south shore of O‘ahu in salinities up to 37 ‰ 

(Englund et al., 2000b). 

Thinophilus hardyi Grootaert & Evenhuis, 1997 Cryptogenic 

This fly was described from O‘ahu based upon 1996 collections. Grootaert & Evenhuis (1997) suggested 

that it was “not a recent introduction, but that due to its small size it was overlooked until 

now.” Subsequently, Englund et al. (2000a) proposed that it was introduced, and native to Austral - 

asia. Further research since Englund shows that T. hardyi may be cryptogenic. Since 1996, the genus 

has been collected extensively in the Indo-Pacific, but T. hardyi has not been found, suggesting it 

may be a Hawaiian endemic (N.L. Evenhuis, pers. comm., February 2007). Specimens of a Thino - 

philus from the Galapagos were identified by Bickel & Sinclair (1997) as T. hardyi, but comparison 

of the male genitalia shows they are a distinct species, and the Galapagos population may be undescribed 

(N.L. Evenhuis, pers. comm., 2007). 

The habitat is “marshy sandplate on top of a porous lava bench protected by a rocky shore from 

the sea. The sand was covered with brown patches of diatomacea. Sea water welled up through the 

lava bench from time to time. The Thinophilus were mainly active on the wet sand around the 

upwelling sea water” (Grootaert & Evenhuis, 1997). Englund et al. (2000b) found this fly to be moderately 

common at a number of south shore O‘ahu stations in salinities ranging from 15 to 43 ‰. 

Psychodidae 

Psychoda salicornia Quate, 1954 

Introduced 

Quate (1954), in describing this fly from San Francisco and Tomales Bays, California, also noted 

specimens collected in light traps in Honolulu, O‘ahu, in 1945 and 1946. In California this species 

is found on the stems and branches of the pickleweed, Salicornia. Hardy (1960) added no new 

records. While there are no further reports of this species in Hawai‘i, we retain it on the list, since 

psychodids are rarely if ever identified to species on the islands, and it may thus still be present. 

Englund et al. (2000a), for example, record a Psychoda sp. from brackish water sties at ‘Aiea 

Stream, E‘o Stream, and Hālawa Stream. 

Ephydridae 

Brachydeutera ibari Ninomya, 1930 

Introduced 

A probable Asian species first collected at Beck’s Cove, Kāho‘olawe Island in 1980 (Evenhuis, 

1987). Specimens were found in estuarine water in E‘o Stream on barnacles at a golf course bridge 

in 1998 (Englund et al., 2000b). 

Ceropsilopa coquilletti Cresson, 1922 

Introduced 

A shore fly first collected on O‘ahu in 1946 and now found in streams in Pearl Harbor (Englund 

2000a) and in streams on the O‘ahu south shore in salinities from 15 to 43 ‰ (Englund et al. 2000a, 

2000b). Described from California, it is considered Nearctic (Englund et al., 2000a). 

Clasiopella uncinata Hendel, 1914 

Introduced 

A shore fly native to Taiwan and Australasia and first found on O‘ahu in 1946. It occurs in shoreline 

habitats in Pearl Harbor, including among mangroves, and in saline streams on the O‘ahu south shore 

on a variety of soft and hard substrates in salinities of up to 39 ‰ (Englund et al., 2000a, 2000b). 

Discocerina mera Cresson, 1939 

Introduced 

This shore fly was first collected in 1948 on O‘ahu. It is strongly associated with saline and estuarine 

habitats on O‘ahu, being found in saline streams on the south shore (up to 39 ‰) and in a variety 

of habitats along the Pearl Harbor shoreline (Englund et al., 2000a, 2000b). It is native to the 

western and/or southern Pacific Ocean. 

Donaceus nigronotatus Cresson, 1943 

Introduced 

Described from Taiwan, this fly was found on O‘ahu in 1958, and now occurs on soft and hard bottoms 

along the Pearl Harbor shore and in saline streams on the O‘ahu south shore, in salinities up to 

39 ‰ (Englund et al., 2000a, 2000b).


123 

Ephydra gracilis Packard, 1871 

Introduced 

= Ephydra cinerea Jones, 1906 

In what would appear to be a classic case of a population explosion by a new invader followed by a 

nearly complete collapse, this Western North America brine fly was found to be breeding by the millions 

in July 1946 in saltwater ponds opposite the Moanalua Gardens, the water in the ponds having 

become increasingly saline as dredging operations had closed the inlet from Ke‘ehi Lagoon (Wirth, 

1947a). Earlier in the year specimens had been found at Hickam Field (April in a light trap) and on 

Sand Island, opposite Honolulu (May). In August 1946 they were found at Iroquois Point, near Pearl 

Harbor, abundant in salt marsh pools. Wirth noted that “the close proximity of favorable breeding 

habitats at California seaplane bases, the large numbers of adults produced at times, their habit of 

swarming into moving vehicles (where they are a pest, as in trains crossing Great Salt Lake), and 

finally the proximity of receptive habitats near the O‘ahu seaplane bases, together develop conditions 

favorable for insect ‘transplantation’.” Wirth thus concluded that these flies were introduced by 

seaplanes from California to O‘ahu seaplane bases. 

Curiously, this fly was not noted again in the islands for the next 50 years. It was rediscovered 

in low numbers in 1998 in Mākaha Stream and Kuapā Pond in a sampling program focused on 

detecting aquatic invasions (Englund et al., 2000b). Habitat destruction may have played a role in 

the demise of the original populations around Hickam Field. 

Ephydra milbrae Jones, 1906 

Introduced 

The second species of brine fly to be introduced to the islands, E. milbrae was first collected in the 

Ala Wai Canal in 1950 (Englund et al., 2000a). It is native to western North America. It occurs on 

the Pearl Harbor shoreline in salinities of up to 44 ‰, and in O‘ahu streams up to 43 ‰ (Englund et 

al., 2000a, 2000b). 

Hecamede granifera Thomson, 1869 

Introduced 

A shore fly collected in 1923 from Mokapu, O‘ahu, and native to the western and/or southern 

Pacific. It is widespread on Pearl Harbor “seashore habitats” and in saline streams (to 43 ‰) on 

Oahu’s south shore ((Englund et al., 2000a, 2000b). 

Mosillus tibialis Cresson, 1916 

Introduced 

A North American shore fly collected at Mapulehu, Moloka‘i, in 1944, and at Waipahu, O‘ahu, in 

1958. In 1998 a specimen was collected over mud and rocks in brackish water of the E‘o canal; it 

has also been collected in the saline streams of Oahu’s south shore (Englund et al., 2000a, 2000b). 

Paratissa pollinosa (Williston, 1896) 

Introduced 

= Paratissa semilutea of authors 

The first records of this Caribbean-South American shore fly are from intertidal beach rocks at 

Lanikai, O‘ahu, in 1945 (Adachi, 1952a; Hardy & Delfinado, 1980, as P. semilutea); Mathis, 1993). 

Additional collections are those from Pearl Harbor in 1998 (Englund et al., 2000a). Hardy & 

Delfinado (1980) note that “Adults have been found mostly on dried seaweed washed up onto the 

each. Adults can often be seen crawling out from under seaweed piles when the seaweed has been 

disturbed.” 

Placopsidella marquesana (Malloch, 1933) 

Introduced 

= Placopsidella cynocephala Kertesz 

A shore fly (described from New Guinea) with a broad Indo-Pacific distribution considered to be 

introduced to the islands and first found on rocks in 1951 in Hanauma Bay, O‘ahu (Adachi, 1952b). 

It remains common and widespread in Pearl Harbor estuarine environments and in highly saline 

O‘ahu south shore streams (up to 42 ‰) (Englund et al., 2000a, 2000b). It is common in seaweed 

and debris on beaches (Englund, 2000b). It is also known from Kaua‘i, Maui, and Hawai‘i.

124 


Psilopa girschneri Roder, 1889 

Introduced 

A broadly distributed Northern Hemisphere (North America-Europe-Asian) shore fly first reported 

in 1952 from O‘ahu. A saline environment species, it is common in marine shorelines of Pearl 

Harbor and on Oahu’s south shore in salinities up to 43 ‰ (Englund et al., 2000a, 2000b). 

Scatella stagnalis (Fallen, 1913) 

Introduced 

Another broadly distributed northern hemisphere (North America-Europe-Asian) shore fly first re - 

ported relatively late (1967) from O‘ahu, with a habitat similar to Psilopa girschneri (Englund et al., 

2000a, 2000b). Englund (2000b) notes that it “remains one of the most common aquatic fly species 

of O‘ahu.” Englund (2002) notes that a first-reported date of 1946 is in error. 

Hostis guamensis Cresson, 1945 

Introduced 

The type locality of this Australasian fly is a China Clipper: the type specimen was taken from the 

airplane when it landed on Guam, with previous ports in California (Alameda, in San Francisco 

Bay), Hawai‘i (Honolulu), and Midway (Hardy & Delfinado, 1980), who suggested that “it is possible 

that it got into the plane at Honolulu. The Honolulu airport is close to the seashore where this 

species is known to occur.” It is widespread throughout the “Afrotropical” region (Mathis, 1993) and 

appears to have been introduced to (at least) the Hawaiian Islands and to Madagascar in the Indian 

Ocean. 

Adachi (1952a) was the first to report it, from specimens collected in 1946 at ‘Ewa, O‘ahu, 

found resting on seaweed on the beach. It also occurs on Maui (Nishida, 2002). 

Collembola (springtails) 

Hypogastruridae 

Oudemansia esakii (Kinoshita, 1932) 

Cryptogenic 

Christiansen & Bellinger (1992) report this Japanese species from Hawai‘i, “found on intertidal 

zones and beaches.” We note it as an example of cryptogenic collembolans. It was collected on 

O‘ahu in 1939 (Ala Moana Canal, on intertidal mud), Laysan (1983, unexposed reef), and Pearl and 

Hermes Reef (1983, lagoon intertidal zone; 1983, raised coral beach). 

ACARINA (mites) 

Eldredge & Miller (1995) listed two nonindigenous species of mites as marine, based upon Nishida 

(1994). However, these are freshwater species. 

PYCNOGONIDA (sea spiders) 

Introduced Pycnogonida 


Ammothella pacifica 1930 SF,BW Indo-Pacific 

Tanystylum rehderi 2000 SF,BW Indo-Pacific 

Pigrogromitus timsanus 1930 SF,BW Unknown 

Endeis biseriata 1960 SF,BW Indo-Pacific 

Endeis nodosa 1924 SF,BW Indo-West Pacific 

Endeis procera 1996 SF,BW Indo-West Pacific 

Anoplodactylus arescus 1998 SF,BW Indo-Pacific 

Anoplodactylus “californicus“ 1937 SF,BW Unknown 

Anoplodactylus digitatus 2000 SF,BW Unknown 

Anoplodactylus erectus


125 

We treat all of the Hawaiian harbor-dwelling sea spiders here as introduced with shipping. For most 

of these species, Hawai‘i is far distant from their apparently natural patterns of occurrence, and no 

shallow-water pycnogonids have been recorded from any high seas drifting material that would suggest 

that they may have naturally drifted into Hawaiian ports and harbors. A number of species 

appear to have arrived in the last half of the twentieth century since Edmondson’s and Hilton’s studies. 

Ammotheidae 

Ammothella pacifica Hilton, 1942 

Introduced 

Hilton (1942a) described this species from the Honolulu Aquarium (now Waikīkī Aquarium) on 

O‘ahu based upon specimens collected in 1930. It has subsequently been reported from the Indo- 

Pacific, including the Caroline Islands, Grand Comoro Island, and Madagascar (Stock, 1968). 

Bishop Museum holdings include specimens collected in 1948 from Honolulu Harbor. Coles et al. 

(2002a) report it from Kāne‘ohe Bay based upon 2000 collections. It is a probable ship-mediated 

introduction. 

Tanystylum rehderi Child, 1970 

Introduced 

This Indo-Pacific sea spider was first recorded in 2000 outside of Kāne‘ohe Bay at Moku Manu islet 

(Coles et al., 2002a), as identified by C.A. Child. There are scattered records throughout the South 

Pacific and Indian Oceans (Arango, 2003, who reports the first record from Australia). It occurs in 

Guam (Child, 1991), where it may also be introduced. We consider it introduced in vessel fouling or 

in ballast water. 

Callipallenidae 

Pigrogromitus timsanus Calman, 1927 

Introduced 

= Clotenopsa prima Hilton, 1942a, described from Hawai‘i, fide Stock, 1968. 

This tramp, ship-dispersed pycnogonid of unknown origin was first described from the Suez Canal, 

and later recorded from the Indian River, Florida (a canal-like environment), the saltwater locks of 

the Panama Canal, and scattered locations in the Indo-Pacific (C.A. Child, in litt. to L.G. Eldredge, 

1996 and to S.L. Coles, 1999). Child (1992) reported it from various stations in the western Gulf of 

Mexico, around Vera Cruz and Yucatan in Mexico. Moazzam (1987) extended it into the Arabian 

Sea. Arango (2003) describes it as “a pantropical-temperate species frequently collected in shallow 

habitats (in the) Indo-Pacific, Caribbean and Mediterranean.” Allan Child refers to it as the “canal 

pycnogonid” [Child (1979) and C.A. Child in litt. to L.G. Eldredge, 1996]. 

It was first collected at Black Point, O‘ahu in 1930 and redescribed as a new species, Clo - 

tenopsa prima by Hilton (1942a). Coles et al. (1997, 1999a) report it from fouling communities in 

Pearl Harbor based upon 1996 collections, and it also occurs in Ala Wai Harbor and Kewalo Basin, 

based upon 1998 material (Coles et al., 1999b). It was found in 1999 in Kāne‘ohe Bay (Coles et al., 

2002a). We consider it introduced to Hawai‘i in vessel fouling or in ships’ ballast water. 

Endeidae 

Endeis biseriata Stock,1968 

Introduced 

Stock described this species from New Guinea, Indonesia, and India; along with a single outlying 

record from Coconut Island, Kāne‘ohe Bay of one female collected in 1960 (Stock, 1968). Stock 

(1970) then extended the distribution well to the east, to the Gulf of Aqaba from the “piers at Eilat”, 

based upon specimens collected in 1970. Child (1996) summarized the known distribution, which 

by then included records from the Red Sea and Madagascar to Indonesia, the Philippines, and 

Australia, with continuing outlying records in Hawai‘i and Brazil. 

Endeis nodosa Hilton, 1942 

Introduced 

William A.Hilton described this species based upon specimens collected in 1924 and again in 1927 

in Kāne‘ohe Bay. Evans et al. (1974) report it from Pearl Harbor, although it was not recollected in

126 


1996 surveys (Coles et al., 1997). Child (1982) reported an additional specimen collected in 1947 in 

Honolulu Harbor “on boat hawser” (a towing or mooring rope). It remained unknown from elsewhere 

until it was collected in 1969 at Enewetak Atoll in the Marshall Islands, on pier pilings and 

on rocks in a lagoon (Child, 1982). Child commented that it “is probably not rare but only rarely 

reported because its habitats are seldom sampled.” Nakamura & Child (1988a) then reported it from 

Naha Harbor, Okinawa, Japan, based upon a specimen collected in 1987. We consider it unlikely to 

be native to the Hawaiian Islands. 

Endeis procera (Loman, 1908) 

Introduced 

This Indo-West Pacific species was reported from Pearl Harbor by Coles et al. (1997) based upon 

collections made in 1996. Stock (1954) noted it from the Kei Islands in sand and shells at about 50 

m depth. 

Phoxichilidiidae 

Anoplodactylus arescus Marcus, 1959 

Introduced 

This species has been recorded from the Atlantic Ocean (Brazil) but is more widespread throughout 

the Indo-Pacific [East Africa (Tanzania), Madagascar, Red Sea (the type locality), Philippines, and 

American Samoa] (Nakamura & Child, 1988b; C.A. Child in litt. to S. Coles, 1999). It was first 

found in Hawai‘i in 1998 at Barbers Point Harbor on pier pilings (Coles et al., 1999b) and again in 

2000 in Kāne‘ohe Bay (Coles et al., 2002a). Identifications were made by C.A. Child. It is often 

found in sandy sediments. It may have been long overlooked in the islands and been transported here 

in sand ballast; alternatively, more modern-day transport with ballast water is possible. It also would 

appear to be an introduction to Brazil. 

Anoplodactylus “californicus” Hall, 1912 

Introduced 

= Anoplodactylus portus Calman, 1927, fide Child, 1987 

= Anoplodactylus projectus Hilton, 1942a, fide Child, 1987 (see Child, 1975, p. 201, for additional basis of 

synonymy) 

This sea spider was described as a new species, Anoplodactylus projectus Hilton, 1942a, based upon 

a specimen collected in 1938 in Pearl Harbor “from bottom of ship.” Earliest Bishop Museum material 

from Pearl Harbor dates from 1937 (identified by J.H. Stock in 1967) with regular collections 

being made thereafter. Grovhoug & Rastetter (1980) reported it (as A. portus) in Kāne‘ohe Bay and 

Pearl Harbor (1976–1977 collections). Coles et al. (1997) reports it from Pearl Harbor. It also is 

found (1997–1998) in Honolulu Harbor, Ala Wai Harbor, Barbers Point Harbor, and Kewalo Basin 

(Coles et al., 1999b) and in Kāne‘ohe Bay (2000 collections; Coles et al., 2002a). 

Child (1987) synonymized the global species A. portus with A. californicus. As such, it is 

reported as a pantemperate-pantropical species from many parts of the world. Hall’s original concept 

of A. californicus was a species that in southern California lived among the bases of the surfgrass 

Phyllospadix on exposed rocky intertidal shores at extreme low tide (Hall, 1913). This is not a habitat 

characterized by cosmopolitan taxa, nor a habitat that either receives or donates invasions. It 

seems probable that the concept of A. californicus involves more than one species, and we thus place 

the species name in quotation marks here and recognize it as a nonnative taxon of unknown origin. 

Anoplodactylus digitatus (Böhm, 1879) 

Introduced 

Coles et al. (2002a) newly report this sea spider from the islands based upon material collected in 

2000 in Kāne‘ohe Bay and identified by C.A. Child. Arango (2003), in recording it for the first time 

for Australia, noted that it was widely known from the Indo-West Pacific, the West Indies (Carib- 

bean), and the Mediterranean Sea. We suggest that is not likely native to the Hawaiian Islands.


127 

Anoplodactylus erectus Cole, 1904 

Introduced 

A species apparently native to the eastern Pacific Ocean (from British Columbia, Canada, to 

Colombia), but also known in the South Pacific from French Polynesia (Tuamotu Islands) and 

American Samoa (Pago Pago Harbor), as well as in the North Pacific in Korea (Child, 1979; 

Nakamura & Child, 1988b). Stock (1975) reported it on fouling panels in shallow water in Val - 

paraiso and Punta Arenas, Chile. 

Hilton (1942b) noted it from “Honolulu” without date or further data. We suggest that it has 

been introduced to at least Korea and the Hawaiian Islands with vessel traffic. 

Anoplodactylus eroticus Stock, 1968 

Deleted 

Stock (1968) described this sea spider from the Gulf of Manaar in India, and from Honolulu Harbor, 

the latter based upon a specimen collected in 1945. It has not reappeared in recent collections of sea 

spiders on O‘ahu. 

Anoplodactylus marshallensis Child, 1982 

Introduced 

Coles et al. (2002a) newly report this sea spider from the islands based upon material collected in 

2000 in Kāne‘ohe Bay and identified by C.A. Child. It was previously known from collections made 

in 1969 from pier fouling on Enewetak Island, Enewetak Atoll, Marshall Islands (Child, 1982). We 

consider it a likely candidate for fouling or ballast water transport. 

Anoplodactylus pycnosoma (Helfer, 1938) 

Introduced 

Coles et al. (1997) report this species from Pearl Harbor (1996), and from Moku Manu islet outside 

of Kāne‘ohe Bay (2000) (Coles et al. 2002a), as identified by C.A. Child. Anoplodactylus pycnosoma 

is a widely distributed Indo-Pacific species, including records in the North Pacific Ocean in 

Korea, Japan, and Guam (Child, 1991, Stock, 1994). Staples (1997) lists records from Western 

Australia and from southern Australia (in Port Philip Bay, a site of many invasions). 

PHORONIDA (phoronids) 

Introduced Phoronida 


Phoronis hippocrepia 1976 SF North/South Atlantic 

Phoronis hippocrepia Wright, 1856 

Introduced 

What appears to be a fouling species of Atlantic phoronid has become established in two locations 

in the Pacific Ocean. The first records of Phoronis hippocrepia are from 1976 in Kāne‘ohe Bay 

(Emig, 1977; Emig & Bailey-Brock, 1987; the latter paper provides the date of 1976). Emig (1982) 

implies a date of 1975 (figure 1A, species 2, location 42), but the date of the paper cited is 1977, not 

1975. Phoronis hippocrepia likely occurred in Hawai‘i well before this date, and it should be expected 

in Pearl Harbor, as well as in other Pacific harbors. 

Phoronis hippocrepia is widely known from the North and South Atlantic Oceans but has been 

recorded only from two stations in the Pacific Ocean, first in Sydney Harbour (Port Jackson), 

Australia in 1893, then in 1976 in Kāne‘ohe Bay. It has also been introduced into the Miraflores 

Locks of the Panama Canal (Emig, 1982; Bailey-Brock & Emig, 2000), where it was collected probably 

in 1972 (see Jones & Dawson, 1973, biotic surveys of that date in the Miraflores Locks). 

This moderate-sized (4 cm) phoronid is a “burrowing and encrusting species” (Bailey-Brock & 

Emig, 2000), occurring in benthic coral rock and burrowing into “oyster shells and barnacles 

attached to harbor pilings to depths of 30 cm below mean tide level” (Emig & Bailey-Brock, 1987, 

who present a figure of a phoronid in situ within an empty barnacle shell). Bailey-Brock & Emig

128 


(2000) specifically note its occurrence “with fouling communities” on Lilipuna Pier, and “in coral 

rock from fringing reefs in south and north regions of” Kāne‘ohe Bay. 

It seems probable that over the course of the past 100 years this Atlantic species has been introduced 

by ship fouling to the Pacific Ocean, and its appearance in the Panama Canal is particularly 

compelling evidence for ship-mediated transport. Colonization is doubtless facilitated by asexual 

reproduction (transverse fission). 

KAMPTOZOA (Entoprocta) 

Introduced Kamptozoa 


Barentsia sp. 1966 SF, BW Unknown 

Barentsiidae 

Barentsia sp. 

Introduced 

We regard this Barentsia, reported as Barentsia gracilis from Kāne‘ohe Bay fouling communities 

(presumably 1966 collections) by Soule & Soule (1968) and from Pearl Harbor and Kāne‘ohe Bay 

fouling in 1976–1977 by Grovhoug & Rastetter (1980) as an introduced species. It is doubtless still 

present but largely overlooked as part of the microfouling fauna. Hawaiian populations should be 

compared to Barentsia benedeni (Foettinger, 1887). 

BRYOZOA (Ectoprocta) (bryozoans) 

Introduced Bryozoa 


Amathia distans 1935 SF Unknown 

Bowerbankia sp., cf. gracilis 1966 SF Unknown 

Bowerbankia sp., cf. B. imbricata 1966 SF Unknown 

Zoobotryon verticillatum


129 

Ctenostomata 

vesiculariidae 

Amathia distans Busk, 1886 

Introduced 

This well-known fouling bryozoan is now so widespread that its origin remains unknown (Cranfield 

et al., 1998, suggest that it is native to the Atlantic coast of the Americas). It occurs in the North and 

South Pacific and North and South Atlantic Oceans. Gordon & Mawatari (1992) and Keough & Ross 

(1999) consider it introduced to New Zealand and Australia, respectively. 

First records for Hawai‘i are 1935 in Kāne‘ohe Bay by Edmondson & Ingram (1939, as Ama - 

thia sp.). Bishop Museum material for Pearl Harbor commences in 1948 and for Honolulu Harbor 

in 1946. Soule & Soule (1968; 1966 collections) noted it from Coconut Island in Kāne‘ohe Bay. 

Long (1974; 1968–1972 collections) found it on fouling panels just offshore of O‘ahu. DeFelice et 

al. (1998) report it from Midway Atoll. Coles et al. (2006) report it from many sites on Kaua‘i, 

O‘ahu, Moloka‘i, Maui, and Hawai‘i based upon collections from 1996 to 2003. Hoover (1998, 

2006) presents a color photograph of specimens in Pearl Harbor. From October 2002 to February 

2003 this bryozoan was observed growing in massive quantities, described as “enormous tumbleweed-like 

colonies” in the brackish water Ala Wai Canal in Waikīkī (C. Zabin, pers. comm., January 

2004). We consider this a ship-fouling introduction that was certainly in Hawai‘i many years before 

its first report. 

Bowerbankia sp., cf. gracilis Leidy, 1855 

Introduced 

This is another now-cosmopolitan ctenostome. While often suggested as being native to eastern 

North America (northwestern Atlantic Ocean)—it was first described from Rhode Island—this 

species is now too widespread to yet suggest its origin, prior to molecular genetic analysis. Gordon 

& Mawatari (1992) provide a general global summary; they consider it introduced to New Zealand. 

While no doubt long present in the islands, first records are only from 1966, based upon collections 

in Kāne‘ohe Bay and the Ala Wai Canal (Soule & Soule, 1968). 

Bowerbankia collected on panels placed just offshore of O‘ahu (1968–1972 collections; Long, 

1974) and reported in Pearl Harbor and in Kāne‘ohe Bay based upon 1976–1977 collections 

(Grovhoug & Rastetter, 1980) are referable to either this species or the next. 

Bowerbankia sp., cf. imbricata (Adams, 1798) 

Introduced 

Soule & Soule (1968) report this species from Kāne‘ohe Bay and Ala Wai Canal, based upon 1966 

collections. It too has a widespread, global distribution (Gordon & Mawatari, 1992). Its origin is 

unknown. 

Zoobotryon verticillatum (Delle Chiaje, 1828) 

Introduced 

= Zoobotryon pellucidum Ehrenberg, 1831 

= Zoobotryon pellucidus of authors (misspelling) 

A bryozoan capable of forming massive colonies in fouling communities, it is either native to the 

Mediterranean (as suggested by Cranfield et al.,1998) or arrived there as an invader on ship bottoms 

from elsewhere by the 18th or early 19th century. We regard its origin as unknown. Robertson (1921) 

noted that she had received specimens from “Honolulu, Hawaiian Islands”; this appears to be the 

first mention of the species from Hawai‘i in the literature. The report by Cohen & Carlton (1995) of 

a Hawaiian record in Robertson (1905) is in error. 

Bishop Museum material dates from 1921 for Pearl Harbor and 1946 for Honolulu Harbor. Coles 

et al. (1999b) also report it form Ala Wai Harbor, Barbers Point Harbor, and Kewalo Basin, based upon 

1998 collections; Coles et al. 2004 report it from Mā‘alea and Kahului Harbors on Maui, based on 2003 

collections. DeFelice et al. (1998) report it from Midway Atoll. Edmondson (1933) reported it (as Z. 

pellucidus) associated with Bugula in fouling communities on O‘ahu. Edmondson & Ingram (1939) 

note that in Kāne‘ohe Bay a colony grew to 50 mm in height in 56 days, and that it was also occasionally 

found on boat bottoms in Pearl Harbor. Soule & Soule (1968) report on collections from Kāne‘ohe 

Bay in 1966, noting that it occurred there “hanging in great tangled streamers.” 

Godwin (2003b) noted that Z. verticillatum was present on a barge towed from San Diego to 

O‘ahu in 1999.

130 


Cheilostomata 

Anasca 

Aeteidae 

Aetea truncata (Landsborough, 1852) 

Introduced 

= Aetea recta of Soule et al., 1987, p. 97 

A tiny creeping stolonate bryozoan which now occurs in all oceans (Osburn, 1950; Winston, 1982), 

and whose origin remains unknown. The earliest record is that of Edmondson & Ingram (1939) 

based on specimens collected in 1935 in Kāne‘ohe Bay. Soule & Soule (1968) continue to record it 

from Kāne‘ohe Bay, based upon 1966 collections, and also report specimens from “off Oahu” at 9.1 

and 27.4 m. Coles et al. (1997, 1999a) report it from Pearl Harbor (1996), and Coles et al. (1999b) 

report it from Honolulu Harbor (1997) and Kewalo Basin (1998). Bailey-Brock (1989) recorded it 

on PVC settlement plates on an artificial reef in open waters at 20 m, 2.4 km off the southeast coast 

of O‘ahu. 

Aetea anguina (Linnaeus, 1758) 

Introduced 

Another now-cosmopolitan species of Aetea, which may involve a species complex (Keough & 

Ross, 1999, who also consider it introduced to Australia). Reported in 1997 from Honolulu Harbor 

by Coles et al. (1999b). 

Bugulidae 

Bugula dentata (Lamouroux, 1816) 

Introduced 

Zabin (1999) reports this blue-green or turquoise species from 1997–1998 collections from Honolulu 

Harbor, Ala Wai Yacht Harbor, and Barbers Point, based upon surveys of O‘ahu harbors (Coles et 

al., 1999b). It also occurs at Koko Marina (Coles et al. 2002b) and in harbors at Kaua‘i, Moloka‘i, 

Maui, and Hawai‘i (Coles et al., 2004). 

It is a species of the Indo-Pacific Ocean (Zabin, 1999) and is also known from Japan (R. Wool - 

la cott, pers. comm., October 1996). Hoover (1998, 2006) presents a color photograph from 3 m at 

Magic Island, O‘ahu (identified as B. stolonifera, but in fact B. dentata; C. Zabin, pers. comm., 

2000). Hoover’s photograph was taken in January 1997 (J. Hoover, pers. comm., January 2004, who 

notes that at that time “they were all over the place at Magic Island boat channel.”) 

Bugula “neritina” (Linnaeus, 1758) 

Introduced 

One of the world’s most commonly recognized fouling organisms, now too widespread to know its 

origin without extensive global genetic analyses. Bugula “neritina” is a species complex (for example, 

Davidson & Haygood, 1999; McGovern & Hellberg, 2003), likely represented by regional 

endemics along with a now-global port-dwelling, ship-dispersed fouling taxon. Earliest Bishop 

Museum specimens are from 1921 in Pearl Harbor. Edmondson (1933) reported it to be common in 

O‘ahu harbors and bays. Edmondson & Ingram (1939) provide growth data in 1935 and later years 

for Kāne‘ohe Bay. Soule & Soule (1968) report it from Kāne‘ohe Bay (1966) and Ala Wai Yacht 

Basin (1966) and note material from Honolulu Harbor as well in Bishop Museum collections (the 

earliest Honolulu specimens are from 1940). Edmondson (1944a) further reported it from Pearl 

Harbor, and Long (1974) found it both on Pearl Harbor fouling panels and in panels just offshore of 

O‘ahu between 1968 and 1972. It remains abundant in Pearl Harbor (Coles et al., 1997, 1999a) and 

is also still common in Honolulu Harbor, Ala Wai Harbor, and Kewalo Basin (Coles et al., 1999b). 

It was also found in 2002–2003 collections in harbors on Kaua‘i and Maui (Coles et al., 2004). 

Mackie et al. (2006) found that Hawaiian specimens from Ford Island (Pearl Harbor) were genetically 

similar to a clade now widespread through the Pacific and Atlantic Oceans. 

Ingram (1939) noted that the snail Atys semistriata deposits its egg masses on (and may feed 

upon) B. neritina in Kāne‘ohe Bay. Entire populations of B. neritina can be grazed down on marina 

floats in a matter of days by Tilapia (R. Woollacott, pers. comm., October 1996).


131 

Bugula minima Norman, 1909 

Introduced 

Zabin (1999) reported (as B. robusta MacGillivrary, 1869) this widespread Bugula from Pearl 

Harbor (1996), Honolulu Harbor (1997), and Barbers Point (1998) based upon the surveys described 

by Coles et al. (1999a, 1999b). It was present in 1995 in Kewalo Basin and Kāne‘ohe Bay (Zabin, 

1999). It also occurs in Nawiliwili Harbor, Kaua‘i, and Kuhului Harbor, Maui (Coles et al., 2004). 

Bugula minima was described from the Eastern North Atlantic, and subsequently widely reported 

from a number of Atlantic and Pacific locations. Hawaiian populations, formerly thought to be B. 

robusta, are now considered to be B. minima (Winston & Woolaccott, 2008) 

Bugula stolonifera Ryland, 1960 

Introduced 

= Bugula californica of authors, including Hawaiian literature 

Another Bugula found in most seas and whose origin remains uncertain pending global analysis of 

distribution and genetic patterns. Cranfield et al. (1998) and Keough & Ross (1999) consider it introduced 

to New Zealand and Australia, respectively. 

Earliest Bishop Museum material is from 1935 (Kāne‘ohe Bay), 1940 (Pearl Harbor), and 1946 

(Honolulu Harbor). Soule & Soule (1968, as B. californica) reported it from Kāne‘ohe Bay and Ala 

Wai Yacht Basin (1966 collections). Long (1974, as B. californica) found it on panels offshore of 

O‘ahu (1968–1972). Coles et al. (1997, 1999a, 1999b) report it from Pearl Harbor, Barbers Point 

Harbor, Ala Wai Yacht Harbor, and Kewalo Basin. Godwin (2003b) noted that B. stolonifera was 

present on a barge towed from San Diego to O‘ahu in 1999. 

Caulibugula dendrograpta (Waters, 1913) 

Introduced 

Zabin (1999) reports this Indo-Pacific species from Honolulu Harbor and Barbers Point (1997– 

1998). Coles et al. (2000a) report it also from Kāne‘ohe Bay. This and the following species represent 

the first Caulibugula reported from the Islands. 

Caulibugula caliculata (Levinsen, 1909) 

Introduced 

This western and Indo-Pacific bryozoan was first collected in 1997 in Honolulu Harbor (Zabin, 

1999). Previous records are for Hong Kong and the Philippines. 

Chorizoporidae 

Synnotum aegyptiacum (Audouin, 1826) 

Introduced 

Soule et al. (1987, p. 119) report this widespread warm-water species from O‘ahu, based upon collections 

presumably in 1966 (see Soule & Soule,1968), noting that it occurs “mixed with hydroids, 

bryozoans, and algae in fouling communities.” Coles et al. (1999b) report it from Honolulu Harbor, 

Ala Wai Harbor, and Barbers Point Harbor, based upon 1997–1998 collections. It likely occurs in 

Pearl Harbor as well. 

Synnotum aegyptiacum, originally described from the Red Sea, is known in the Eastern Pacific 

from southern California to Peru, in the Western Atlantic from Cape Hatteras to Brazil, including the 

Caribbean and the Gulf of Mexico (Osburn, 1950; Winston, 1982), as well as in the Indo-Pacific and 

Australia. Its original home remains unknown. 

Scrupocellariidae 

Caberea boryi (Audouin, 1826) 

Introduced 

Soule et al. (1987, p. 98, as Caberia) report this species as being epizoic on the bryozoans Amathia 

and Zoobotryon based upon collections in 1966 in Kāne‘ohe Bay [although this species is not mentioned 

in Soule & Soule (1968)]. It remains present in Kāne‘ohe Bay (Coles et al., 2002a). Coles et 

al. (2006) report it from Moloka‘i, based upon 2003 collections. It is now reported from most global 

shores, and its origin is at this time not known.

132 


Ascophora 

Cryptosulidae 

Cryptosula pallasiana (Moll, 1803) 

Introduced 

Soule et al. (1987, p. 98) record this common North Atlantic species from Kāne‘ohe Bay fouling 

communities in 1966, although it is not reported in their earlier fouling study (Soule & Soule, 1968). 

It occurs widely in other Pacific harbors and bays, including the North American Pacific coast, 

Japan, New Zealand and Australia (Cohen & Carlton, 1995). It is a probable global species complex. 

Saviignyellidae 

Savignyella lafontii (Audouin, 1826) 

Introduced 

= Catenaria lafontii in Edmondson & Ingram, 1939 

A global species of uncertain origin recorded from fouling communities in many parts of the world. 

First recorded in 1935 in Kāne‘ohe Bay (Edmondson & Ingram, 1939, as Catenaria lafontii; identified 

by R.C. Osburn). Soule & Soule (1968) record it again from Kāne‘ohe Bay (1966) and note 

Bishop Museum specimens from Pearl Harbor and Honolulu Harbor. Long (1974, collections from 

1968–1972) recorded it on panels set just offshore of O‘ahu. Coles et al. (1999b, 2002a, 2002b, and 

2004) report it from a number of locations on O‘ahu, Moloka‘i, and Maui. 

The photographs of this species in Hoover (1998, pp.87 and 89; 2006) in deeper waters off 

Maui are instead Vittaticella uberrima Harmer, 1957 (C. Zabin, pers. comm., January 2004). 

Escharellidae 

Hippopodina tahitiensis Leca & d’Hondt, 1993 Introduced 

= Hippopodina feegeensis of Hawaiian authors, not of Busk, 1884 

Tilbrook (2006) determined that the Hippopodina long reported from the Islands as H. feegeensis is 

H. tahitiensis, described relatively recently from French Polynesia. It was first collected in the 

Islands in 1948 in Honolulu Harbor (Bishop Museum collections; this material was noted by Soule 

& Soule, 1968, without date). It appears to have been next collected in 1966 in raft fouling in shallow 

water in Kāne‘ohe Bay (Soule & Soule, 1968). It was also collected in 1996 in Pearl Harbor (as 

Schizoporella sp. in Coles et al., 1997; specimens reexamined and identified by C. Zabin). It was 

observed in 1994 at the Waikīkī Yacht Club, and for the first time in 1994 at Kewalo Basin (R. 

Woollacott, pers. comm., October 1996). Coles et al. (1999b) report it from Honolulu Harbor, Ala 

Wai Harbor, and Kewalo Basin, based upon 1997 and 1998 collections. 

Schizoporellidae 

Schizoporella sp. 

Introduced 

One or more species of Schizoporella, none native, are present in Hawaiian fouling communities. 

Uncertainty has attended the identification of species in this genus since the earliest attempts: in 

1936, R.C. Osburn examined specimens for C.H. Edmondson collected in 1935–1936 from 

Kāne‘ohe Bay and reported them as “Schizoporella unicornis or a closely related species” (A. 

Wetmore in litt. to C.H. Edmondson, 12 September 1936; letter on file at Bishop Museum). 

Schizoporella unicornis (Johnston, 1847) has been reported by many workers since Edmondson 

& Ingram’s (1939) studies in Kāne‘ohe Bay (specimens collected since 1935). Soule & Soule (1968, 

based upon 1966 collections) record S. unicornis from Kāne‘ohe Bay and Ala Wai Yacht Basin and 

note additional Bishop Museum specimens from Pearl Harbor and Honolulu Harbor. Hurlbut 

(1991c) reported on settlement and juvenile survival in Pearl Harbor. Schizoporella errata (Waters, 

1878) is also reported from many stations in Hawaiian harbors and reefs (Coles et al., 1997, 1999a, 

1999b, 2002a, b, 2004, 2006). DeFelice et al. (1998) reported S. errata from Midway Atoll. Godwin 

(2003b) noted that S. unicornis was present on a barge towed from San Diego to O‘ahu in 1999. 

The identification of these and other Schizoporella species has often presented many challenges 

(Keough & Ross, 1999), and a thorough reexamination of Hawaiian material is required. Dick et al. 

(2005), for example, note that many records of S. unicornis from the north Pacific Ocean are attributable 

to S. japonica Ortmann, 1890.


133 

Hippothoidae 

Trypostega venusta (Norman, 1864) 

Introduced 

Soule et al. (1987, p. 98) record this cosmopolitan species (Osburn, 1952; Winston, 1982) from 

Kāne‘ohe Bay fouling communities in 1966, although it is not reported in their earlier fouling study 

(Soule & Soule, 1968). 

Watersiporidae 

Watersipora edmondsoni Soule & Soule, 1968 

Introduced 

This Watersipora was described as a new species from collections made in 1966 from the hull fouling 

of houseboats in the Ala Wai Yacht Harbor in Honolulu. Watersipora edmondsoni is known from 

Pearl Harbor, as well as from panels placed in waters just offshore (Long, 1974, Grovhoug, 1979, 

and Coles et al., 1997, 1999a), from Kāne‘ohe Bay [Dade & Honkalehto, 1986 (1983 collections)]; 

Coles et al., (2002a; Bishop Museum material since 1975); from Kaua‘i (Soule et al., 1987, page 

129, plate caption), from Ke‘ehi Lagoon (Coles et al., 1999b, 2002a (1997), and from Waikīkī, Koko 

Marina, and Hawai‘i Kai (Coles et al., 2000b), and from Molokai’s Kaunanakai Main Dock (Coles 

et al., 2004). Mackie et al. (2006) record W. edmondsoni from Waiahole Reef in Kāne‘ohe Bay, 

based upon collections made in 1999 (J. Mackie, pers. comm., January 2007). 

It remained known only from the Hawaiian Islands until reported from several Indo-Pacific 

locations by Winston & Heimberg (1986). Winston & Heimberg (1986) reported it from Sanur in 

Bali, based upon collections made in 1984 (J. Winston, pers. comm., January 2004; the date does not 

appear in the paper). They also interpreted specimens illustrated by Soule & Soule (1975) as falling 

within the concept of W. edmondsoni. These included material collected in 1898 in Torres Strait 

(between New Guinea and Northeast Australia) and in 1905 in Sri Lanka. It would thus appear that 

W. edmondsoni is Indo-Pacific in origin. Soule & Soule (1968) suggested that it was introduced 

between 1956 and 1966, based upon collections made by personnel from the Bishop Museum up 

until 1956 and the failure to collect this species prior to that. 

Dick (2006) reported this species as “Watersipora subovoidea sensu Harmer, 1957,” from rocky 

intertidal sites at Kapa‘a Beach Park, on the Big Island, and tentatively placed the Soules’s name in 

synonymy. It may well be that W. subovoidea, in part, captures W. edmondsoni, and may eventually 

be found to be a senior synonym. 

Watersipora arcuata (Banta, 1969) 

Introduced 

Coles et al. (1999b, 2004) report the fouling bryozoan Watersipora arcuata from Kewalo Basin 

(1998 collections) and from Kahului Harbor, Maui (2003 collections). Mackie et al. (2006) genetically 

confirmed W. arcuata from both Ala Wai Harbor and Hale‘iwa Harbor on O‘ahu; this material 

was collected in 2000 (J. Mackie, pers. comm., January 2007). Watersipora arcuata is native to the 

warm waters of the Eastern Pacific and has been introduced to Australia as well (Mackie et al., 

2006). 

Watersipora subtorquata (d’Orbigny, 1852) 

Introduced 

Soule et al. (1987, p. 129, figure caption) illustrated Watersipora “subovoidea” based upon 1966 

collections in Pearl Harbor and Ala Wai Marina. The caption description and the photograph of the 

aperture match W. subtorquata, a species believed to be native to the Caribbean and also introduced 

to Australia and New Zealand (Mackie et al., 2006).

134 


Echinodermata 

Introduced and Cryptogenic Echinodermata 


Ophiuroidea 


Ophiactis savignyi 1847–1849 SF Indo-Pacific 


Ophiactis modesta 1902 SF 

Ophiuroidea 

Ophiactidae 

Ophiactis savignyi (Muller & Troschel, 1842) 

Introduced 

= Ophiactis sexradia Grube, 1857, described from Hawai‘i, synonymy fide Clark, 1949 

This widespread ophiuroid is native to the Indo Pacific; its modern-day distribution has been strongly 

influenced by dispersal in ship fouling, and we suggest here that it was an early ship-borne introduction 

to the Islands. It was described as a new species, Ophiactis sexradia, from Honolulu by 

Grube (1857), a record repeated by a number of early authors thereafter (Clark, 1949). Grube attributed 

the specimens to “Dr. Ed. Lenz.” Lenz sailed in the Pacific aboard the Russian skiff Achta in 

1847–1849 (Lenz, 1863; von Goguslawski, 1884), but we have not yet determined which year he 

was in the Islands. Clark (1915) reported it specifically from Pearl Harbor but without further data. 

Earliest records for Hawai‘i in Bishop Museum collections are from 1924 (Kāne‘ohe Bay) and 

1929 (Pearl Harbor); the species still occurs in both locations (Coles et al., 1997, 2002a), and in all 

of Oahu’s commercial harbors and public marinas (Coles et al., 1999b). Ely (1942) noted that it was 

“common in Kaneohe Bay and Pearl Harbor, habitually concealed in the canals of sponges, or even 

in tangled masses of sea weed or crevices in dead coral blocks. it is a gregarious species, and many 

individuals may be found within a piece of sponge the size of a man’s hand.” MacKay (1945) noted 

the number as “50 to 75 in a piece of sponge the size of a man’s fist.” 

Edmondson (1933) noted that it was “very common” and that “Large numbers of specimens 

have been taken among the sponges common on piling and buoys in Kaneohe Bay and Pearl 

Harbor.” Clark (1949) noted that the Albatross collected it in 1902 in Pearl Harbor (O‘ahu), Puakō 

Bay (Hawai‘i), as well as at two deeper water stations between Maui and Lāna‘i, in ‘Au‘au Channel, 

in 51–79 m (this latter material bears re-examination, relative to the possibility of being a similarlooking 

but cryptic species). Coles et al. (2006) report it from reefs at Kaua‘i and Moloka‘i, and from 

har areas at Nawiliwili, Kaua‘i, Kaunakakai Main Dock, Moloka‘i, and Hilo Harbor, based upon 

2002–2003 collections. 

Rowe & Gates (1995) noted it was a tropical circumglobal species occurring from 0 to 550 m. 

Based upon genetic analyses Roy & Sponer (2002) concluded that this ophiuroid had been introduced 

to the Atlantic Ocean by shipping. Unpublished nested clade analyses further suggest that 

human-mediated dispersal is much more likely to have carried this species to distant outlier sites, 

such as South Africa, Panama, and the Hawaiian Islands (M. Roy, pers. comm., August 2008). Given 

the genetic evidence combined with its fouling habitat we treat Ophiactis savignyi as introduced. 

Ophiactis modesta Brock, 1888 

Cryptogenic 

In reporting this brittle star for the first time from Hawai‘i, Ely (1942) made the following interesting 

statement: “O. modesta has previously been taken at Misaki, Japan, Amboina, Thursday Island, 

and along the Australian coast from Darwin to Lagrange Bay. It is now recorded for the first time 

from Hawai‘i. H.L. Clark (personal letter) believes that, as both members of this genus are carried 

about on boat bottoms, their arrival here may have been fairly recent. It is quite difficult to say how 

recently this species may have arrived in Hawaii.” Ely noted that this species and O. savignyi were


135 

collected in almost identical situations (see above, at O. savignyi) in shallow water in Pearl Harbor 

and Kāne‘ohe Bay. The only Bishop Museum material is from 1938 collected in Pearl Harbor. Coles 

et al. (2002a) list it for Kāne‘ohe Bay, but Coles et al. (1997) do not report it further for Pearl Harbor. 

Austin H. Clark (1949) then reported on the presence of O. modesta in the islands as a deeper 

water species, noting a modest number of specimens collected by the Albatross in 1902 from off 

Moloka‘i in 146–406 m, from between Maui and Lāna‘i in 51–79 m, from between Hawai‘i and 

Maui in 90–322 m, and from off Maui in 82–95 m and in 104–124 m. Rowe & Gates (1995) report 

O. modesta from Australia with a depth range of 0–22 m and note a broad distribution in the tropical 

west Pacific Ocean, including West India and Pakistan. Price & Rowe (1996) add Sri Lanka to 

the known locations. Clark’s (1949) deeper water Hawaiian material could possibly bear re-examination 

relative to their assignment to O. modesta, given the shallow-water records that otherwise 

characterize this species in Pearl Harbor and Kāne‘ohe Bay, and in Australia as well. 

CHORDATA 

Introduced and Cryptogenic Tunicata (Ascidiacea) 



Didemnum candidum 1930 SF Northeast Atlantic 

Didemnum perlucidum 1999 SF Unknown 

Didemnum psammathodes 1998 SF Indo-West Pacific 

Diplosoma listerianum 1900 SF Unknown 

Lissoclinum fragile 1962 SF Unknown 

Ciona intestinalis 1933 SF North Atlantic 

Ecteinascidia imperfecta 1979 SF Western Pacific 

Corella minuta

136 


Tunicata (Urochordata): Ascidiacea (sea squirts) 

Didemnidae 

Didemnum candidum Savigny,1816 

Introduced 

This seasquirt, possibly of Eastern North Atlantic origin, was first found in the Hawaiian islands on 

Pearl and Hermes Reef, based upon collections by Paul S. Galtsoff in 1930 (Tokioka, 1967). 

Eldredge (1967) recorded it from Kāne‘ohe Bay, Ala Wai Yacht Harbor, and Pearl Harbor, based 

upon 1961–1963 collections. It was also collected off Barbers Point in 1962 at 40 m (Eldredge, 

1967). Additional records include further reports from Pearl Harbor (Long, 1974; 1968–1972 collections; 

Hurlbut, 1990, 1991a, 1991b), Kāne‘ohe Bay (Straughan, 1969b; 1967–1968 collections; Fitz - 

hardinge & Bailey-Brock, 1989; 1985 collections, colonizing artificial substrates in abundance), and 

Honolulu Harbor, Ke‘ehi Lagoon, Ala Wai Harbor, and Ke‘ehi Basin (Coles et al., 1999b, 1997– 

1998 collections), 1999–2000 collections in Kāne‘ohe Bay (Coles et al. 2002a), and 2001 collections 

off Waikīkī (Coles et al., 2002b). Bailey-Brock (1989) recorded it on PVC settlement plates on an 

artificial reef in open waters at 20 m, 2.4 km off the southeast coast of O‘ahu. Coles et al. (2006) 

report it from Kaua‘i and Moloka‘i and Maui,based upon 2003 collections, and from 2005 collections 

on Lāna‘i (Coles et al., 2005). Hurlbut (1990, 1991a, 1991b) studied the larval abundance, settlement, 

and juvenile mortality of this species in Pearl Harbor. 

Didemnum perlucidum Monniot, 1983 

Introduced 

Godwin & Lambert (2000) report this species from Ke‘ehi Lagoon on O‘ahu based upon collections 

from floating docks in 1999. This is a now widespread species. Although described from the 

Caribbean, “the origin is ... unknown. It is a fouling species that (has) colonized many harbors in the 

Atlantic (Brazil, West Indies, Senegal) as well as the Indian Ocean (Maldive Islands, Zanzibar) and 

the Pacific (New Caledonia, Polynesia) and now Indonesia” (Monniot & Monniot, 1996). It also 

occurs in the Gulf of Mexico on the Texas coast (Godwin & Lambert, 2000). 

It was found in 1999–2000 in Kāne‘ohe Bay (Coles et al., 2002a) and in 2001 at Kuapā Pond- 

Maunalua Bay (Coles et al., 2002b). It likely occurs in other O‘ahu harbors, including Pearl Harbor 

but was probably mixed with D. candidum in earlier studies (S. Godwin, pers. comm., 2000). 

Didemnum psammathodes (Sluiter, 1895) 

Introduced 

Didemnum psammathodes was found in Ke‘ehi Lagoon in 1998 (G. and C. Lambert) and at Waikīkī 

in 2001 and in 2002 at Kuapā Pond-Maunalua Bay (Coles et al., 2002b). It occurs throughout the 

Indo-West Pacific, from the Caribbean to Brazil (Kott, 1998; Monniot et al. 2001), and in Sierra 

Leone (Monniot & Monniot, 1994), to which Atlantic locations it is likely introduced. Lambert 

(2003) recorded it from Guam, as cryptogenic. 

Diplosoma listerianum (Milne-Edwards, 1841) Introduced 

= Diplosoma macdonaldi Herdman, 1886 

Another common compound ascidian of O‘ahu fouling communities and now too widespread to yet 

determine its original home (although Monniot et al. 2001, refer to this as a “European species”). 

Eldredge (1967) provided the following records, here arranged chronologically: 

1900 Honolulu L. Miller 

1902 Honolulu, from tug boat Albatross expedition 

1940 Pearl Harbor, hull of USS Dobin D. P.Abbott 

1941 Honolulu Harbor D. P. Abbott and C. B. Mills 

1942 O‘ahu: Queens Beach D. P. Abbott 

1946 O‘ahu: Halona Blowhole D. P. Abbott 

1961 Pearl Harbor L. Eldredge 

1961–1962 Kāne‘ohe Bay L. Eldredge 

Eldredge (1967) also reported specimens from Midway Point, Midway, collected in 1963.


137 

Various workers have continued to report it from Kāne‘ohe Bay (Rastetter & Cooke, 1979; 

Grovhoug & Rastetter, 1980; Coles et al., 2002a) and Pearl Harbor (Grovhoug, 1979; Grovhoug & 

Rastetter, 1980), in Oahu’s commercial harbors and public marinas (Coles et al., 1999b), off Waikīkī 

(Coles et al. 2002b), and in Kahului Harbor, Maui (Coles et al., 2004). Bailey-Brock (1989) recorded 

it (as D. macdonaldi) on PVC settlement plates on an artificial reef in open waters at 20 m, 2.4 

km off the southeast coast of O‘ahu. 

Hurlbut (1990, 1991a, 1991b) studied the larval abundance, settlement, and juvenile mortality 

of this species in Pearl Harbor. 

Lissoclinum fragile (Van Name, 1902) 

Introduced 

= Diplosoma (Lissoclinum) fragile 

Eldredge (1967) reported this now-widespread species from Kāne‘ohe Bay on O‘ahu based upon 

material collected in 1962 and 1963. It occurs throughout the Western Pacific, South Pacific, and 

Indian Ocean; described from Bermuda, it is also known from throughout the Caribbean, Brazil, and 

Sierra Leone on the West African coast (Eldredge, 1997; Monniot & Monniot, 1994). 

Cionidae 

Ciona intestinalis (Linnaeus, 1767) 

Introduced 

Edmondson (1933, Honolulu Harbor) and Abbott (1941, Pearl Harbor) are among the early reports 

of this species from Hawai‘i. It is widely considered to be native to the North Atlantic Ocean (Mon- 

niot et al., 2001), having been ship-dispersed to many harbors around the world. Abbott et al. (1997) 

note that it is found on boat hulls and docks in Pearl Harbor, Honolulu Harbor, and Kāne‘ohe Bay. 

D.P. Abbott collected it in 1940 along with Diplosoma listerianum, Styela canopus, and Ascidia sydneiensis 

on the hull of the USS Dobin in Pearl Harbor (Bishop Museum collections, and identification 

reconfirmed by Abbott in November 1980). 

Perophoridae 

Ecteinascidia imperfecta Tokioka, 1950 

Introduced 

This ascidian was described from Palau Island in the Western Pacific Ocean and later recorded from 

the Great Barrier Reef (Kott, 1998). Abbott et al. (1997) report it growing on worm tubes, dead coral, 

bryozoans, algae, and solitary ascidians in Kāne‘ohe Bay, Pearl Harbor, and the Ala Moana Yacht 

Basin. Bishop Museum collections include specimens from Kāne‘ohe Bay collected in 1979. 

Corellidae 

Corella minuta Traustedt, 1882 

Introduced 

= Corella japonica of authors 

Abbott et al. (1997) report this species from floating docks in Kāne‘ohe Bay. Coles et al. (2002a, 

2000 collections) continue to report it from there; Coles et al. (1999b, 1997 collections) also report 

it from Honolulu Harbor and from Kahukui Harbor, Maui (Coles et al., 2004). It occurs in the central 

and western Pacific, as well as in the Caribbean-Florida region (Abbott et al., 1997), to which 

latter region it is likely also introduced. 

Ascidiidae 

Phallusia nigra Savigny, 1816 

Introduced 

This Red Sea-Indian Ocean species has been introduced to Micronesia (Guam; Lambert, 2002) and 

to Hawai‘i. Charles H. Edmondson photographed this species in the 1930s in Pearl Harbor (Bishop 

Museum Department of Invertebrate Zoology files, photographs identified by G. Lambert in August 

2002). Long’s (1974) report of Ascidia melanostoma may belong here (Abbott et al., 1997, p. 36). 

Abbott et al. (1997) report it from rocks and dead coral “on barely subtidal mudflats” in 

Kāne‘ohe Bay and on floats and pilings in Kāne‘ohe Bay, Pearl Harbor, and Ke‘ehi boat harbor. 

Coles et al. (1999b) also report it from 64 sites throughout the main Hawaiian Islands, including 

Pearl and Honolulu Harbors, Ke‘ehi Lagoon, Ala Wai Harbor, Barbers Point Harbor, and Kewalo

138 


Basin, based upon 1997–1998 collections. Additional records include Kāne‘ohe Bay (Coles et al., 

2002a), Waikīkī and Koko Marina (Coles et al., 2002b), an Nawiliwili, Kahului and Hilo Harbors 

(Coles et al., 2004). Hurlbut (1991c) reported on settlement and juvenile survival in Pearl Harbor. 

Ascidia archaia Sluiter, 1890 

Introduced 

= Ascidia corelloides (Van Name, 1924) 

Abbott (1941) reported this western Pacific and Indo-Pacific species (as A. corelloides) from the 

Honolulu Aquarium; 50 years later, he referred to this incident when he noted that “Some years ago, 

small, clear specimens of Ascidia archaia grew on the walls of aquaria in the old Honolulu 

Aquarium” (Abbott et al., 1997). We take the earliest date to be 1940. Abbott et al. (1997) note that 

“recently” it was collected on solitary ascidians and oysters in the seawater run-off trenches of the 

Kewalo Marine Laboratory in Honolulu. They also suggest that it lives subtidally off Honolulu. 

Coles et al. (2002a) record it from Kāne‘ohe Bay. 

Ascidia sydneiensis Stimpson, 1855 

Introduced 

This southern hemisphere (Abbott et al., 1997) species has been introduced to Micronesia (Guam; 

Lambert, 2002) and to Hawai‘i. Charles H. Edmondson photographed this species in the 1930s in 

Pearl Harbor (Bishop Museum Department of Invertebrate Zoology files, photographs identified by 

G. Lambert in August 2002). It was recorded by Abbott (1941), probably based upon specimens he 

collected in Pearl Harbor in 1940 on the hull of the USS Dobin (Bishop Museum collections; specimens 

reconfirmed by D. P. Abbott in November 1980). Abbott et al. (1997) report it the “commonest 

large ascidian” of calm O‘ahu harbor waters, noting that it “exploits docks, floats, boat hulls, 

solid debris on mudflats, and other firm substrates” in Kāne‘ohe Bay, Pearl Harbor, and Ke‘ehi Boat 

Harbor. Coles et al. (1999a) report it from Pearl Harbor, 1996 collections, and Coles et al. (1999b) 

report it from Honolulu Harbor, Ke‘ehi Lagoon, Ala Wai Harbor, Barbers Point Harbor, and Kewalo 

Basin, based upon 1997–1998 collections. It also occurs in Kāne‘ohe Bay (2000 collections; Coles 

et al., 2002a), Koko Marina (Coles et al., 2002b), and Nawiliwili, Kaunakakai, and Hilo Harbors 

(Coles et al., 2004). 

Monniot et al. (1991) also emphasize the lower-energy habitat of this species, noting that it 

“flourishes not only on buoys in ports but also on broken shells in sheltered lagoon waters.” In this 

regard, we note the report by Hoover (1998, page 342, 2006) of an ascidian identified as Ascidia sydneiensis 

from exposed, higher-energy rocky shores at about 10-m depth at Pūpūkea, O‘ahu, at a site 

“favored by scuba divers”, where it is usually found “in crevices along vertical walls with only the 

siphons exposed.” Hoover noted that the color of these animals in open water was yellow-green, as 

opposed to the silt-covered, darker animals of pilings and floats. Hoover also remarked that “specimens 

were dredged from 150–250 feet by the steamer Albatross in the early 1900s” [1902]. 

Although introduced ascidians do occur in Hawai‘i in more exposed locations, both of these habitats 

are very unusual for A. sydneiensis and would require further investigation. We have not been 

able to locate the source of the statement that the Albatross collected this species in deep water, 

which could be in error (J. Hoover, pers. comm., February 2007). 

Ascidia species A 

Introduced 

Ascidia species B 

Introduced 

Abbott et al. (1997) report two additional Ascidia species as “fairly common on floats and subtidal 

reefs in protected habitats” in Kāne‘ohe Bay and Pearl Harbor. They suggest that Ascidia sp. B may 

be Ascidia gemmata Sluiter, 1895, a widespread tropical western Pacific species. Coles et al. (1999b) 

further report Ascidia sp. A from Honolulu Harbor, Ke‘ehi Lagoon, Ala Wai Harbor, and Kewalo 

Basin, and a reef site and Port Allen Harbor on Kaua‘i based on 2002 collections (Coles et al., 2004, 

2006). Ascidia sp. B is known from Pearl Harbor (Coles et al., 1997, 1999a), Honolulu Harbor, 

Ke‘ehi Lagoon, Ala Wai Harbor (Coles et al., 1999b), Kāne‘ohe Bay (Coles et al., 1999a), and 

Kahului Harbor (Coles et al., 2004).


139 

Ascidia sp. 

Introduced 

Godwin & Lambert (2000) report a third unidentified, very long and yellow-green Ascidia sp., collected 

in 1998 in Kewalo Basin, not previously recorded from Hawai‘i. It was found attached to 

pieces of rug hanging in the water. 

Styelidae 

Styela canopus Savigny, 1816 

Introduced 

= Styela partita (Stimpson, 1852) 

Donald P. Abbott collected this Western Pacific species on the hull of the USS Dobin in 1940 in Pearl 

Harbor (Bishop Museum collections; Abbott, 1941). Abbott (1941) noted it occurred on boat bottoms 

and wharf pilings in Pearl Harbor and Kāne‘ohe Bay. Coles et al. (1999b) report it from 

Honolulu Harbor, Ke‘ehi Lagoon, Ala Wai Harbor, Barbers Point Harbor and Kewalo Basin, based 

upon 1997–1998 collections. It remains in Kāne‘ohe Bay as well (Coles et al., 2002a), and was 

found in 2003 in Kahului Harbor, Maui (Coles et al., 2004). Earliest Bishop Museum material from 

the islands is from Pearl Harbor based upon collections made in 1929 by C.H. Edmondson. 

Eusynstyela hartmeyeri Michaelsen, 1904 

Introduced 

= Eusynstyela aliena Monniot, 1991 

Godwin & Lambert (2000) report this species (as E. aliena) from Pearl Harbor (1998), Honolulu 

Harbor (1998), and Ke‘ehi Lagoon (1998). Coles et al. (1999b) also report it from Honolulu Harbor, 

Ke‘ehi Lagoon, and Barbers Point Harbor, based upon 1997–1998 collections. Coles et al. (2002b) 

further report it in Kāne‘ohe Bay and Hawai‘i Kai. Coles et al. (2006) report it from Kaua‘i, based 

upon 2003 collections. It may have been overlooked in an earlier (Coles et al., 1997) 1996 survey 

of Pearl Harbor (S. Godwin, pers. comm., 2000). Monniot & Monniot (2001) synonymized E. 

aliena, described from New Caledonia in the southwest Pacific Ocean, with the earlier E. hartmeyeri 

from the Red Sea and the Indian Ocean. 

Cnemidocarpa irene (Hartmeyer, 1906) 

Introduced 

= Cnemidocarpa areolata in Abbott et al., 1997 

Abbott et al. (1997) report this species from docks in Kāne‘ohe Bay, and noted it was “found occasionally 

on the walls of tanks in the old Honolulu Aquarium, presumably from settlement of larvae 

entering the aquarium’s seawater intake at Waikīkī.” We take the latter reference to be the same as 

the one for Ascidia archaia, and thus a species first observed by Abbott in 1940–1941. Coles et al. 

(1999b) report it from Honolulu Harbor, Ke‘ehi Lagoon, Barbers Point Harbor, and Kewalo Basin, 

based upon 1997–1998 collections. Coles et al. (2002a) report it from Kāne‘ohe Bay, and Coles et 

al. (2002a) report it form Waikīkī and Hawai‘i Kai, and Coles et al. (2004) report it from Nawiliwili 

and Kawaihae Harbors. 

Botrylloides simodensis Saito & Watanabe, 1981 Introduced 

Godwin & Lambert (2000) report this Western and South Pacific species from O‘ahu in November 

1998; the location, not mentioned, is Pearl Harbor (S. Godwin, pers. comm., 2000). Coles et al. 

(1999b) also report it from Ala Wai Harbor and Kewalo Basin, based on 1998 specimens, and from 

2001 collections at Waikīkī (Coles et al., 2002b). 

Botryllus sp. or spp. 

Introduced 

Botrylloides sp. or spp. 

Introduced 

At least two additional botryllids, one each in each of these two genera, are present in fouling communities 

in Pearl Harbor and Kāne‘ohe Bay (Straughan, 1969b; Grovhoug, 1979; Grovhoug & 

Rastetter, 1980) with collections dating from 1973 (Coles et al., 1997). These species remain unresolved 

taxonomically (Abbott et al., 1997).

140 


Polyandrocarpa sagamiensis Tokioka, 1953 

Introduced 

Abbott et al. (1997) report this ascidian from Kāne‘ohe Bay and Pearl Harbor on floating docks. It 

occurs in the western Pacific and the south Pacific. It has also been found in 1999–2000 collections 

in Kāne‘ohe Bay (Coles et al., 2002a) and Hawai‘i Kai (Coles et al., 2002b; 2001 collections). Coles 

et al. (1999b) report it from Honolulu Harbor, Ke‘ehi Lagoon, Barbers Point Harbor, and Kewalo 

Basin, based upon 1997–1998 collections, and from 2002–2003 collections in Nawiliwili and 

Kawaihae Harbors (Coles et al., 2004). 

Polyandrocarpa zorritensis (Van Name, 1931) 

Introduced 

Abbott et al. (1997) report this ascidian from Kāne‘ohe Bay, Pearl Harbor, and Kewalo Bay in fouling 

communities. It occurs also in Peru, southern California, and the Mediterranean (Abbott et al., 

1997). Coles et al. (1999b) report it from Ke‘ehi Lagoon based on 1997 collections and from 1999 

collections in Kāne‘ohe Bay (Coles et al., 2002a). 

Polycarpa aurita (Sluiter, 1890) 

Cryptogenic 

This very widespread western Atlantic and Indo-Pacific species is reported from floats in Kāne‘ohe 

Bay, subtidally in the Honolulu Harbor channel, in the seawater system of the Kewalo Marine 

Laboratory in Honolulu, under rocks in tidepools at Halona, O‘ahu, and on the island of Hawai‘i 

under “nearshore rocks” at Napo‘opo‘o in Honaunau Bay (Abbott et al., 1997). Coles et al. (1999b) 

report it from Honolulu Harbor, Ke‘ehi Lagoon, Barbers Point Harbor and Kewalo Basin, based 

upon 1997–1998 collections. Coles et al. (2002a) report it from Kāne‘ohe Bay, and Coles et al. 

(2000b) further report if from Waikīkī and Hawai‘i Kai. Lambert (2002) considers it cryptogenic in 

Guam. 

Symplegma reptans (Oka, 1927) 

Introduced 

This seasquirt, previously known from Japan and Australia (Kott, 1998) was reported by Coles et al. 

(1999a, 1999b) from Pearl Harbor based on 1996 collections. We suspect it was introduced from 

Japan, given its much wider distribution there (Kott, 1985). 

Symplegma brakenhielmi (Michaelsen, 1904) 

Introduced 

= Symplegma oceania Tokioka, 1961 

= Symplegma connectans of authors 

The first record of this Indo-Pacific species appears to be that of Straughan (1969b), based upon 

specimens collected in Kāne‘ohe Bay in 1967. Abbott et al. (1997) report it for Kāne‘ohe Bay, Pearl 

Harbor, and Honolulu Harbor in fouling communities. Grovhoug (1979; 1978 collections) and 

Grovhoug & Rastetter (1980), both as S. connectans, report it for Pearl Harbor and Kāne‘ohe Bay 

as well. Coles et al. (1997, 1999a) also report it from 1996 collections in Pearl Harbor, and it was 

also found in 1997–1998 collections in Honolulu Harbor, Ke‘ehi lagoon, Kewalo Basin, and Ala Wai 

Yacht Harbor (Coles et al., 1999b), 2001 collections off Waikīkī (Coles et al., 2002b) and 2003 collections 

in Kawaihae Harbor (Coles et al., 2004). 

Symplegma sp. 

Introduced 

A third Symplegma species, as yet not identified, was collected in Pearl Harbor (1998 collections) 

(G. Lambert & S. Godwin, pers. comm., 2000). 

Polyclinidae 

Polyclinum constellatum Savigny, 1816 

Introduced 

= (?) Polyclinum vasculosum of Tokioka, 1967, Pearl Harbor records vide Abbott et al., 1997 

This tropical and subtropical cosmopolitan species has long been present in Hawai‘i. Charles H. 

Edmondson photographed this species in the 1930s in Pearl Harbor (Bishop Museum Department of 

Invertebrate Zoology files, photographs identified by Gretchen Lambert in August 2002). Bishop 

Museum material dates from 1960 in Kāne‘ohe Bay. Abbott et al. (1997) note that it “sometimes


141 

abundant” in Kāne‘ohe Bay, Pearl Harbor, Honolulu Harbor, and Ala Moana yacht harbor, “growing 

on dock pilings, floats, dead corals, or coral rubble lying on sandflats.” They also note populations 

in “more exposed places, such as Honolulu’s Black Point,” where small colonies grow under rocks. 

Abbott et al. (1997) imply that Tokioka’s (1967) identification of Polyclinum vasculosum from 

Pearl Harbor are likely P. constellatum, and we tentatively so treat those reports here. Polyclinum 

vasculosum Pizon, 1908, is known from the tropical western Pacific. Regardless of whether the specimen 

are P. constellatum or P. vasculosum, they represent some of the earliest introduced ascidians 

collected in Hawai‘i. Tokioka notes material collected in Pearl Harbor by P. Bartsch in September 

1920 in USNM collections, as well as “three large colonies”, the largest measuring 14.5 cm by 11.0 

cm, collected on O‘ahu by T.H. Streets, but without noting a date. As discussed under the crab 

Carcinus maenas, we know that Streets collected on O‘ahu only once, in 1873–1874. 

Coles et al. (1997, 1999a) report P. constellatum from Pearl Harbor; Coles et al. (1999b) report 

it from Honolulu Harbor (1997) and Ala Wai Harbor (1998); from 1999–2000 collections in 

Kāne‘ohe Bay (Coles et al., 2002a), and from 2001 collections off Waikīkī (Coles et al., 2002b). 

Pyuridae 

Microcosmus exasperatus Heller, 1878 

Introduced 

Abbott (1941, as Microcosmus claudicans exasperatus) reported this now-cosmopolitan species as one 

of the most common ascidians with collections from Honolulu Harbor, Kāne‘ohe Bay, and Pearl Harbor. 

We take the earliest date to be 1940. Abbott et al. (1997) note that this species occurs in the seawater 

runoff trenches in Honolulu of the Kewalo Marine Laboratory and that it is common on floats and docks 

in Kāne‘ohe Bay. Coles et al. (1997, 1999a) report it from Pearl Harbor, and Coles et al. (1999b) report 

it from Honolulu Harbor, Ke‘ehi Lagoon, Ala Wai Harbor, Barbers Point Harbor and Kewalo Basin, 

based upon 1997–1998 collections. Abbott et al. (1997) also note the description of Microcosmus miniaceus 

Sluiter, 1900, from Pearl Harbor and its failure to be recorded again. Sluiter’s original material 

requires re-examination to determine if these represent early records of M. exasperatus. 

Mastrototaro & Dappiano (2005) report M. squamiger from Hawai‘i, misciting Godwin (2003b) 

as Godwin (2003a). Godwin (2003b) reported M. squamiger in fouling on vessels arriving from 

California in the Hawaiian Islands. 

Herdmania mauritiana (von Drasche, 1884) 

Introduced 

= Herdmania insolita Monniot & Monniot, 2001 vide Nishikawa (2002) 

This species was found on floats in Ke‘ehi lagoon in July 1985 by G. Lambert. Lambert (2003) treated 

this species (as H. insolita) as cryptogenic in Hawai‘i. Herdmania insolita is broadly distributed 

across the Indo-Pacific (Kott, 2002). 

Herdmania momus (Savigny, 1816) 

Introduced 

Charles H. Edmondson photographed this Indo-Pacific (Kott, 2002) species in the 1930s in Pearl 

Harbor (Bishop Museum Department of Invertebrate Zoology files, photographs identified by G. 

Lambert in August 2002). Abbott et al. (1997) record it from the Honolulu Harbor channel and in 

Kāne‘ohe Bay and Ke‘ehi Lagoon on mudflat debris and in fouling communities. Long (1974) also 

reported it from fouling panels set just offshore of O‘ahu. Coles et al. (1999b) report it from Ke‘ehi 

Lagoon, Ala Wai Harbor, Barbers Point Harbor, and Kewalo Basin, based upon 1997–1998 collections, 

and note records since 1991 for Honolulu Harbor. It also occurs in Waikīkī and Hawai‘i Kai 

(Coles et al., 2002b). Earliest Bishop Museum material for Kāne‘ohe Bay is 1963. 

Herdmania pallida (Savigny, 1816) 

The Indo-Pacific Herdmania pallida (Kott, 2002) was found in 1990, growing alongside Herdmania 

momus, on floats in Ke‘ehi Boat Harbor, O‘ahu, by G. and C. Lambert (Abbott et al., 1997, page 54; 

this is the species listed as Herdmania sp., vide G. Lambert, pers. comm., December 2006). It has 

also been collected in Pearl Harbor in 1998. Coles et al. (2002a) report it from Kāne‘ohe Bay 

(1999–2000), and from 2002–2003 collections from reefs and harbors on Kaua‘i, Moloka‘i, Maui, 

and Hawai‘i (Coles et al., 2004, 2006).

142 


PISCES 

Introduced Fish 


Herklotsichthys quadrimaculatus 1972 R West Pacific: Marshall Is 

Sardinella marquesensis 1955 R South Pacific: Marquesas 

Valamugil engeli 1955 R South Pacific: Marquesas 

Poecilia latipinna 1905 R Texas 

Poecilia salvatoris x P. mexicana -group: 

hybrid complex >1960 R North America 

Limia vittata 1950 R Cuba 

Gambusia affinis 1905 R Eastern North America 

Cephalopholis argus 1956 R S Pacific: Society Islands 

Lutjanus fulvus 1956 R S Pacific: Society Islands 

Lutjanus gibbus 1958 R S Pacific: Marquesas/ 

Moorea 

Lutjanus kasmira 1958 R South Pacific: Marquesas 

Upeneus vittatus 1955 R South Pacific: Marquesas 

Oreochromis mossambicus 1951 R Singapore 

Sarotherodon melanotheron 1962 R West Africa 

Parablennius thysanius 1971 SF, BW Indo-Pacific 

Omobranchus ferox 1998 SF, BW Philippines, S China Sea 

Omobranchus rotundiceps obliquus 1951 R Indo-Pacific (Samoa) 

Mugilogobius cavifrons 1987 SF, BW Indo-Pacific 


Abudefduf vaigiensis 1991 drift nets; R? Indo-Pacific 

Status: Waif with drift nets? 

Centropyge flavissima 1990 R Indo-Pacific 

Status: Establishment unknown 

Chrysiptera taupou 2002 R Southwest Pacific 


Favonigobius sp. 1990 BW Australasian? 


Ten species of anchovies, bass, groupers, snappers, emperors, and flagtails (see table, below) 

Status: Released not established 

Valuable summaries or reviews of the introduced fish of the islands are those of Randall (1960, 1981, 

1987, 1996), Randall & Kanayama (1972, 1973), Oda & Parrish (1982), Maciolek (1984), Eldredge 

(1987), Randall et al. (1993), and Mundy (2005, p. 83, Tables 7 and 20, and individual species treatments). 

Friedlander et al. (2006) provide data on habitat patterns in marine protected areas of several 

introduced species. Additional papers (treated below) cover recent invasions of some ballast water 

introductions. 

Fisheries Releases 

In what would now be environmentally and politically impossible, in the 1950s and early 1960s the 

United States Bureau of Commercial Fisheries (now the NOAA National Marine Fisheries Service), in 

cooperation with the then-named State of Hawaii Division of Fish and Game, undertook an extensive 

program to introduce, largely from the South Pacific, both game fish and bait fish into the Islands. No 

fewer than 11 species of snappers and groupers—classic Indo-Pacific members of which were absent 

from the Hawaiian fauna—were released into open waters of the state between 1955 and 1961 (Oda & 

Parrish, 1982; Eldredge, 1987; Randall, 1987; see also Brock, 1962). Upon arrival in Honolulu in Sep - 

tember 1956, one group of fish was treated for five days, while still in the livewell, with copper sulphate, 

to remove external parasites (Randall & Kanayama, 1972, 1973). This “1950s epoch” was preceded by 

earlier attempts in the 1920s and 1930s to introduced anchovy and striped bass from California (see 

table, below).


143 

Randall & Stender (2002) commented upon one release as an example of the difficulties in predicting 

when a species will become established. They noted that 469 dwarf spotted groupers (Epinephelus 

merra) from Moorea in the Society Islands were released into Kāne‘ohe Bay and 132 off Wainiha, 

Kaua‘i (see table below), “a proper choice in terms of habitat because the species occurs primarily in 

lagoons [and] the introduced adults lived out their lives, but no young ever appeared.” Temperature, they 

noted, did not appear to be a factor. “Therefore, some unknown factor or factors other than temperature 

prevented their becoming established in Hawaiian waters (which was fortunate because the species was 

not a good choice for introduction due to its small size).” We note that it was also fortunate because of 

the unknown ecological impacts an introduction may have had. 

Hourigan & Reese (1987) noted that of the above 11 species, only four were introduced in numbers 

exceeding 2000 individuals—and three of these became established. Planes & Lecaillon (1998) 

reviewed the population genetics of the three successful species, finding that no significant change in 

genetic diversity had occurred, although some gene selection had occurred in the bluestripe snapper 

(ta‘ape) Lutjanus kasmira. 

Also in the 1950s, a program to introduce the Marquesan sardine for tuna bait was undertaken; 

remarkably, two additional, untargeted fish were released at the same time, resulting in a total of three 

more fish introductions. As late as the 1970s, intentional albeit unsanctioned releases occurred: a vessel 

chartered by the National Marine Fisheries Service loaded goldspot herring in the Marshall Islands to 

use as tuna bait on the voyage back to Hawai‘i—and subsequently released these fish into Kewalo Basin, 

O‘ahu! 

Impact considerations of these intentional releases were voiced: Murphy (1960), in discussing the 

introduction of the Marquesan sardine, wrote that “Ordinarily, such an expensive project as transferring 

a marine fish from 10° south of the Equator to 20° north of the Equator would not be undertaken without 

an extensive investigation of the fish’s life history ... to weigh thoroughly the possibility that the 

species might do harm in the new area.” However, he then simply noted that “The possibility of harm 

seemed remote.” Relative to the large groupers and snappers, Randall & Kanayama (1972) remarked 

that the species with which these introduced fish would compete, or the prey they would eat, were of little 

commercial or economic importance; today, the potential for such introductions to compete with or 

prey upon noncommercial species would focus on the unknown ecological cascades that could result 

from such introductions. Randall & Kanayama (1972, 1973) concluded both essays with a strong caveat 

about the risk of introducing organisms into the ocean given the improbability of their removal if they 

were to succeed. 

The table below lists the species of fish which were intentionally released during the above periods 

of operation but did not become established. 

Fisheries Releases: Species Not Established in Hawaiian Waters 

(from Brock, 1960; Maciolek, 1984; Randall, 1987; Mundy, 2005) 

Species Date Introduced from 

Engraulidae (anchovies) 

Anchoa compressa 1932 California 

Moronidae (bass) 

Morone saxatilis 1922 California 

Serranidae (groupers) 

Cephalopholis urodeta 1958, 1961 Marquesas 

Epinephelus fasciatus 1958, 1961 Marquesas 

Epinephelus hexagonatus 1958, 1961 Marquesas 

Epinephelus irroratus 1958 Marquesas 

Epinephelus merra 1956, 1961 Society Islands 

Lutjanidae (snappers) 

Lutjanus guttatus 1960 Mexico 

Lethrinidae (emperors) 

Lethrinus sp. 1958 Marquesas Islands 

Kuhliidae (flagtails) 

Kuhlia rupestris 1957–1958 Guam

144 


Aquarium Releases 

With the post-World War II increased interest in and capability of moving aquarium hobby fish 

around the world, an increasing number of such species have been released by the public or industry 

into open waters, resulting in either established populations or—in regions where large numbers 

of knowledgeable observers are in the water, as in the Hawaiian Islands—the discovery of either 

cast-out pets or (as suspected in some cases) the intentional release of such fish. In subtropical or 

tropical regions, distinguishing aquarium releases from waifs (potential colonists carried far afield 

by ocean currents) may be a challenge: here, evidence such as distance from a possible source, the 

number of individuals found, their presence over widespread locations over a number of years, and 

so forth, may be mustered as evidence to attempt to distinguish between these two categories. 

We have not attempted to list here all such discarded or released aquarium animals; the table 

below represents examples only. It is to be expected that there will be a continual record of openwater 

discoveries of exotic species in the islands, based upon such illegal releases. Mundy (2005), 

in discussing the occasional appearances of anemonefish (clownfish) in the islands, remarked that, 

“In at least one such case, an aquarium-fish wholesaler has been reported to have purposefully 

released Amphirion into [Kāne‘ohe] Bay with the intent of establishing a breeding population that 

could then be ‘ranched’ for sale.” 

Aquarium Releases: Discarded or Released Animals Found in Open Water 

in the Hawaiian Islands (Not Known to be Established) 

(Randall, 1981; Mundy, 2005) 

Species Date Remarks 

Serranidae 

Cromileptes altivelis 1977–1978 Hawai‘i Island, Maui, 1980s; 

O‘ahu early 1990s 

Lutjanidae 

Lutjanus sebae 2002 One individual, O‘ahu 

Pomacanthidae 

Apolemichthys xanthopunctatus 1980s? O‘ahu, several individuals: purposely 

released by an aquarium fish 

importer 

Chaetodontoplus mesoleucus 1994 One individual, O‘ahu 

Pomacanthus semicirculatus 2001 One individual, O‘ahu 

Pygoplites diacanthus 1997 One individual, O‘ahu 

Pomacentridae 

Amphiprion sp. 1980s–1990s Kāne’ohe Bay, O‘ahu 

(see text) 

Ephippidae 

Platax tiera 2003 One individual, Maui 

Acanthuridae 

Acanthurus leucosternon 2003 One individual, O‘ahu 

Paracanthurus hepatus 1994–1996 O‘ahu, Maui; possibly waifs? 

Fouling Intercepts 

Mundy (2005, Table 7, page 23) listed Pseudochromus ?tapeinosoma, Cirrhitichthys falco, 

Cirrhitichthys oxycephalus, Enneapterygius bahasa/nigricauda, and Ecsenius bicolor as “nonindigenous 

fish species that were (or likely were) released into open waters” in Hawai‘i “but that 

failed to establish reproducing populations.” These five species were collected from a USN concrete 

barge, the YON-146, which came into Pearl Harbor from Guam in 1950.


145 

Clupeidae 

Herklotsichthys quadrimaculatus (Ruppell, 1837) Introduced 

The goldspot sardine (also known as the goldspot herring) was introduced in 1972 when the contents 

of the livewell aboard the tuna fishing vessel Anela was released into Kewalo Basin, O‘ahu (Randall, 

1987). The fish had been taken up as potential tuna bait in the lagoon of Jaluit in the Marshall 

Islands; the vessel had been chartered for exploratory fishing by the National Marine Fisheries 

Service. Baldwin (1984) and Williams & Clarke (1983) provide documentation on the introduction. 

It is now an abundant fish in the islands. Greenfield (2003) noted it from Kāne‘ohe Bay based upon 

1990–1995 collections. Mundy (2005) gives the Hawaiian distribution as Lāna‘i to O‘ahu at 1–13 

m. 

Sardinella marquesensis Berry & Whitehead, 1968 Introduced 

= Harengula vittata of authors 

The Marquesan sardine was intentionally released into Hawaiian waters from the Marquesas 

between 1955 and 1959 to establish a tuna bait fishery (Brock, 1960; Randall, 1987, 1996). A total 

of 143,800 fish were released around O‘ahu in five years; it is believed to have become established 

by 1958 (Brock, 1960). Murphy (1960) and Hida & Morris (1963) provide details of the introduction 

and early establishment. Murphy (1960) noted that wide publicity attended the releases, including 

a special poster that was distributed to fishermen. It remains present but not abundant in the 

islands. Mundy (2005) gives the Hawaiian distribution as Maui to Kaua‘i, in shallow water. 

Mugilidae 

Valamugil engeli (Bleeker, 1858) 

Introduced 

= Moolgarda engeli, = Chelon engeli 

The kanda, a mullet, is another species released in 1955 with the first shipment of Marquesan sardines, 

although not detected in Hawai‘i until 1966 (Randall, 1987). Randall (1981) noted that this 

mullet was of “little or no commercial value (but) seems to have increased in abundance at the 

expense of the commercially important mullet Mugil cephalus.” Maciolek & Timbol (1981) reported 

it from the Kahana Estuary, O‘ahu, based on collections made from 1969 to 1971. Mundy (2005) 

notes that it has spread throughout the main islands in estuaries. 

Poecilidae 

Poecilia latipinna (LeSueur, 1821) 

Introduced 

= Mollienesia latipinna in Brock, 1960 

The sailfin molly, or tabai, was introduced into Moanalua Stream, O‘ahu, in 1905 for mosquito control 

purposes, from Texas (Randall, 1987; Yamamoto & Tagawa, 2000). It is established in brackish 

water areas of O‘ahu and Moloka‘i (Brock, 1960; Randall, 1987), Maui (Randall, 1987), and Ha - 

wai‘i and Kaua‘i (Mundy, 2005). 

Yamamoto & Tagawa (2000) note that, “Old-timers will probably remember the large numbers 

of sailfin mollies established in the brackish water canals and lagoons of Ala Moana Beach Park, 

O‘ahu, during the sixties and seventies. This species disappeared from this waterway by the late seventies, 

to be replaced by the Liberty Molly and the Cuban Molly” (two introduced freshwater fish). 

Coles et al. (1999a) record it from Pearl Harbor. Englund et al. (2000a) report it as locally common 

in 1 to 9 ‰ in Pouhala Marsh in Pearl Harbor. 

Poecilia salvatoris Regan, 1907 x Poecilia mexicana Steindachner, 1863-group: 

hybrid complex 

Introduced 

The shortfin molly was introduced sometime after 1960 [Maciolek (1984), who suggests this based 

on the absence of its treatment by Brock in his 1960 paper; Randall (1987) cites Maciolek as the 

basis for this introduction occurring “sometime before 1950”, but this information does not appear 

in Maciolek’s paper.] Based upon Randall’s statement (which seems to be in error) and earlier surveys, 

Englund (2002) indicated the introduction took place between 1940 and 1950. We use “after

146 


1960”, following Maciolek. The shortfin molly is native to North America and is a probable aquarium 

release. Yamamoto & Tagawa (2000) note that the populations now on O‘ahu appear to represent 

a hybrid of “two or more species from the P. salvatoris/mexicana complex”; we retain the above 

name here for purposes of convenience. Englund & Baumgartner (2000) report Poecilia mexicana 

in 35–36 ‰ in Hālawa Stream, Pearl Harbor, co-occurring with the introduced blenny Omobranchus 

ferox. As both the salvatoris-clade and the mexicana-clade are largely rooted on the Atlantic coasts 

of North and Central America, we treat this introduction biogeographically as Atlantic in origin. 

Englund et al. (2000a) note that this species “was one of the most common fish found in lower 

stream and estuarine areas of Pearl Harbor” in salinities up to 36 ‰. 

Limia vittata (Guichenot, 1853) 

Introduced 

= Poecilia vittata of Hawaiian literature 

The Cuban molly (or Cuban limia, Cuban topminnow, or tabai) was present on O‘ahu by 1950 

according to Randall (1987). Brock (1960) noted that it “is found in the streams flowing through 

Honolulu which include Moanalua, Kapalama Canal, Nuuanu, and Manoa streams. It is found in the 

lower portions of the streams and in salt water.” Englund et al. (2000a) found it to be relatively 

uncommon in Pearl Harbor estuarine streams. It is another probable aquarium release. 

Gambusia affinis (Baird & Girard, 1853) 

Introduced 

The well-known eastern North American mosquitofish was released intentionally for mosquito control 

at Moanalua, O‘ahu, in 1905; the specimens originated from Texas (Brock, 1960). Today it is 

common in brackish water habitats (Yamamoto & Tagawa, 2000). Englund et al. (2000a) found it to 

be in “extremely high densities in the lower reaches of O‘ahu streams and always in the presence of 

Poecilia mexicana”; they occurred in salinities of up to 40 ‰. Mundy (2005) gives its distribution 

as Hawai‘i, Maui, Moloka‘i, O‘ahu, and Kaua‘i, “in all low current, freshwater habitats and in brackish-water 

lagoons and channels.” 

Serranidae 

Cephalopholis argus Bloch & Schneider, 1801 Introduced 

= Cephalopholis guttatus of authors 

In 1956 bluespotted grouper (or roi, its Tahitian name; also known as the peacock grouper or peacock 

rockcod) was intentionally released from the livewell of the vessel Hugh M. Smith that had 

transported the fish from Moorea in the Society Islands. The fish were released off Brown’s Camp, 

O‘ahu (571 individuals) and off Keāhole Point, Hawai‘i (400); establishment was thought to have 

occurred by 1958 (Randall, 1987). It is most common on the island of Hawai‘i (Randall, 1987, 

1996); it has not become abundant anywhere (Planes & Lecaillon, 1998). Coles et al. (2002a) report 

it from Kāne‘ohe Bay; further records are from off Waikīkī (Coles et al. 2002b), Hanauma Bay and 

various neighbor island locations on Kaua‘i, Moloka‘i,Maui, Hawai‘i (Coles et al. 2004) and Lāna‘i 

(Coles et al. 2005). Mundy (2005) reports its Hawaiian range as Hawai‘i Island to French Frigate in 

6–12 m. It naturally occurs from the East Africa to French Polynesia (Planes & Lecaillon, 1998). 

Lutjanidae 

Lutjanus fulvus (Forster in Bloch & Schneider, 1801) Introduced 

The blacktail snapper (or toau, its Tahitian name) was intentionally released from the same vessel as 

the bluespotted grouper; 239 fish from the Society Islands were released into Kāne‘ohe Bay from 

Moorea in 1956 (Randall, 1987; Planes & Lecaillon, 1998). In 1958 “additional fish”, from the 

Marquesas, were released into Kāne‘ohe Bay (Randall, 1987). It is established on O‘ahu and all 

other islands in the Archipelago (Randall, 1987). Hoover (1993) noted that it “has become common 

throughout the Hawaiian Islands”, and presented a photograph of an individual at 12 m at Honaunau, 

Hawai‘i. Hoover (1993) notes that “large schools of these showy snappers are a common sight 

around wrecks and other popular dive sites, especially where fishes have been hand-fed.” Russo 

(1994) notes that in Hawai‘i “it is usually found alone under ledges or in caves mixed in with sol-


147 

dierfishes or in the open cruising over the reef.” Coles et al. (1999a) record it from Pearl Harbor; 

further locations include Honolulu and Ala Wai Yacht Harbors (Coles et al., 1999b) Kāne‘ohe Bay 

(Coles et al., 2002a), Koko Marina (Coles et al., 2002b) and various reef sites and harbors on Kaua‘i, 

Moloka‘i, Maui, Hawai‘i (Coles et al., 2004, 2006) and Lāna‘i (Coles et al., 2005). DeFelice & 

Parrish (2003) provide data on its occurrence and diet in Hanalei Bay on the island of Kaua‘i. 

Mundy (2005) gives the Hawaiian distribution as from Hawai‘i Island to French Frigate Shoals, 

from 3 to 128 m. 

Lutjanus gibbus (Forsskål, 1775) 

Introduced 

This humpback snapper was released in O‘ahu in relatively small numbers (40 in 1958 from the 

Marquesas and 137 in 1961 from Moorea), and was long thought not to have not become established 

(Randall et al., 1993; Mundy, 2005). Surprisingly, living individuals were discovered commencing 

in the late 1980s: one fish was taken in 1989 off the Hamākua coast of Hawai‘i Island, and about six 

fish were caught in the early 1990s off Ka‘ena Point, O‘ahu, at night in 18 m. An additional fish was 

observed at 15 m on the south shore of Kaua‘i in April 1992. It appears to be established, but whether 

from the original introduction has not been determined. Genetic comparison of Hawaiian populations 

with those from French Polynesia would thus be of interest. 

Lutjanus kasmira (Forsskål, 1775) 

Introduced 

The bluestripe or blueline snapper (or ta‘ape, its Tahitian name) was intentionally released in 1958 

into Kāne‘ohe Bay, based upon 2,435 individuals transported from the Marquesas (Brock, 1960; 

Randall, 1987). It was considered to have become established by 1960. Oda & Parrish (1982) reported 

ecological observations and food habits in Hawai‘i and its “mild commercial success.” Randall 

(1987) noted that it occurs as far as Laysan and has attained great abundance on the island of 

Hawai‘i. It has also invaded deeper waters, occurring to depths of 200 m (Randall, 1987) and 275 m 

(Randall, 1996). Coles et al. (2006) report it from reef sites on all the main Hawaiian Islands. Mundy 

(2005) noted the distribution as being from Hawai‘i Island to Midway, from 4 to more than 200 m 

depth. 

The bluestripe snapper has become very abundant locally. Randall (1996) also noted that it 

forms large semi-stationery schools during the day, and disperses to feed at night “mainly on crustaceans 

and small fishes.” Randall (1987) remarked that “it is very unpopular with fishermen who 

are convinced that its increase has been at the expense of more valuable species, such as the goatfish 

Parupeneus porphyreus and P. multifasciatus. Also, it enters fish traps readily and comes quickly 

to baited hooks, thus catches of more desirable fish are reduced.” Schumacher & Parrish (2005) 

reported that Lutjanus kasmira displaces the native yellowtail goatfish Mulloidichthys vanicolensis 

from shelter, causing the goatfish to occur higher in the water column when Lutjanus was present, 

and thus potentially making Mulloidichthys more vulnerable to predators and fishermen. 

Morales-Nin & Ralston (1990), Friedlander et al. (2002), and DeFelice & Parrish (2003) provide 

data on age, growth, biology, and ecology of L. kasmira in Hawai‘i. Morales-Nin & Ralston 

(1990) report that it has one of the highest growth rates of any lutjanid fish in the Pacific, perhaps, 

they suggest, because its growth “may have been enhanced by the relative lack of competitors in the 

depauperate Hawaiian marine fish community.” Friedlander et al. (2002) describe its occurrence 

across a variety of habitats, as the “second most abundant species by numbers and biomass over a 

hard substratum.” DeFelice & Parrish (2003) detail the habitat occurrence, abundance, and prey of 

L. kasmira in Hanalei Bay on the island of Kaua‘i, based upon studies conducted in 1993–1994. 

Lutjanus kasmira was the most numerous species, and “was observed during 100% of nocturnal censuses 

on transects 0–25 m from the reef.” It was the most abundant species over soft-bottom habitats. 

Both L. kasmira and L. fulvus were characterized as “important predators upon soft-bottom 

invertebrates in the bay”, but noted that “it is difficult to speculate on competition for food between 

L. kasmira and native species”: dietary overlap between L. kasmira and other species was relatively 

low, “and there is no indication that food resources are limiting in this habitat.”

148 


Mullidae 

Upeneus vittatus (Forsskål, 1775) 

Introduced 

The striped goatfish (or yellowbanded goatfish) was introduced in 1955 on O‘ahu with the releases 

of the Marquesan sardine (Randall, 1981, 1987). Randall (1996) notes that it inhabits muddy bottoms. 

The first specimens were collected in 1976 and 1977 in Kāne‘ohe Bay (Randall, 1981). 

Cichlidae 

Oreochromis mossambicus (Peters, 1852) 

Introduced 

= Tilapia mossambica 

The Mozambique tilapia was intentionally introduced to O‘ahu to control “aquatic plants which clog 

irrigation systems, as a food fish, and as a tuna baitfish” in 1951 and 1952 from Singapore (Brock, 

1960; Randall, 1987). It is now well established in fresh and brackish water on most of the islands 

(Mundy, 2005). Coles et al. (1999a) record it from Pearl Harbor; additonal records include Honolulu 

Harbor, Kewalo Basin, Ala Wai Yacht Harbor (Coles et al., 1999b) and Kāne‘ohe Bay (Coles et al. 

2002a). 

Sarotherodon melanotheron Rüppell, 1852 

Introduced 

= Tilapia melanotheron 

= Tilapia macrocephala (Bleeker, 1862) 

The silvery or blackchin tilapia, native to West Africa, was imported in 1962 by the Bureau of 

Commercial Fisheries for holding in experimental tanks (to study its potential as tuna bait) at Sand 

Island, from stock from the American Museum of Natural History in New York City (Randall, 1987). 

“Al though there was no intentional release to the sea, this tilapia somehow escaped—perhaps 

through a dislodged screen on an outflow pipe to Ke‘ehi Lagoon” (Randall, 1987). Englund (2002) 

reports a date of 1951 in error. 

It is now a very abundant fish in Pearl Harbor (Coles et al., 1999a), the Ala Wai Yacht Harbor 

(Coles et al., 1999b), and Kāne‘ohe Bay; Randall (1987) further notes that “Tuna fishermen are very 

unhappy with its abundance, for they believe it may be feeding in part on young nehu.” Yamamoto 

& Tagawa (2000) note that it is “now one of the most abundant species of tilapia in Hawai‘i, having 

displaced the previously ubiquitous O. mossambicus.” Englund et al. (2000a) refers to this tilapia as 

“the dominant estuarine fish in Pearl Harbor and (the rest of) O‘ahu.” “The blackchin tilapia is one 

of the most harmful introduced fish in low-elevation areas of Hawaiian streams, wetlands, and estuaries 

and likely has caused more negative ecosystem effects in lower Pearl Harbor streams and wetlands 

than any other introduced aquatic species” (Englund et al., 2002a). Mundy (2005) notes that it 

occurs on Kaua‘i as well. 

Pomacanthidae 

Centropyge flavissima (Cuvier, 1831) 


Coles et al. (1999b, p. 47) note that “The only occurrence of the Lemon Peel fish Centropyge flavissima 

in this study was an observation near the [Ala Wai Yacht] harbor boat ramp.” However, Richard 

Pyle (pers. comm.) has sighted this species infrequently over at least the past ten years at Ke‘ehi 

Lagoon, Kewalo Basin, the Ala Wai Canal, and Kāne‘ohe Bay” (see also Mundy, 2005). Coles et al. 

(2002a) also report it from Kāne‘ohe Bay. Richard Pyle informs us (pers. comm., January 2004) that 

over this period of time he has never seen juvenile fish. Randall et al. (1990) give the distribution as 

Cocos-Keeling Islands to southeastern Oceania and north to the Ryukyu Islands, noting that it is 

“common in island groups of the tropical central and western Pacific.” We take the first date of 

occurrence as approximately 1990. It is a probable aquarium release. 

Centropyge loriculus (Günther, 1874) Kāne‘ohe Population Introduced 

Mundy (2005, pp. 43, 83, and 405) reports a population of this otherwise-indigenous flaming 

angelfish in He‘eia Kea Harbor (He‘eia Boat Harbor), Kāne‘ohe Bay, O‘ahu with a distinctive color 

pattern “that are probably the descendents of aquarium fishes that were released in Kāne‘ohe Bay”


149 

(p. 405), although more strongly so assigned in an earlier discussion (“These are known to be the 

descendants of fish released from imported aquarium stock”, p. 83). Coles et al. (2002a) report it 

from near He‘eia Kea Marina, Kāne‘ohe Bay. 

Pomacentridae 

Abudefduf vaigiensis (Quoy & Gaimard, 1825) Waif with Drift Nets? 

This damselfish, known as the “Indo-Pacific sergeant major,” or mamo, was first recorded in the 

Hawaiian Islands in 1991 at Molokini, Maui (J.E. Randall, pers. comm., January 2004). Randall 

(1996) notes that “small breeding colonies have been observed at the islet of Molokini and on 

O‘ahu.” Basch (2002) reports observing A. vaigiensis in a south Maui survey in 2002, and 237 individuals 

were noted in a two day period in October 1998 in a survey of Molokini Shoal [see http: 

//cramp.wcc.Hawai‘i.edu/Study_Sites/Maui/Molokini/1998_Survey/]. Mundy (2005, pp. 21, 425) 

reviews its increasing spread and establishment on a number of islands and north to French Frigate 

Shoals. 

The pattern of arrival and spread has suggested to Hawaiian ichthyologists that this is a “natural” 

colonist that may have arrived associated with drifting nets and debris (J. Randall & J. Hoover, 

pers. comm., January 2004), with which this fish is known to be associated (Kimura et al.,1998, 

Mundy, 2005; see also Donohue et al., 2001). While on the one hand it is unusual that it would have 

failed to arrive as a waif in the Islands previous to the 1990s, the increasing prevalence of high seas 

drifting debris, including derelict fishing gear, might provide a mechanism for transport that was less 

frequent in earlier decades. It is of interest to note in this regard the collection of the Asian sea 

anemone Diadumene lineata on a drift net in the lagoon at Pearl and Hermes Reef in the North - 

western Hawaiian Islands in 2000 (see discussion at that species). 

This fish is occasionally found in the aquarium trade and is sold as an aquarium fish in Hawai‘i 

(see http://www.raingarden.us/marine.htm [accessed January 2007]) and thus may have been an 

intentional release, or aquarium releases may have contributed to part of its modern-day occurrence 

in Hawai‘i. 

Maruska & Peyton (2007) report interspecific spawning between A. vaigiensis and the endemic 

Hawaiian sergeant fish, Abudefduf abdominalis. 

Chrysiptera taupou (Jordan & Seale, 1906) 


Mundy (2005) notes that specimens of this damselfish (known as the “South Seas devil”) have been 

found in 2002 and 2004 in Hanauma Bay, O‘ahu, and possibly in 2002 at Ala Moana Beach Park, 

O‘ahu. The records are from shallow water. Mundy remarked that it “might be established in shoreline 

waters of Honolulu.” 

Blenniidae 

Parablennius thysanius (Jordan & Seale, 1907) Introduced 

This Indo-Pacific blenny (the “tasselled blenny”) was first collected in 1971 in Kāne‘ohe Bay and 

by 1990 had become abundant in “the fouling on buoys floating in about 10 m of water and about 

300 m from shore” (Springer, 1991). In 1990 it was also said to occur on the fouling of wharf pilings 

in Kāne‘ohe Bay, and was reported from Pearl Harbor (Springer, 1991). Greenfield (2003) notes 

that it is found in Kāne‘ohe Bay “in fouling communities ... on dock floats.” 

Parablennius thysanius is known from Pakistan, Sri Lanka, Oman, Thailand, and the Philip - 

pines. Mundy (2005) considered it likely to have been introduced from the Philippines. We consider 

it a ballast water, or sea chest fouling, introduction to O‘ahu. 

Omobranchus ferox (Herre, 1927) 

Introduced 

Englund & Baumgartner (2000) report the establishment of the fang-toothed blenny Omobranchus 

ferox in Pearl Harbor. This fish is native to the Philippines and South China Sea. It was found on a 

rocky shoreline interspersed with mangroves in Halawa Stream in 35–36 ‰; the first specimens 

were collected in 1998. It remains present in Pearl Harbor and the Ala Wai region (R. Englund, pers.

150 


comm., January 2004). Englund & Baumgartner (2000) suggested transport in ballast water or sea 

chest fouling. While not mentioned by Englund & Baumgartner (2000), Mundy (2005, p. 465) notes 

that the site of collection was “near a large, floating dry dock that had been moved to O‘ahu from 

the Philippines a few years earlier ... . This species was most likely introduced to O‘ahu on the hull 

of the dry dock.” This appears to refer to the movement of the dry dock Machinist to Pearl Harbor 

in 1992. Englund & Baumgartner (2000) do mention the incident of a dry dock towed from Subic 

Bay, Philippines to Guam, which arrived with three species of gobies (including Omobranchus elongatus), 

none native to Guam. Englund & Baumgartner (2000) speculate that this introduced goby 

could compete with the native estuarine goby Oxyurichthys lonchotus. 

Omobranchus rotundiceps obliquus (Garman, 1903) Introduced 

= Omobranchus elongatus of Strasburg (1956, 1966) fide Springer & Gomon, 1975, p.69 

Specimens of this Indo-Pacific blenny (known as the mangrove blenny or roundhead blenny) were 

accidentally imported from “near the Samoan Islands” in 1951, when coralline rock, the clam Tri - 

dacna, and other living materials, in which the blenny was hiding, from the island area were placed 

in a concrete tank on the shore of Kāne‘ohe Bay with an open seawater system (Strasburg, 1956). 

By 1963 it had rounded O‘ahu, and a specimen was taken from a reef in Pearl Harbor (Strasburg, 

1966). It remains common in Kāne‘ohe Bay on sheltered patch reefs, and in rocky and mangrove 

habitats but is most abundant in fouling communities (Greenfield, 2003). It remains restricted to 

O‘ahu [Englund & Baumgartner (2000), based upon pers. comm. from J.E. Randall and J.P. Hoover]. 

Gobiidae 

Mugilogobius cavifrons (Weber, 1909) 

Introduced 

= Mugilogobius parvus (Oshima, 1919) fide Larson, 2001 

This Indo-Pacific marine and brackish water goby was collected in 1988 at Coconut Island in 

Kāne‘ohe Bay in a shallow mangrove habitat (Randall et al., 1993). Additional specimens and observations 

then came to hand from Ala Moana drainage canal (1989) and from a Pearl Harbor drainage 

canal (1987). Englund (2002) reports that this goby is now abundant in Pearl Harbor and is common 

in a number of other estuarine streams in western and southern O‘ahu. Mundy (2005) notes that it 

has spread to the Kaua‘i estuaries. 

Randall et al. (1993) regards it as a ballast water introduction, to which we add the possibility 

of transport in sea chest fouling as well. 

Favonigobius sp. 


Greenfield & Randall (2004) report the collection in 1990 of “several small specimens” of this perhaps 

Australasian goby in Kahana Bay. The specimens were about 14 mm in length. Greenfield & 

Randall note that there were “attempts to collect more specimens” and that these attempts were not 

successful; these attempts were in June 1997 using a fine-mesh seine (D.W. Greenfield, pers. comm., 

March 2007). 

The presence of several specimens at a site (Kahana Bay) distant from ballast water release suggests 

that a reproducing population had become established. That said, there have been no further 

known collections since 1990, although small gobies may not attract sufficient attention, nor be 

brought, if collected, to the attention of professional ichthyologists. If this small species is still present, 

searches in nearby Kāne‘ohe Bay might be fruitful.


151 

ALGAE (Seaweeds) 

Introduced and Cryptogenic Algae 



Rhodophyta 

Acanthophora spicifera 1952 SF, R Indo-Pacific 

Eucheuma denticulatum 1970 R Philippines 

Kappaphycus alvarezii 1974 R Philippines 

Kappaphycus striatum 1970 R Philippines, Pohnpei 

Asparagopsis “taxiformis” - Lineage 4 1991 SF? Indo-Pacific 

Gracilaria salicornia

152 


Rich arenas for exploration for introduced or cryptogenic species may be found among the filamentous 

reds and greens in Hawai‘i, many of which bear cosmopolitan names and now require 

genetic analyses. Given the centuries of global shipping and the importation of many animals and 

plants for fisheries purposes, it is difficult to imagine that no smaller algae have been introduced. 

Prominent introduced species include Acanthophora spicifera, Hypnea musciformis, Gracilaria 

salicornia, Kappaphycus spp., and Avrainvillea amadelpha. The spread and growth in abundance of 

most of these species have resulted in extensive ecological and societal impacts, including competition 

with and overgrowth of native algae and coral reefs, fouling, habitat change, and (for Hypnea 

and Gracilaria) masses of rotting seaweed washing ashore. These effects have attracted media and 

public attention since the 1980s. For example, on 24 August 2002, a “Waikīkī Alien Algae Clean-Up 

Event” was held, for volunteers to remove massive amounts of Gracilaria salicornia from the waters 

in front of Waikīkī Aquarium, and to evaluate methods needed to control introduced algae on 

Hawai‘i’s reefs. Five thousand pounds of algae were removed on this day. As of late 2005, the ‘A’ohe 

Limu’e, No Alien Algae program had removed about 100 tons of Graciliaria from the area (see 

http://www.hawaii.edu/malamalama/2006/01/f4_algae.html [accessed February 2007]). 

These efforts were foreshadowed more than 25 years earlier, when a similar but smaller effort 

was made to remove Kappaphycus striatum from an area of Kāne‘ohe Bay, as discussed below. 

Numerous materials—brochures, cards, posters, videos, et cetera—have been produced to in - 

form the public about “alien seaweeds” in the islands. For example, the Hawaii Division of Aquatic 

Resources produced (with the assistance of the “Hawai‘i Marine Algae Group”, or HiMAG) in 2003 

a set of 10 small waterproof plastic cards (held together by an electrician’s tie) on “Alien Seaweeds”, 

with the “title card” describing that “alien seaweeds harm our native marine life, reduce fisheries 

habitat, [and] cost millions [of dollars] to Hawai‘i’s economy.” Five nonnative species were assigned 

common names as follows: 

Graciliaria salicornia 

Avrainvillea amadelpha 

Acanthophora spicifera 

Kappaphycus spp. 

Hypnea musciformis 

Gorilla Seaweed 

Leather Mudweed 

Prickly Seaweed 

Smothering Seaweed 

Hookweed 

The cards advise the public to “help keep these aliens from spreading throughout Hawai‘i”, 

identifying, interestingly, three modes of interisland dispersal: by boats, by fishing, and by recreational 

diving. Individuals are asked to remove seaweed from anchors, mooring lines, outboard propellers, 

and the bilge, and by keeping boat hulls clean; to inspect fishing lines, hooks, nets, traps, and 

“catch bags” for “small fragments of seaweed”, and, while snorkeling or diving, to inspect wetsuits, 

footwear, gloves, buoyancy compensator, fins, and gear bags. Another card (“What You Can Do to 

Help Protect Hawai‘i’s Coral Reefs from Alien Seaweeds”) further advises the public to “get in the 

habit of thoroughly drying your dive gear, wetsuits, and dive bag after each use“, to “take steps to 

decrease overfishing. Many reef fish eat seaweed and help keep them from overgrowing the reef,” 

and “Never dump aquarium animals or plants into streams or coastal waters.” 

Recent eradication attempts of Eucheuma, Gracilaria, and other species have focused on an 

under water removal technique using barge-mounted vacuum cleaners (Holden, 2007), causing 

Goreau (2008) to question its efficacy, leading to rationales for eradication attempts by Smith et al. 

(2008) in a high-profile journal, Science. Smith et al. (2008) noted that “areas cleared by the Super 

Sucker remain clear of alien algae and have increased coral recruitment after just 2 years,” noting 

that removal strategies buy “time for the development of biological control methods with the native 

sea urchin Tripneustes gratilla,” citing Hawaiian studies by Stimson et al. (2007) on the potential for 

this urchin to feed on Gracilaria and other species. 

Vroom et al. (2006) found no nonnative algae at French Frigate Shoals, Northwestern Hawaiian 

Islands, although all four Ulva species occurring there (clathrata, flexuosa, paradoxa, and prolifera) 

we regard as of uncertain history in the Islands.


153 

Vermeij et al. (2009) experimentally studied the effects of herbivorous fish on Acanthophora 

spicifera and Hypnea musciformis on O‘ahu and Maui, and in their native Caribbean habitats, finding 

that in Hawai‘i these algae experienced reduced herbivore pressure, helping to facilitate their 

success. 

RHODOPHYTA (red algae) 

Rhodomelaceae 

Acanthophora spicifera (Vahl, 1802) Borgesen 1910 Introduced 

Doty (1961) reported the discovery of this Indo-Pacific and cosmopolitan “spiny seaweed” based 

upon specimens discovered in 1952 in Pearl Harbor. Although with a Caribbean type locality (St. 

Croix, Virgin Islands), it is now widespread globally “throughout tropics into warm temperate areas” 

(Abbott, 1999). How much of this now cosmopolitan distribution is due to ship-mediated dispersal 

appears not to have been investigated. Doty reported the following chronology of its early discovery 

and spread as follows: 

Early Discovery and Spread of Acanthophora in Hawai‘i, 

1952–1960 

Date Location Comments 

1952 (fall) O‘ahu south (leeward) shore: a small fragment 

Pearl Harbor 

1953 (April) O‘ahu south shore: Waikīkī Beach “much larger piece” 

1953 (May) O‘ahu south shore: Ke‘ehi Lagoon in abundance 

(between Pearl Harbor and Port 

of Honolulu) 

1953 (May) O‘ahu north (windward) shore: dense growth 

Hau‘ula 

1953–1956 O‘ahu south shore “so common that it has not often been 

preserved” 

1954–1955 Kaua‘i 

1956 (February) O‘ahu: north shore: Kāne‘ohe Bay substrate for eggs of native snail Conus 

quercinus 

1956 (June) O‘ahu northwest shore: Mokuleia “ubiquitous some distance within the 

reef”, very rare in 1955 

1956 (November) Kaua‘i 

1960 (August) Kaua‘i: Po‘ipū Beach 

1960 (November) Lāna‘i north shore washed ashore in abundance, “free or 

attached to shells, coral, or even rocks 

up to 2 lb. in weight” 

Doty (1961) felt that this alga arrived in hull fouling and specifically suggested that a heavily fouled 

barge, the Yon 146, which arrived in Pearl Harbor in February 1950 may have been the actual vessel 

involved. Russell (1987) also suggested that it may have arrived with algal-covered substrates 

imported with fish to the Waikīkī Aquarium. 

Acanthophora is now the most abundant and ubiquitous introduced alga in the Hawaiian Islands 

(Smith et al., 2002). It is common in many habitats in shallow and intertidal reefs. Russell (1992) 

noted that Acanthophora often outcompetes the native algae Laurencia spp. and Hypnea cervicornis. 

Smith et al. (2002) note that “it is most common in intertidal regions and in semiprotected tide 

pools, where it may escape spatially from herbivory. This species was commonly observed fouling 

ship hulls in harbors throughout the state during field surveys. Although this plant was a common 

component of the marine flora and is clearly displacing native species in Hawai‘i, it does not appear 

to form large, monospecific nuisance blooms.”

154 


Stimson & Larned (2001) found that the native green “bubble” alga, Dictyosphaeria cavernosa 

remained abundant in parts of Kāne‘ohe Bay, despite nutrient-enriching sewage abatement in the 

1970s, due to decreased herbivory: herbivorous fish prefer introduced algae such as Acanthophora, 

reducing predation on adjacent populations of Dictyosphaeria. 

Russell & Balazs (1994) documented the green sea turtle’s (Chelonia mydas) use as food of 

Acanthophora and Hypnea in Hawai‘i. O’Doherty & Sherwood (2007) provide the first analysis of 

the genetic population structure of A. spicifera in Hawai‘i, reporting that populations in the islands 

form discrete genetic groups. 

Solieriaceae 

In an act that would not be permitted today, the State of Hawaii issued permits in the 1970s for the 

late Professor M.S. Doty of the University of Hawaii to import and release three species of eu - 

cheuma, largely from the Philippines, into Kāne‘ohe Bay, for the purposes of open water farming for 

the production of kappa-carrageenan and agar. All three have become established; a fourth, Eu - 

cheuma isiforme, did not (Russell, 1992). The importation data from Russell (1992) are given below 

each species. 

Eucheuma denticulatum (Burman, 1768) Collins 

et Harvey, 1917 

Introduced 

Released into Kāne‘ohe Bay, Honolulu Harbor, and other O‘ahu sites from October 1970 to “late 

1976” stock from the Philippines. The type locality of E. denticulatum is the Indian Ocean, and it 

also occurs in the Western Pacific. 

Kappaphycus alvarezii (Doty, 1985) Doty ex Silva, 1996 Introduced 

= Eucheuma striatum “tambalang variety” of Russell, 1983, and others 

Released into the above sites from September 1974 to “late 1976”; stock from the Philippines. The 

type locality of K. alvarezii is Malaysia, and it also occurs in the Sulu Sea and southern Philippines. 

Kappaphycus striatum (Schmitz, 1895) Doty 

ex Silva, 1996 

Introduced 

Released into the above sites from August 1970 to “late 1976,” stock from the Philippines and 

Pohnpei. The type locality of K. striatum is also the Indian Ocean, and it occurs widely in Indonesia, 

the western Pacific, and Micronesia (Abbott,1999). Kappaphycus striatum also went under the earlier 

name of Eucheuma cottonii (Abbott, 1999), and was also known as the “elkhorn” variety. 

Kappaphycus alvarezii was also known as the “tambalang” variety (Russell, 1983, in that paper as 

Eucheuma striatum; Eldredge, 1994). 

Doty (1973a, 1973b, 1977) touted the commercial, economic, and social values of introducing 

and growing eucheuma, even referring to the “rational utilization of the subtidal environment” of 

Kāne‘ohe Bay (Doty, 1977, figure 3 caption). Doty (1977) presented maps of the sites where 

eucheuma had been released and growth rate data for Kappaphycus alvarezii (as the “tambalang 

strain of Eucheuma”). He also discussed post-release control measures and the government bureaucracy 

that would constrain releases of nonnative species. 

While E. denticulatum and K. striatum had been imported as early as 1970, it was the later 

importation of K. alvarezii that caused initial concern. Material of this species from the Philippines 

imported in September 1974 (Russell, 1983, as E. striatum—tambalang form) and/or May 1975 

(Russell, 1979). Initially placed in wire holding pens on the northwestern reef bordering Coconut 

Island in Kāne‘ohe Bay, the pens broke down and the algae were freed (A.H. Banner in litt. to L.G. 

Eldredge, October 1982). By March 1976 an estimated 50 to 100 metric tons were on the reef 

(Russell, 1979) resulted in a day-long effort, organized by Dr. E. Reese (A.H. Banner in litt. to L.G. 

Eldredge, October 1982), that found 50 volunteers removing about four tons of seaweed (Russell, 

1983). Russell (1983) and Glenn & Doty (1990, 1992) described aspects of the ecology and growth 

of euchemas in Kāne‘ohe Bay. Zuccarello et al. (2006) provide recent data on the genotypes of 

Hawaiian euchemas.


155 

All three species remain in Kāne‘ohe Bay, and at least two, K. alvarezii and K. striatum abundantly 

so, having spread within the Bay but not outside of it (Rodgers & Cox, 1999, for K. alvarezii 

and K. striatum). Smith et al. (2002) note that Kappaphycus spp. in Kāne‘ohe Bay are “clearly competitively 

dominant, occupying up to 80% of the substrate. Because of its large structure, Kappa - 

phycus also appears to be competing with coral and may be able to overgrow live coral colonies.” 

Conklin & Smith (2005) provide further data on the increased spread through Kāne‘ohe Bay of 

Kappaphycus spp. between 1999 and 2002, noting that it “can be found in a variety of reef habitats 

overgrowing and killing corals.” They suggest that management action be undertaken, given that 

Kappaphycus continues to spread and can have over 50% cover on some coral reefs. 

Bonnemaisoniaceae 

Asparagopsis “taxiformis” — Lineage 4 

Introduced 

As Abbott (1999) notes, this well-known red alga has a long history in Hawaiian culture, and has 

been formally recorded in the flora since the 19th century. With a type locality of Alexandria, Egypt, 

and said to occur in all oceans, it is not surprisingly a species complex (Andreakis et al., 2007). 

Sherwood (2008) establishes that one of these lineages appears to be a relatively new introduction: 

while two other clades of A. “taxiformis” occur relatively widely in the Islands, lineage 4 is restricted 

to a localized area on the south shore of O‘ahu. The first record of this lineage is 1991 (Andreakis 

et al., 2007). It is otherwise known from tropical and subtropical regions in the Indo-Pacific ranging 

from Sri Lanka to Panama and Costa Rica (Sherwood, 2008). 

The other two lineages have curious distributions: lineage 1 occurs in Hawai‘i and the Pacific 

coast of Central America, whereas lineage 2 is known from the Indo-Pacific and the Mediterranean, 

as well as Hawai‘i (lineage 3 does not occur in Hawai‘i).This pattern is suggestive of earlier invasions, 

whose directions and tempo remain to be worked out. 

Ceramiaceae 

Wrangelia bicuspidata Borgesen, 1916 

Failed(?) 

Russell (1992) notes that this tropical alga, known from the Indo-Pacific and the Caribbean, was discovered 

in Kāne‘ohe Bay in about 1974 (as “1974 (?)”) and was established (“successful”). Abbott 

(1999), however, does not further report it from Hawai‘i and reports to us (I. Abbott, pers. comm., 

April 2000 and January 2004) that there are no recent collections. 

Graciliariaceae 

Gracilaria salicornia (C. Agardh 1820) Dawson, 1954 Introduced 

Smith et al. (2002) suggest that this Indo-Pacific alga is not native to the Hawaiian Islands, noting 

that prior to its intentional transplantation from the Big Island of Hawai‘i to O‘ahu, it was known 

from only two populations on Hawai‘i, one in Hilo Bay and another in Kapoho “before 1950.” They 

note that “the origin of these populations is unknown but may be tied to early harbor arrivals in Hilo 

from the Philippines.” Smith et al. (2004, page 327) remark that “There is some support for the idea 

that this Hawai‘i Island population was an early twentieth-century ballast introduction from ships 

originating in the Philippines ... .” 

In April 1971 and again in September 1978, this alga was transplanted intentionally for aquaculture 

purposes from Hawai‘i to Waikīkī and Kāne‘ohe Bay on O‘ahu (Russell, 1992), where it 

became established (Abbott (1999) gives these dates as “about 1980”, but see Russell, 1992). 

However, Abbott (1999) notes the presence of G. salicornia at Kaloko, on the eastern shore of 

O‘ahu, in 1946, prior to its later transplantation. Smith et al. (2002) further note that “sometime in 

the 1980s” G. salicornia was transplanted from O‘ahu to Puko‘o fishpond on the island of Moloka‘i, 

based upon information provided by I.A. Abbott. Abbott (1999) further reported it from Ala Moana 

Park. Rodgers & Cox (1999) discuss the rate of spread of this species and its distribution in 

Kāne‘ohe Bay. 

Smith et al. (2002) report as follows: “This nonindigenous algal species currently has the most 

discontinuous distribution of all species examined in this study. It is now found on three islands, with

156 


no obvious continuity among locations. At most of the sites where G. salicornia was found, this alga 

was highly dominant over a distinct area. It was very common in southern Kāne‘ohe Bay but was 

not found in the north bay. Gracilaria salicornia was dominant in Waikīkī in front of the Aquarium 

but was not present at adjacent sites such as Ala Moana Beach Park or at Kahala. It seems that, once 

introduced, this species may have the ability to spread within a site laterally and become locally 

dominant but does not have great success at dispersing larger distances between sites or islands, over 

this two-decade time frame.” 

Smith et al. (2004) report on the ecology of this invasion on O‘ahu, noting that 30 years after 

its release on two reefs in the 1970s, “this species has spread from the initial sites of introduction 

and is now competing with native marine flora and fauna.” They note that “substantial amounts of 

G. salicornia become dislodged from the reef during large wave events and periodically become 

deposited onto the beach in front of the Waikīkī Aquarium ... over 20,000 kg of alien algal fragments 

were removed from this location in five 4-hr cleanup events.” 

Conklin & Stimson (2004) add herbivorous fish to Gracilaria reefs at Kāne‘ohe Bay to experimentally 

address herbivory as a means of controlling fleshy macroalgae, but found that fish did not 

remain on site due to lack of shelter. 

Gracilaria tikvahiae McLachlan, 1987 

Introduced 

Russell (1992) reports that this Western Atlantic Ocean alga had been intentionally imported for 

commercial mariculture purposes from Florida and released in Kāne‘ohe Bay and Kahuku on O‘ahu 

in the “mid 1970s.” Abbott (1999) reported the introduction as having occurred in 1987, “to help 

alleviate the low market supplies of G. parvispora.” Abbott (1999) reports further unidentified 

Gracilaria existing in the University of Hawai‘i Fisheries Laboratory (‘Ānuenue) and in the Kahuku 

mariculture operations “that are not like G. tikvahiae”, and for which “it is uncertain how many 

species are involved, or if any are native.” Abbott notes a collection of G. tikvahiae from near the 

seawater discharge of the Oceanic Institute near Makai pier in the vicinity of Makapu‘u Point, 

O‘ahu. 

Hypneaceae 

Hypnea musciformis (Wulfen in Jacquin, 1791) 

Lamouroux, 1813 

Introduced 

This seaweed, which is largely an epiphytic species with recurved hooks on its branch tips, was 

transplanted from Florida to Kāne‘ohe Bay in January 1974 (Russell, 1992) “as part of an aquaculture 

project that was later abandoned” (Smith et al., 2002). It apparently came as part of a shipment 

of Eucheuma, but was then interpreted as a potential farming candidate (Russell & Balazs, 1994, p. 

54, “Hypnea musciformis came with a shipment of two morphs of Eucheuma … .”). Russell & 

Balazs (1994) further noted that Hypnea became so abundant in Kāne‘ohe Bay that it began growing 

among Eucheuma that were then exported to other countries in the Pacific, resulting in the yet 

further spread of this seaweed. It is now one of the most pestiferous seaweed invaders in Hawai‘i. It 

occurs in the Philippines and Indian Ocean and from the Caribbean to Uruguay; curiously enough, 

its type locality is Trieste, Italy (Abbott,1999). 

Abbott (1987) reported its early history from 1974 to 1982 as follows: 

“This past summer [1987] many residents of Kuau, a coastal section just southeast of Paia, Maui 

Island, were up in arms over the appearance of masses of a ‘new’ seaweed on their beaches. Indeed, 

the culprit alga was new to Maui, having come over from Oahu Island where it had been spreading 

since its introduction to Kaneohe Bay about 1974. The red alga alien is Hypnea musciformis ... . 

Stories about its introduction are not agreed to by all parties, one saying that it was accidental, having 

come with another species of algae as an epiphyte from the Caribbean. I began to record its spread 

in the fall of 1978 when I found it at Kaaawa (not far from the mouth of Kaneohe Bay), and in the 

fall of 1979 it was a Laie Point; in 1980 at Waikiki; 1981 at Ewa Beach and by 1982 ... it seemed to 

have reached every nook and cranny of intertidal Oahu.” 

Abbott (1987) further notes that it made up between one-third and one-half of the wet biomass of


157 

drift algal piles on O‘ahu beaches, and further suggested that it might account “for larger masses than 

previously of Sargassum on the beaches, since it adds drag and weight to the Sargassum plants. 

Sargassum was frequently in the drift in late summer and fall in pre-Hypnea times.” 

Russell & Balazs (1994) detail the spread of Hypnea around O‘ahu, Moloka‘i, Maui, and Lāna‘i 

from 1977 to 1991. They note it was found as an epiphyte on Sargassum echinocarpum, S. polyphyllum, 

and Acanthophora spicifera, or in windrows of beach algal drift. Russell & Balazs (1994) further 

documented the use as food of Hypnea and Acanthophora by the green sea turtle Chelonia 

mydas in the Hawaiian Islands, noting that it was the first documentation of introduced algae being 

incorporated into the diet of this turtle. Hypnea was also considered to be inhibiting the growth of 

the native Hypnea cervicornis and Laurencia nidifica. 

Abbott (1999) remarks that by 1987 it had become established near Pa‘ia, Maui, “where it is 

most commonly found in large nearly unialgal masses cast ashore in windrows up to 0.5 m depth on 

many beaches and considered to be an odiferous pest by local inhabitants and tourists alike.” Obser - 

vations in 1996 on windward and leeward beaches of Maui revealed that Hypnea formed at least 

two-thirds of the drift algal biomass. In 2003, the Environmental Protection Agency provided 

$250,000 to purchase a seaweed-removal machine to be used on Maui (Carroll, 2004). 

Smith et al. (2002) reports that this alga was the second most common introduced algae in the 

islands, although it had not yet spread to all of the islands, and was not as abundant as Acanthophora, 

although it was becoming more common. Abbott (1999) also notes that it was not yet on either 

Kaho‘olawe or Hawai‘i. It blossoms at “discrete locations” and when abundant co-occurs with Ulva 

fasciata, “a known weedy species in a genus known to require high nutrient flux for growth” (Smith 

et al., 2002). On the basis of this, Smith et al. (2002) suggest that Hypnea blooms may be related to 

land use activities and nutrient input. Hypnea propagates vegetatively, especially among smaller 

algal pieces. 

Huisman et al. (2007) note that in 2003 and 2004, H. musciformis was found at Necker Island 

(Mokumanamana), “probably carried there by currents. Managing this potentially destructive invasive 

species is a major challenge.” 

Gigartinaceae 

Mazzaella volans (C. Agardh, 1820) J. Agardh, 1846 Escaped; Not Established 

Abbott (1999) reports that this eastern Pacific red alga, known from Oregon to Mexico, was collected 

at the outfall of experimental aquaculture tanks holding the California kelp Macrocystis, at 

Keāhole Point, Kona, Hawai‘i. The material was collected by W.H. Magruder on 5 April 1987 (I. 

Abbott, pers. comm., April 2000). Abbott (1999) writes that “Keāhole Pt. is the location of several 

commercial aquaculture operations. Spores of this common western North American species could 

have come with Macrocystis, imported to feed abalone (Haliotis sp.), also imported for mariculture 

purposes, which might be a source of spores as well.” Isabella A, Abbott reports (pers. comm. 

January 2004) that it has been looked for since 1987 and not found (see also Abbott & Huisman, 

2004, page 239). We consider it not established. 

CHLOROPHYTA (green algae) 

Udoteaceae 

Avrainvillea amadelpha (Montagne) A. Gepp 

Introduced 

& E. Gepp, 1908 

Brostoff (1989) report that the first collections of this Indo-Pacific (Abbott & Huisman, 2004) 

species were in 1981 (a three square meter patch in 13 m of water at Kahe Point, O‘ahu, on sandcovered 

dead coral) in 1985 (from sand and rock substrates in 10 m at Maunalua Bay, O‘ahu), and 

in 1987 and 1988 (intertidal zone at Hawai‘i Kai). While Brostoff notes that “ship introduction is 

unlikely”, the transport of drift material in ballast water may be possible, either as parts of adult 

plants torn loose from the substrate, or possibly as gametes. Alternatively, anchors or anchor chains 

with entangled specimens may have provided a vector from the Indo-Pacific.

158 


Smith et al. (2002) report that it has persisted and “has spread laterally from Koko Head to 

Kahala on Oahu’s south shore from Kahe Point north on the west shore.” They also newly report it 

as a “small population” at Prince Kuhio Beach Park on Kaua‘i. “This species inhabits soft or sandy 

bottom habitats where the majority of the plant biomass is subsurface. This nonindigenous alga frequently 

serves as a substrate for many native species of epiphytic algae and as habitat for many 

invertebrates. The endemic Hawaiian sea grass Halophila hawaiiana and A. amadelpha now cooccur 

in areas that were once H. hawaiiana meadows. This may prove to be a considerable conservation 

and management problem, and more research is needed to determine A. amadelpha’s ecological 

strategies and impacts on the native biota” (Smith et al. 2002). 

Ulvaceae 

Rich grounds for exploration of cryptogenic and introduced species are the species of Ulva, taxa well 

known for centuries to be associated with ship fouling and oyster shells, and easily transported as 

smaller pieces in ballast water. Included are species formerly assigned to Enteromorpha, a genus 

now synonymized with Ulva (Hayden et al., 2003). Species assigned to both genera, as well as a 

number of other brackish-water green algae, have long been known from the islands (Abbott, 1947, 

1980; Abbott & Huisman, 2004). 

We regard the following Ulva as cryptogenic, with the exception of Ulva expansa. We include 

species of Ulva with native Hawaiian names that have been long-embedded in Hawaiian culture: 

here, our consideration is that if these species were present on the Islands aboriginally, modern-day 

populations may have been genetically altered and modified by the influx of global genomes over 

the past 200 years. Geographic ranges and unattributed quotations below are all from Abbott & 

Huisman (2004): 

Ulva clathrata (Roth, 1806) Greville, 1830 

All temperate seas; “intertidal and more exposed areas.” 

Cryptogenic 

Ulva compressa (Linnaeus, 1753) Nees, 1820 

Cryptogenic 

Warm and temperate seas; “mostly on shallow nearshore rocks or limestone.” 

Ulva fasciata Delile, 1813 

Cryptogenic 

Known as limu palahalaha and lipahapaha, this Ulva is one of only three species of green algae used 

in traditional Hawaiian food preparation; the others are Ulva prolifera and Codium edule (Abbott & 

Huisman, 2004). Occurring over a wide variety of habitats, it is “the commonest sea lettuce in the 

Islands” If introduced, it is one of the earliest documented invasions, having been found in 1819 by 

C. Gaudichaud (St. John & Titcomb, 1983). Widespread in the Pacific and Indian Oceans, it is also 

recorded from the Mediterranean and Caribbean (Abbott & Huisman, 2004). Aguilar-Rosas et al. 

(2005) regard it as introduced to the Pacific coast of Mexico, with the earliest specimens from there 

being collected in 1970, suggesting that this species is transportable by human activity. 

Aguilar-Rosas et al. (2005) noted that U. fasciata “is considered as an introduced species” in 

Hawai‘i (and Australia), citing several references, none of which mention its introduction to Hawai‘i. 

Raúl Aguilar Rosas informs us (pers. comm., February 2007) that they intended to say that U. fasciata 

was considered an invasive native rather than introduced, citing the “Marine Algae of Hawai‘i” 

web page of the University of Hawai‘i (http: //www.hawaii.edu/reefalgae/invasive_algae/chloro/ 

chlorophyta.htm). 

Ulva flexuosa (Wulfen, 1803) J. Agardh, 1883 

Cryptogenic 

= Ulva tubulosa (Kutzing, 1856) Kutzing 

= Ulva lingulata of Hawaiian authors 

Temperate and tropical seas of Atlantic and Pacific Oceans; “Attached to sandy high to mid-intertidal 

rocks, to 2 m deep, with freshwater intrusion; epiphytic in brackish fishponds.”


159 

Ulva intestinalis (Linnaeus, 1753) Nees, 1820 

Cryptogenic 

Temperate and tropical seas; “Brackish water to marine; in canals, ponds, on rocks, often floating.” 

Ulva linza (Linnaeus, 1753) J. Agardh, 1883 

Cryptogenic 

Temperate seas, not common in tropics; “Shallow quiet water; drift.” 

Ulva paradoxa (C. Agardh, 1817) Kutzing, 1845 Cryptogenic 

Temperate and tropical seas; “Intertidal rocks, tidepools; also in brackish water.” 

Ulva prolifera (O. F. Muller, 1778) J. Agardh, 1883 Cryptogenic 

Alaska to Mexico; Australia to Japan; “Intertidal sandy rocks, concrete pilings, breakwaters, and in 

brackish water.” With a native Hawaiian name, limu ‘ele‘ele. 

Ulva reticulata Forsskål, 1775 

Cryptogenic 

Widespread in Pacific and Indian Oceans and Red Sea; “Intertidal, epiphytic; common in drift, rare 

on rocks, shallow subtidal to deepwater (dredge collection from 91–128 m).” 

Records suggest that there were populations of this species on the islands in the 19th century, 

but there may have been one or more introductions during or since World War II, accounting for its 

easier detection by the 1950s. The earliest records are “prior to 1882” (exact location on the islands 

not known), followed by specimens collected in the 1920s and 1930s (Abbott & Huisman, 2004). 

Unaware of these records (unpublished until 2004), Gilbert (1962), in first reporting Ulva reticulata 

from O‘ahu, Maui, Kaua‘i, Hawai‘i, and Moloka‘i, all based upon collections he made in 1959, 

remarked that “It is difficult to understand why this well-known Ulva has not been reported previously 

from Hawaii. I found it one of the commonest elements in the green algal flora of the five 

islands I visited.” Doty (1973a) noted that he had “made a particular search for this species in the 

world’s herbaria and among local collections for specimens from the islands. The specimen having 

the earliest date, 1933, was found in a book given the author by Dr. Otto Degener.” Doty (1973a) 

reported specimens from 1945–1946 from the leeward side of Oahu. He further noted that the 

1945–1946 specimens had “very slender” thalli “in contrast to what is found today,” suggesting (p. 

185) that it had changed in form over time. Abbott & Huisman (2004) note, however, that specimens 

from the 1920s and 1930s “are similar to those found today, evidence that the species is not of recent 

introduction and has not changed form in recent years, as was suggested by Doty.” Doty (1973a) 

made the further interesting remark that “For a long time only one species, Ulva fasciata, was found 

in Hawaii,” but this proves to be incorrect (Abbott & Huisman, 2004). 

Ulva rigida C. Agardh 

Cryptogenic 

“Pacific coast of North America, Chile, Australia; Indian Ocean; other warm seas”; “Basalt boulders; 

usually only few plants present in the same location.” 

Ulva taeniata (Setchell) Setchell & Gardner 

Cryptogenic 

American Pacific coast from Oregon to central California; Australia and New Zealand; “basalt rocks 

on exposed shores.” 

Ulva expansa (Setchell, 1905) Setchell & Gardner, 1920 Introduced 

First recorded from the islands by Tilden (1901, as Ulva lactuca laciniata, vide Abbott & Huisman, 

2004), who collected it in 1900 at Waialua, O‘ahu on the reef at low tide. It has the most disjunct 

distribution of all of the enteromorphoid and ulvoid Ulva species in the islands, otherwise being 

known only from the Pacific coast of North America (southern British Columbia to La Paz, Baja 

California), as opposed to the broad temperate, subtropical, and tropical distribution characterizing 

most of the other ulvacean species noted here (Abbott & Huisman, 2004), who also note a native 

Hawaiian name for U. expansa, limu pakaiea.

160 


Ulva expansa is now widespread through the archipelago, where it apparently sinks to considerable 

depths (to 200 m) at times (Abbott & Huisman, 2004). Gilbert (1965) noted that it formed “a 

distinct zone at high tide level on cement wall of pier running out from Natatorium-end of Kuhio 

Beach, Waikiki.” Two physiological conditions argue against its naturally surviving the many 

months it would take to be transported naturally on floating objects from the American Pacific coast 

to the islands: (1) its inhibition to ultraviolet-B light (Grobe & Murphy, 1994), as would be encountered 

in ocean surface waters, and (2) its weak holdfast, leading to its blades soon becoming detached 

(Abbott & Huisman, 2004). 

Cladophoraceae 

Chaetomorpha indica (Kutzing, 1849) Kutzing Cryptogenic 

A species of the “warm Pacific” (including Australia) and the Indian Ocean (its type locality), newly 

recorded for Hawai‘i by Abbott & Huisman (2004) from O‘ahu [Hawai‘i Kai, Ka‘alawai, and 

Wai‘anae Boat Harbor, and from Maui (Lahaina Boat Harbor)]. It was found “floating in 2 m depth 

near shore, and in a small boat harbor” (Abbott & Huisman, 2004), suggestive of the possibility of 

recreational boat traffic carrying it to the islands. Additional material in the Bishop Museum collections 

comes from the Big Island (Coconut Island, Hilo, 2003). The earliest material appears to be the 

Hawai‘i Kai material from 1988 (BISH 635517). It is curious that if present naturally it was not collected 

prior to the 1980s and is not more widespread through the Islands. 

Abbott & Huisman (2004) tentatively assign Gilbert’s (1965) record of C. indica (collected in 

1959 at Hau‘ula Park, O‘ahu) to C. capillaris. 

PHAEOPHYTA (brown algae) 

Lessoniaceae 

Macrocystis pyrifera (Linnaeus, 1771) Agardh, 1820 Escaped; Not Established 

This well-known giant kelp from the California coast was introduced into aquaculture operations 

at Makapu‘u in 1972 and Keāhole Point in 1980 (Russell, 1992). While Russell (1992) notes 

that the commercial value of this alga is for alginates, Abbott (1987) remarked that Macrocystis was 

“the food of choice for the abalone that are being grown under aquaculture conditions near Keahole 

Point.” Of interest is that Macrocystis escaped from the facility, presumably in the early to mid 

1980s, and “occasionally small sporophytes (the spore-bearing kelp plant) [were] seen” (Abbott, 

1987). However, mechanical eradication was undertaken (“such plants have been physically 

removed” before spore-production stages were reached, Abbott, 1987, p. 61). There appears to be little 

about this incident in the published literature. 

Dasyaceae 

Nemacystus decipiens (Suringar, 1872) Kuckuck, 1929 Introduced 

Doty (1973a) reports this Indo-Pacific species as introduced from the Western and South Pacific (the 

Japanese name is mozuku). It was first collected in the islands in May 1963 on O‘ahu at Swansea 

Beach Park, just north of Kāne‘ohe Bay [Russell (1992) gives a date of “1950s?”, meaning a possible 

time of introduction]. Doty (1973a) provided a history of spread from Kāne‘ohe Bay and the 

Makapu‘u region to the south, noting that it became abundant “within three years.” In 1970 it was 

collected at Diamond Head Beach Park and in 1971 it was abundant on Sargassum echinocarpum 

near the seaward edge of the Waikīkī reef. Doty (1973a) provided quantitative abundance data for 

the Waikīkī area in 1971. 

Abbott & Huisman (2004), however, felt that the species is “certainly an indigenous alga and 

should not be listed as an alien species;” they do not cite Doty’s 1973 paper regarding its history but note 

that Russell (1992) had listed it as an alien species. They report collections from Midway to Moloka‘i,


161 

and suggest that it is native based upon the “numerous collections made throughout the Hawaiian chain.” 

Broad distributions, especially for species that could drift over decades amongst the islands, do not provide 

evidence for or against a species’ recent biogeographic history. We retain it as a potential invasion. 

This relatively large alga does not appear to have been known prior to the 1960s on O‘ahu and would 

not likely have been overlooked (R.T. Tsuda, pers. comm., 2005). Unlike many other potential cryptogenic 

algae in the Hawaiian Islands, N. decipiens is not known from any of the nearest island systems, 

such as Micronesia, and otherwise occurs either in China-Japan or Australia-New Zealand, as well as 

southwest Asia and the Indian Ocean (see http://www.algaebase.org/speciesdetail.lasso?species_id 

=3948&sk=60&from=results&-session=abv3:89A504B3032cf22D26Xth22AA782 [accessed February 

2007]). 

The mode of introduction of this alga is unclear, but it is a popular edible, cultivated seaweed, 

and may have been intentionally released. 

Pilayellaceae 

Pilayella littoralis (Linnaeus, 1753) Kjellman, 1872 Introduced 

Abbott & Huisman (2004) report this widespread species (“warm and cool temperate seas”) from 

pier pilings in Honolulu Harbor, O‘ahu. They also note records of this species “from hulls of ships 

traveling between the western Pacific and Pacific North American ports”, with stops in Hawai‘i. 

There is no date of collection for the Honolulu pier piling record. The specimen number cited for the 

Honolulu piling record, BISH 773638, refers to the collections noted by Abbott & Huisman (2004) 

from visiting barges (“ships”), and the material that forms the basis of the Honolulu record cannot 

be located at the Bishop Museum as of September 2005. We regard it as a ship-introduced fouling 

species. 

Dictyotaceae 

Dictyota flabellata (Collins, 1901) Setchell 

Introduced 

& N.L.Gardner, 1924 

Godwin (2003) reports that this American brown alga “was documented to have survived and 

become established in a discrete area of Barbers Point Harbor,” on the south shore of O‘ahu. The 

alga arrived on the hull of a floating dry dock towed from San Diego to Barbers Point Harbor in 

December 1999. Colonization of the adjacent harbor retaining wall was observed in 2000 (Abbott & 

Huisman, 2004): “in collections made six months after arrival, this species was growing ... in the surrounding 

environment.” Abbott & Huisman (2003) report that “fertile specimens ... were noted and 

collected in the spring of 2002, 2 years after arrival” and that it has a “pervasive presence in the area 

around the barge near Barbers Point.” It was still present in 2006 (S. Godwin, pers. comm., February 

2007), and we consider it established. 

Dictyota flabellata is recorded from central California to Panama (Abbott & Huisman, 2004), as 

well as Japan and Pakistan (Abbott & Huisman, 2003), and Chile (Ramirez & Santellces, 1991), the 

latter three locations, if the species is correctly identified, certainly representing additional invasions. 

Sargassaceae 

Sargassum muticum (Yendo, 1907) Fensholt, 1955 Intercept 

Abbott & Huisman (2003) report this species as an “accidental introduction”; Abbott & Huisman 

(2004) list this species as a “new record” for the islands. It was collected on a barge towed from 

southern California (where the species is common, and introduced there from Asia) to Pearl Harbor 

in 1999 (Godwin, 2003b). It has not been found in the wild in Hawai‘i. We consider it a “ship intercept,” 

one of hundreds of such species of marine life that arrive on ships’ hulls (or in ships’ ballast 

water) that could be listed as “new records” for the islands. We do not further consider it here.

162 


ANTHOPHYTA (flowering plants) 

Introduced Flowering Plants 


Halophila decipiens 1979 BW?/AN? Indo-Pacific, Caribbean 

Spergularia marina 1909 SBA, R Eurasia-North America 

Pluchea indica 1915 SBA, R Asia 

Pluchea carolinensis 1931 SBA, R Caribbean, S America 

Batis maritima 1859 SBA Caribbean 

Atriplex semibaccata 1895 R Australia 

Atriplex suberecta 1923 R Australia, South Africa 

Typha latifolia 1979 R Eurasia, North America 

Paspalum vaginatum 1936 R Unknown 

Bruguiera sexangula 1922 R Philippines 

Rhizophora mangle 1902 R Florida 

Conocarpus erectus


163 

reports around the world of H. decipiens since the 1970s (Kuo et al., 1995), and particularly its 

arrival in Japan, where it was first detected in 1990 (Kuo et al., 1995). These may be suggestive of 

range expansions induced by climate change. The arrival of H. decipiens in the Hawaiian Islands is 

nestled in this global pattern. 

Of no small value would be reexamination of pre-1970s herbarium material of the native seagrass, 

Halophila hawaiiana Doty & Stone, 1966 (which prior to the 1960s will be in most herbaria 

collections under the name H. ovalis) to determine if H. decipiens is present, or if present when it 

first appears in collections. We predict that no H. decipiens will be represented in 19th century or 

early 20th century collections of Halophila from Hawai‘i. 

The mechanism of introduction of this species to the islands is unclear, but introduction via 

anchors or anchor chains by visiting yachts or other vessels may be possible. Anchors and chains 

often “capture” benthic plants, which can get deeply embedded and wrapped around these structures 

(J.T. Carlton, pers. observ.). Vessels having last anchored in H. decipiens source areas in the Indo- 

Pacific, and then dropping anchor again in the Islands, could provide potential transport. Alter - 

natively ballast water is possible, in the form of uptake of seeds or of fragments of floating plants 

with seeds. 

Caryophyllaceae 

Spergularia marina (Linnaeus, 1758) Grisebach, 1843 Introduced 

This Eurasian-North American salt marsh plant, known as “saltmarsh sand spurry,” was first collected 

on O‘ahu in 1909 (Wagner et al., 1990). Wagner et al. (1990) note that in Hawai‘i it is found in 

“coastal and low elevation” areas, and in “dry areas” on Kure Atoll, Midway Atoll, French Frigate 

Shoals, Kaua‘i, O‘ahu, Moloka‘i, and Maui. 

Asteraceae 

Pluchea indica (Linnaeus, 1753) Lessing, 1831 Introduced 

The “Indian fleabane” or “Indian pluchea” is native to southern Asia but has been “widely introduced 

in the tropics, usually in saline places” (Wagner et al., 1990). Wagner et al. (1990) report it 

“in Hawai‘i naturalized in low elevation, dry, coastal habitats on Midway Atoll, Laysan, and probably 

all of the main islands but not documented from [the island of] Hawai‘i.” Since Wagner et al. 

(2002) report a record from the Big Island, collected in 2000 “near sea level, in degraded Scaevola 

coastal shrubland” at Keaukaha (South Hilo District). It was first collected on O‘ahu in 1915. 

Englund et al. (2000a) note that this species, along with mangroves, pickleweed, and the Pluchea 

below, dominate the shoreline areas of Pearl Harbor. 

Pluchea carolinensis (Jacquin 1789) G. Don 1839 Introduced 

= Pluchea symphytifolia (Miller, 1731) Gillis, 1977 

This pluchea, known as “sourbush” was first collected on O‘ahu in 1931, where it occurs from “relatively 

dry, coastal areas... ranging up to about 900 m in mesic to wet forest on Kure Atoll, Midway 

Atoll, French Frigate Shoals, and all of the main islands” (Wagner et al., 1990). It is native to 

Mexico, the West Indies, and northern South America, and thus forms, in part, another “Caribbean” 

element in the Hawaiian introduced biota. In Pearl Harbor it occurs, along with several other introduced 

plants as noted above, as one of the shoreline dominants in maritime conditions. 

Bataceae 

Batis maritima Linnaeus, 1758 

Introduced 

This Caribbean pickleweed was first collected in 1859 by W.F. Hillebrand “in the saltmarshes of 

Prison Island [Sand Island] near Honolulu, and has since extended to Fisherman’s Point and to

164 


Quarantine Island, where it grows with Lycium sandwicense, a plant much like it in appearance” 

(Hillebrand, 1888). Hillebrand noted that it was a “native of the West Indies (Florida, Bahamas, 

Jamaica, Venezuela)” but had also been reported from the west coast of Mexico and Baja California. 

Wagner et al. (1990) add the Galapagos Islands to this distribution, suggesting it is native there (but 

it would seem more likely to be an introduction, as it has been to the Hawaiian Islands). Wagner et 

al. (1990) reported it as “naturalized in coastal area on all of the main islands.” Hosaka (1937) noted 

that this species was one of the major plants of the “maritime zone” flora of Kipapa Gulch, which 

flows into the West Loch of Pearl Harbor. Englund et al. (2000a) note the continued abundance of 

pickleweed in Pearl Harbor. Starr (2003) stated that it “forms monotypic stands over hundreds of 

acres of Keālia Pond on Maui, and in virtually every other coastal wetland in the state of Hawai‘i.” 

Chenopodiaceae 

Atriplex semibaccata R. Brown, 1810 

Introduced 

The Australian saltbush was introduced to Lāna‘i from Australia about 1895 “as an experimental forage 

for cattle (and is) now naturalized in dry to seasonally wet areas, 0–150 m, on all of the main 

islands” (Wagner et al., 1990). It was first collected on Moloka‘i in 1910 (Wagner et al., 1990). 

Atriplex suberecta Verdoom, 1954 

Introduced 

Another saltbush species, native to Australia and South Africa (or perhaps introduced to one or the 

other?). Wagner et al. (1990) report that in Hawai‘i it occurs in “dry, disturbed areas, 0–1,920 m, on 

French Frigate Shoals, O‘ahu, Moloka‘i, Lāna‘i, Maui, and Hawai‘i.” It was first collected on O‘ahu 

in 1923. It occurs in maritime conditions as well in Hawai‘i. 

Typhaceae 

Typha latifolia Linnaeus, 1753 

Introduced 

The common cattail was first collected in O‘ahu in 1979 (Wagner et al., 1990). It is a native of 

Eurasia, northern Africa, and North America, and in Hawai‘i is “sparingly naturalized in low elevation, 

marshy sites, at least along the Wailua River, Kaua‘i, and in the Salt Lake and Pearl Harbor 

areas, O‘ahu, and perhaps also on Hawai‘i” (Wagner et al., 1990). In Pearl Harbor it occurs in the 

maritime zone. 

Poaceae 

Paspalum vaginatum Swartz, 1788 

Introduced 

The seashore paspalum, first collected on O‘ahu in 1936 and now in Hawai‘i is “naturalized in 

coastal sites in shallow brackish water and on brackish sands, often forming pure stands at sea level, 

on Kaua‘i, O‘ahu, and Hawai‘i” (Wagner et al., 1990). It is now “widely distributed in warm temperate 

to tropical sea coasts and brackish marshes worldwide” (Wagner et al., 1990). Its native range 

is not yet known. 

Rhizophoraceae 

The history of releasing no fewer than seven species of mangroves into the Hawaiian Islands has 

been reviewed by Wester (1981) and Allen (1998). Of these, three species survived, and these are 

treated below. Allen et al. (2000) noted that a fourth introduced species, Bruguiera parviflora, may 

still be on O‘ahu: “… it is possible that a small number of B. parviflora may be present in Hawai‘i, 

although the last herbarium specimens collected for this species were in 1948, and it is unlikely that 

more than a few individuals still exist.” Bruguiera parviflora was released in 1922 from the 

Philippines (Allen, 1998); any remaining trees would therefore be old and there is no evidence of 

reproduction. James Allen reports (pers. comm., January 2004) that he and K. Krauss “have explored 

virtually all the sites known to have B. sexangula and never found any other Bruguiera species …. 

if it persisted much longer than 1948 it eventually fell victim to the people who cut B. sexangula for 

access to their flowers or for poles.”


165 

Mangrove Releases: Species Not Established in Hawai‘i 

(Wester, 1981; Allen, 1998) 

Species Date Introduced from 

Bruguiera parviflora (Roxburgh) Griffiths, 1836 1922 Philippines 

Last recorded in 1948 

Rhizophora mucronata Lamarck, 1804 1922 Philippines 


Ceriops tagal C. B. Robbins 1908 1922 Philippines 


Conocarpus erectus sericeus (Grisebach) Jimenez, 1953 1946 Bahamas 

Not established from 1946 planting, but “still widely planted as an ornamental” (Allen, 1998). 

Bruguiera sexangula (Loureiro, 1790) Poiret, 1816 Introduced 

= Bruguiera gymnorrhiza of Hawaiian authors; see Allen, 1998 

The Oriental mangrove was introduced from the Philippines in 1922 by the Hawaiian Sugar Plan - 

ters’ Association (Allen, 1998, Wagner et al., 1990) in the “hopes of reclaiming” mudflats (Wester, 

1981). It is presumed to be native from tropical Africa and Madagascar to Asia, Australia, Micro - 

nesia, and Polynesia (Wagner et al., 1990). It is found in only four sites on O‘ahu, including saltwater 

marshes at He‘eia, O‘ahu (Wagner et al., 1990, Allen, 1998). Allen & Krauss (2006) examined 

the role of propagule dispersal on the restriction of this species to O‘ahu and concluded that B. 

sexangula propagules could float long enough to colonize other islands, suggesting that “factors 

other than flotation ability are responsible for the failure of B. sexangula to become established on 

other Hawaiian islands” (these factors, they speculate, might include the lack of propagules to adequately 

disperse from O‘ahu, as they “may be stranded in the vicinity of their parent trees”, as well 

as the “limited number of B. sexangula trees capable of producing propagules relative to R. mangle.” 

Rhizophora mangle Linnaeus, 1753 

Introduced 

The American or Red Mangrove was introduced from Florida in 1902 by the American Sugar Com - 

pany to stabilize mudflats and as a source of honey flora on southwestern Moloka‘i (Wester, 1981 

Wagner et al., 1990). MacCaughey (1917), Wester (1981), and Allen (1998) review the history of this 

introduction. This mangrove is native to Florida, the Caribbean (West Indies), and South America. It 

occurs in maritime environments on Kaua‘i, O‘ahu, Moloka‘i, Lāna‘i, and Hawai‘i (Wagner et al., 

1990). It is now an abundant and dominant plant along island shores (Allen, 1998, Cox & Allen, 1999, 

Allen et al. 2000). Impacts include reduction in habitat for endangered aquatic birds, elimination of 

habitat for native species, overgrowing native Hawaiian archaeological sites, and other shoreline alterations 

(Allen, 1998). Chimner et al. (2006) found that mangroves continue to expand at a rapid rate on 

O‘ahu, and “have colonized many different landforms, including tidal flats, riverbanks, fishponds, 

canals, protected reefs, embayments, lagoons, and other protected areas.” Approximately 70% (102 

hectares) of all mangroves on O‘ahu occur in Pearl Harbor (Chimner et al., 2006). 

Demopoulos et al. (2007) examined food web structure in R. mangle forests in Puerto Rico and 

Hawai‘i. 

Combretaceae 

Conocarpus erectus Linnaeus, 1753 

Introduced 

The sea mulberry, or buttonwood, is a native of southern North America and South America, ranging 

from Florida to Mexico to Brazil; it is also in Ecuador and in tropical western Africa (Wagner et 

al., 1990), which we suggest may be introductions there. Wagner et al. (1990) note that in Hawai‘i 

it is “cultivated and sparingly naturalized in coastal areas on O‘ahu, Lāna‘i, and Maui.” It was intentionally 

introduced twice, once before 1910 (possibly from Florida) and once in 1946 (from the 

Bahamas) (Wester, 1981; Allen, 1998) for ornamental purposes. It was first found on Maui in 1935 

(Wagner et al.,1990). Lorence & Flynn (1997) newly report it as a small population on Kaua‘i, in 

the Koloa District, at the Kukuiula small boat harbor, amongst littoral vegetation on a lava flow near 

sealevel, based on 1996 collections.

166 


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196 


INDEX 

(synonymys are in italics) 

abiuma, Namalycastis, 51, 55 

abnormis, Platynereis, 51, 55 

Acarina, 124 

acheilognathi, Bothriocephalus, 48, 50 

acherusicum, Corophium, 94 

acherusicum, Monocorophium, 94 

Acropora, 40 

aculeata, Crepidula, 72 

acutifrons, Caprella, 92 

adamsi, Bulla, 68, 77 

adherens, Ophryotrocha, 51, 54 

adriatica, Colurella, 48 

adriaticus, Bathydrilus, 51, 52 

aegyptiacum, Synnotum, 128, 131 

affinis, Gambusia, 142, 146 

affinis, Lyrodus, 78, 87 

aliena, Eusynstyela, 159 

alii, Vermetus, 68, 71 

aloha, Gonodactylus, 114 

altivelis, Cromileptes, 144 

alvarezii, Kappaphycus, 151, 154 

amadelpha, Avrainvillea, 151, 152, 157 

amboinensis, Namanereis, 51, 56 

amoena, Mesovelia, 116, 118 

amphitrite, Amphibalanus, 104, 105 

amphitrite, Balanus, 105 

anatifera, Lepas, 105 

andrewsi, Metafolliculina, 22, 23 

andrewsi, Paralimnoria, 23, 98, 100 

andromeda, Cassiopea, 44, 45 

anguina, Aetea, 128, 130 

angularis, Parajassa, 94 

angularis, Ruffojassa, 94 

angulatus, Canaceoides, 116, 120 

annectans, Thicanus, 120 

Annelida, 51 

annulata, Halofolliculina, 23 

Anthophyta, 162 

Anthozoa, 40 

aquayoi, Lamellodiscorbis, 22 

arabica, Cypraea, 74 

archaia, Ascidia, 135, 138 

arctica, Hiatella, 78, 85 

arcuata, Watersipora, 128, 133 

arenaceodentata, Neanthes, 51, 54 

arenaceodonta, Neanthes, 54 

arenaria, Mya, 79, 89 

areolata, Cnemidocarpa, 139 

arescus, Anoplodactylus, 124, 126 

areyi, Sertularella, 32, 38 

argus, Cephalopholis, 142, 146 

armata, Mycale, 29 

Ascidiacea, 135 

aspera, Schizophrys, 108, 114 

asperum, Echinodictyum, 25, 31 

atypha, Protula, 60 

aurantiacus, Suberites, 21, 24, 26 

aurita, Polycarpa, 135 

auritum, Conchoderma, 149 

australis, Hipponix, 68, 72 

baconi, Corophium, 95 

baconi, Laticorophium, 95 

Bacteria, 21 

bahasa/nigricauda, Enneapterygius, 144 

bartschi, Teredo, 78, 88 

beanii, Halecium, 32, 35 

benedicti, Streblospio, 51, 58 

bensoni, Teredo, 89 

bicolor, Ecsenius, 144 

bicuspidata, Laomedea, 37 

bicuspidata, Obelia, 37 

bicuspidata, Wrangelia, 151, 155 

bidens, Pistorius, 98, 100 

bidentata, Obelia, 32, 37 

bincinctus, Cricotopus, 116, 121 

bipalmulata, Bankia, 78, 87 

biseriata, Endeis, 124, 125 

bispinosa, Grandidierella, 95 

Bivalvia, 78 

blackburni, Corixa, 118 

blepharosora, Keissleriella, 51, 61 

blepharospora, Etheirophora, 22 

bori, Tectidrilus, 51, 53 

boryi, Caberea, 128, 131 

brachyacantha, Hydroides, 131 

brachyacanthus, Hydroides, 51, 61 

brakenhielmi, Symplegma, 135, 140 

brasiliensis, Ericthonius, 93 

brasiliensis, Janua, 66 

brasiliensis, Podocerus, 96 

brevicornis, Namalycastis, 52, 56 

bryani, Kleidotoma, 116, 119 

Bryozoa, 128 

caliculata, Caulibugula, 128, 131 

californica, Bugula, 131 

“californicus”, Anoplodactylus, 124 

calyptraeformis, Bostrycapulus, 68 

candidum, Didemnum, 135, 136 

canopus, Styela, 135, 139 

capitaneus, Conus, 68, 75 

carneola, Cypraea, 74 

carolinensis, Pluchea, 162, 168 

caspia, Cordylophora, 32, 33 

Cassiopea, 22 

caudata, Neanthes, 54 

cavernosa, Dictyosphaeria, 153 

cavifrons, Mugilogobius, 142, 150 

cecilia, Mycale, 24, 29 

cervicornis, Hypnea, 153 

chinensis, Fenneropenaeus, 108


197 

chinensis, Penaeus, 107, 109 

ciliata, Polydora, 57 

Ciliophora, 23 

cincta, Boonea, 68, 76 

cinerea, Ephydra, 123 

cingulata, Branchiomma, 58, 72 

Cirripedia, 104 

civile, Enallagma, 116, 117 

clandestina, Cypraea, 74 

clappi, Teredo, 78 

clathrata, Ulva, 151, 158 

claviger, Limnodriloides, 52, 53 

Cnidaria, 32 

coerulea, Halichondria, 24, 27 

coerulea, Haliclona, 24, 28 

coerulea, Sigmadocia, 28 

communis, Cephaloidophora, 24 

compressa, Anchoa, 148 

compressa, Ulva, 151, 158 

conchaphila, Ostrea, 79, 90 

conica, Sabia, 72 

connectans, Symplegma, 140 

constellatum, Polyclinum, 135, 140 

Copepoda, 90 

coquilletti, Ceropsilopa, 116, 122 

coreanica, Sphenia, 78, 84 

corelloides, Ascidia, 138 

cornicina, Dynamena, 32, 38 

coronatus, Pyrgophorus, 68, 71 

corrugata, Haliotis, 69 

corrugata, Plumularia, 36 

corrugata, Spirorbis, 66 

costata, Holmesimysis, 104 

cotti, Camallanus, 47 

couchii, Halophiloscia, 98, 102 

cracherodii, Haliotis, 69 

crassicirrata, Myrianida, 56 

crenatus, Balanus, 106 

cribraria, Cypraea, 73 

crisioides, Dynamena, 32, 38 

crocea, Tridacna, 79, 89 

cruciger, Hydroides, 61 

crucigera, Hydroides, 51, 61 

Crustacea, 90 

Ctenophora, 47 

cucullata, Ostrea, 82 

“cucullata”, Saccostrea, 78, 82 

culebrae, Littorophiloscia, 98, 102 

culebrae, Philoscia, 98, 102 

culebrae, Vandeloscia, 73 

Cumacea, 104 

cylindrical, Cypraea, 73 

cynocephala, Placopsidella, 123 

cytherea, Acropora, 22 

danilevskii, Caprella, 92 

Decapoda, 107 

decipiens, Halophila, 162 

decipiens, Nemacystus, 151, 160 

dendrograpta, Caulibugula, 128, 131 

dentata, Bugula, 128, 130 

denticulatum, Eucheuma, 151, 154 

depressa, Cypraea, 74 

derzhavini, Incisocalliope, 93 

derzhavini, Perapleustes, 93 

diacanthus, Pygoplites, 144 

dianae, Paradella, 98, 100 

diaphana, Halopteris, 32, 36 

diaphana, Sertularella, 32, 38 

dichotoma, Antipathes, 42 

dichotoma, Obelia, 32, 37 

diegensis, Teredo, 88 

diffusa, Callyspongia, 25, 28 

digitatus, Anoplodactylus, 124, 126 

Dinoflagellata, 22 

diramphus, Hydroides, 51, 61 

disjunctus, Ericthonius, 93 

distans, Amathia, 128, 129 

distans, Tridentata, 32, 39 

disticha, Halocordyle, 32 

disticha, Pennaria, 32, 34 

“dubia”, Leptochelia, 98, 99 

dura, Halichondria, 24, 27 

dura, Topsentia, 27 

dysteri, Salmacina, 62 

eburneus, Amphibalanus, 104, 106 

eburneus, Balanus, 106 

echinocarpum, Sargassum, 157 

Echinodermata, 134 

edmondsoni, Sarcothelia, 42 

edmondsoni, Watersipora, 128, 133 

elatensis, Chama, 83 

elegans, Hydroides, 51, 62 

ellipticus, Armadilloniscus, 98, 102 

elongatus, Omobranchus, 150 

engeli, Chelon, 145 

engeli, Moolgarda, 145 

engeli, Valamugil, 142, 145 

enigmatica, Mercierella, 60 

enigmaticus, Ficopomatus, 51, 60 

epihippisora, Gracilaria, 151 

equilibra, Caprella, 92 

erectus, Anoplodactylus, 124, 127 

erectus, Conocarpus, 162, 165 

eroticus, Anoplodactylus, 124, 127 

errata, Schizoporella, 132 

esakii, Oudemansia, 116, 124 

eucosmia, Iolaea, 68, 76 

eusiphona, Pellina, 27 

exasperatus, Microcosmus, 135, 141 

excavata, Gyractis, 43 

exotica, Ligia, 98, 103 

expansa, Ulva, 151, 158 

exsul, Dolichopus, 116, 121 

fagei, Mesonerilla, 51, 54 

fakaravensis, Pachygrapsus, 107, 113 

falcata, Jassa, 93

198 


falcatus, Gonodactylaceus, 107, 114 

falco, Cirrhitichthys, 144 

fasciata, Ulva, 151, 158 

fasciatus, Epinephelus, 143 

fastigiata, Lioconcha, 78, 85 

feegeensis, Hippopodina, 132 

fennica, Pedalia, 48 

ferox, Omobranchus, 142, 149 

fibrosa, Gelliodes, 24, 28 

fibula, Chama, 78, 83 

filicina, Grateloupia, 151 

fimbriatus, Cercyon, 116, 119 

Fishes, 135 

flabellata, Dictyota, 151, 161 

flavissima, Centropyge, 142, 148 

flexible, Syntormon, 116, 121 

flexuosa, Ulva, 151, 158 

floridana, Plumularia, 32, 36 

foraminosa, Neodexiospira, 51, 65 

formosanus, Centrocestus, 49 

fragile, Diplosoma, 137 

fragile, Lissoclinum, 135, 137 

franciscana, Diadumene, 40, 44 

fruticosus, Thyroscyphus, 32, 39 

fucata-martensi, Pinctada, 79, 89 

fulgens, Haliotis, 69 

fulleri, Teredo, 78, 89 

fulvus, Lutjanus, 142, 146 

Fungi, 21 

furcifera, Teredo, 78, 88 

furcillatus, Teredo, 78, 89 

gallensis, Stenothoe, 96 

galloprovincialis, Mytilus, 79 

gaspardi, Cypraea, 74 

Gastropoda, 68 

gemmata, Ascidia, 138 

gibbus, Lutjanus, 142, 147 

gigas, Crassostrea, 78, 81 

girschneri, Psilopa, 116, 124 

glomerosa, Heteropia, 25, 32 

gracilis, Barentsia, 128 

gracilis, Cypraea, 74 

gracilis, Ephydra, 116, 123 

grandimanus, Macrobrachina, 109 

grandis, Antipathes, 

grandis, Mycale, 24, 29 

granifera, Hecamede, 116, 123 

granifera, Tarebia, 68, 70 

gregoryi, Teredo, 89 

grisescens, Medetera, 116, 121 

guamensis, Hostis, 116, 124 

gurwitschi, Thalassodrilides, 51, 52 

guttatus, Cephalopholis, 146 

guttatus, Lutjanus, 143 

gymnorrhiza, Bruguiera, 165 

halecioides, Ventromma, 32, 36 

hardyi, Thinophilus, 116, 122 

hartmeyeri, Eusynstyela, 135, 139 

hastata, Lecane, 48 

havaica, Sabella, 58 

hawaiensis, Martesia, 86 

hawaiensis, Photis, 94 

hawaiensis, Saxicava, 85 

hawaiensis, Teredo, 88 

hawaiiana, Halophila, 162 

hawaiiensis, Balanus amphitrite, 105 

hawaiiensis, Bankia, 87 

hawaiiensis, Haplostomides, 91 

hawaiiensis, Namalycastis, 51, 55 

helleri, Charybdis, 108, 111 

helvola, Cypraea, 74 

hemisphaerica, Clytia, 32, 37 

hendersoni, Chama, 83 

hendersoni, Gonodactylellus, 108, 115 

hepatus, Paracanthurus, 144 

herbstii, Panopeus, 111 

hermaphroditus, Gyratrix, 49 

hexagonatus, Epinephelus, 143 

hiloensis, Teredo, 88 

hippocrepia, Phoronis, 127 

hirundo, Cypraea, 74 

holoprasinus, Goeldichironomus, 116, 121 

honoluluensis, Teredo, 88 

horii, Moerisia, 32, 34 

horii, Ostroumovia, 34 

horvathi, Caecijaera, 98, 101 

humilis, Garveia, 34 

humilis, Micracanthia, 116, 118 

hupferi, Tridentata, 32, 39 

hyotis, Hyolissa, 82 

ibari, Brachydeutera, 116, 122 

ignis, Tedania, 30 

immigrans, Atergatopsis, 108, 113 

imperfecta, Ecteinascidia, 135, 138 

inclusus, Amphithalamus, 68, 71, 76 

indica, Caloria, 68, 77 

indica, Chaetomorpha, 151, 160 

indica, Hinemoa, 68, 76 

indica, Learchis, 77 

indica, Pluchea, 162, 163 

indica, Sabellastarte, 59 

indicus, Fenneropenaeus, 108 

inflata, Trochammina, 22 

Insecta, 116 

insidiosum, Corophium, 95 

insidiosum, Monocorophium, 95 

insolita, Herdmania, 141 

insulae, Tropichelura, 97 

intermedia, Armandia, 51, 58 

intestinalis, Ciona, 135, 137 

intestinalis, Hyattella, 25, 31 

intestinalis, Ulva, 159 

irene, Cnemidocarpa, 135, 139 

irregularis, Anthipathes, 94 

irroratus, Epinephelus, 143 

Isopoda, 98


199 

istiblenni, Spirocamallanus, 47 

jacobsoni, Atrichopogon, 116, 121 

japonica, Branchiomma, 51, 58 

japonica, Corella, 137 

japonica, Grandidierella, 95 

japonica, Tapes, 74 

japonicus, Marsupenaeus, 108 

japonicus, Penaeus, 109 

japonicus, Spinther, 51, 54 

japonicus, Telmatogeton, 116, 121 

Kamptozoa, 128 

kasmira, Lutjanus, 48, 142, 149 

kauaiensis, Teredo, 88 

knightjonesi, Janua, 65 

knightjonesi, Leodora, 51, 65 

koehleri, Pileolaria, 67 

koehleri, Vinearia, 51, 67 

konaensis, Bankia, 87 

kraussii, Pomatoleios, 51, 62 

kuroharai, Cypraea, 68, 74 

labiata, Aurelia, 44 

labrolineata, Cypraea, 74 

lacerata, Tramea, 116, 117 

lacustris, Cordylophora, 33 

lacustris, Panopeus, 107, 111 

lafontii, Catenaria, 132 

lafontii, Savignyella, 128, 132 

lamellatus, Porcellio, 98, 103 

lar, Macrobrachium, 107, 109 

latifolia, Typha, 162, 163 

latipinna, Mollienesia, 145 

latipinna, Poecilia, 142, 145 

latitheca, Clytia, 32, 37 

lazarus, Chama, 78, 83 

leucolena, Diadumene, 40, 43 

leucosternon, Acanthurus, 144 

lignicola, Eufolliculina, 24 

lignorum, Limnoria, 24, 101 

ligulata, Sertularia, 39 

limnoriae, Mirofolliculina, 24 

lineata, Diadumene, 40, 43 

lingulata, Ulva, 158 

linza, Ulva, 151, 159 

listerianum, Diplosoma, 135, 136 

litoralis, Armadilloniscus, 102 

littoralis, Namanereis, 51, 56 

littoralis, Pilayella, 151, 161 

loculosa, Tridentata, 32, 39 

loriculus, Centropyge, 148 

luciae, Haliplanella, 44 

lugubris, Myzobdella, 52, 53 

lunifer, Eupomatus, 61 

lunulifera, Hydroides, 53, 61 

macdonaldi, Diplosoma, 136 

macerophylla, Chama, 78, 83 

macrocephala, Tilapia, 148 

macrodactyla, Tedania, 148 

maenas, Carcinus, 108, 110 

makena, Neomicrodeutopes, 95 

mangle, Rhizophora, 21, 162, 165 

manilaensis, Haplophragmoides, 22 

marginata, Tridentata, 32, 39 

marina, Labyrinthomyxa, 23 

marina, Spergularia, 162 

marinum, Dermocystidium, 23 

marinus, Perkinsus, 23 

marinus, Pseudodiaptomus, 90 

marioni, Spirorbis, 51, 67 

maritima, Anisolabis, 116, 117 

maritima, Batis, 162, 163 

marmorea, Bulla, 77 

marquesana, Placopsidella, 116, 123 

marquesensis, Sardinella, 142, 145 

marshallensis, Anoplodactylus, 124, 127 

mauiensis, Melania, 70 

maunakea, Mycale, 29 

maunakea, Mycale, 29 

mauritiana, Herdmania, 135, 141 

megathecum, Synthecium, 32, 37 

melanodocia, Halichondria, 24, 27 

melanotheron, Sarotherodon, 142, 148 

melanotheron, Tilapia, 148 

melleni, Neobenedenia, 48, 50 

mera, Discocerina, 116, 122 

mercenaria, Mercenaria, 79, 85 

meretrix, Meretrix, 79, 89 

merra, Epinephelus, 143 

mesoleucus, Chaetodontoplus, 144 

microadriatacum, Symbiodinium, 22 

micronesiae, Leucothoe, 96 

midwayensis, Teredo, 88 

milbrae, Ephydra, 116, 123 

miliaris, Vitularia, 68, 74 

militaris, Caloria, 77 

militaris, Pileolaria, 51, 53, 66 

milleri, Plumularia, 36 

milleri, Teredo, 87 

miniaceus, Microcosmus, 141 

minima, Bugula, 128, 131 

minor, Hipponix, 77 

minusculus, Smithsonidrilus, 51, 53 

minuta, Corella, 135, 137 

minutum, Nanosesarma, 107, 113 

minutus, Pomataoceros, 60 

mirablis, Syncoryne, 33 

mirafloresensis, Acantholobulus, 111 

modesta, Ophiactis, 134 

molestus, Pectinodrilus, 51, 52 

Mollusca, 68 

momus, Herdmania, 135, 141 

monodon, Penaeus, 21, 108 

mossambica, Tilapia, 148 

mossambicus, Oreochromis, 142, 148 

mucronata, Rhizophora, 165 

mulsanti, Mesovelia, 116, 118 

multiformis, Vallicula, 47

200 


mundulus, Cyclodinus, 116, 120 

murphyi, Parathroscinus, 116, 119 

musciformis, Hypnea, 151, 152, 156 

muscus, Bougainvillia, 32, 34 

mutatus, Gonodactylaceus, 114 

muticum, Sargassum, 151, 161 

Myscidacea, 104 

navisa, Ividella, 76 

Nematoda, 47 

neritina, Bugula, 128, 130 

nidifica, Laurencia, 159 

nigra, Phallusia, 135, 137 

nigromaculata, Branchiomma, 58 

nigronotatus, Donaceus, 116, 122 

niloticus, Trochus, 68, 69 

nipponica, Janua, 66 

nipponica, Neodexiospira, 51, 66 

nobilis, Anomia, 78, 79 

nodosa, Endeis, 124, 125 

nordgardi, Metafolliculina, 23 

norvegica, Hydroides, 53, 62 

nuchalis, Polydora, 51, 57 

nummiforme, Discosoma, 40 

nummiformis, Actinodoscus, 40 

nutricula, Turritopsis, 32, 33 

nuttalli, Clinocardium, 89 

oahuensis, Alloniscus, 98, 102 

oahuensis, Pilumnus, 107, 112 

obelisus, Trochus, 70 

obliquus, Omobranchus rotundiceps, 142, 150 

oceania, Symplegma, 140 

oceanicus, Metopograpsus, 107, 113 

okupi, Tapes philippinarum, 84 

Oligochaeta, 52 

oodes, Pyrgulina, 68, 76 

orientalis, Eulaeospira, 51, 65 

ornata, Tethya, 24, 26 

oxycephalus, Cirritichthys, 44 

oxyuris, Hexarthra, 48 

pachycera, Myrianida, 51, 56 

pacifica, Ammothella, 124, 125 

pacifica, Chama, 78, 83 

pacificus, Acantholobulus, 107, 111 

pacificus, Cotylorhizoides, 45 

pacificus, Panopeus, 111 

pagenstecheri, Janua, 51, 65 

pallasiana, Cryptosula, 128, 132 

pallida, Cypraea, 74 

pallida, Herdmania, 135, 141 

paradoxa, Ulva, 151, 158 

parasitica, Anthohebella, 37 

parishii, Mycale, 24, 30 

parksi, Teredo, 89 

partita, Styela, 139 

parviflora, Bruguiera, 164 

parvus, Mugilogobius, 150 

peasiana, Bulla, 77 

pectenicrus, Elasmopus, 96 

pedicellatus, Lyrodus, 78, 88 

pedispinis, Parapseudes, 98, 99 

pedroana, Peristichia, 68, 76 

pellucida, Okenia, 68, 78 

pellucidum, Zoobotryon, 78, 129 

penantis, Caprella, 92 

penicillatus, Hemigrapsus, 108, 113 

peninsularis, Megabalanus, 105 

pentodon, Sphaeroma, 100 

perca, Cuthona, 68, 77 

perlucidum, Didemnum, 135, 136 

philippinarum, Paphia, 84 

philippinarum, Tapes, 78 

philippinarum, Venerupis, 78 

Phoronida, 127 

phyteuma, Lytocarpia, 32, 36 

platensis, Orchestia, 98 

platensis, Platorchestia, 98 

Platyhelminthes, 48 

plicatilis, Brachionus, 48 

poindimieri, Phyllodesmium, 68, 78 

pollinosa, Paratissa, 116, 123 

Polychaeta, 53 

polymorpha, Halopteris, 32, 36 

polyphyllum, Sargassum, 157 

poraria, Cypraea, 74 

portus, Anoplodactylus, 126 

prima, Clotenopsa, 125 

proboscidea, Boccardia, 52, 59 

procera, Clathria, 25, 29 

procera, Endeis, 124, 126 

projectus, Anoplodactylus, 126 

prolifera, Ulva, 151, 158 

proteus, Chthamalus, 104, 106 

Protoctista, 22 

psammathodes, Didemnum, 135, 136 

pseudoclavus, Pileolaria, 51, 66 

pseudocorrugata, Janua, 66 

pseudocorrugata, Neodexiospira, 51, 66 

pseudomilitaris, Pileolaria, 67 

pseudomilitaris, Simplicaria, 51, 67 

pugettensis, Upogebia, 108 

punctata, Phyllorhiza, 44, 45 

pupu, Odostomia, 76 

pusilla, Paracaprella, 93 

Pycnogonida, 124 

pycnosoma, Anoplodactylus, 124, 129 

pyrifera, Macrocystis, 151, 160 

quadridentata, Dynamena, 32, 38 

quadrimaculatus, Herklotsichthys, 142, 145 

quoianum, Sphaeroma, 99, 100 

quoyanum, Sphaeroma, 100 

rachelfitzhardingeae, Culicia, 40, 41 

radiatum, Cladonema, 32, 34 

ramburii, Ischnura, 116, 117 

ramosa, Bougainvillia, 34 

“rapax”, Elasmopus, 96 

rayi, Gnorimosphaeroma, 98, 99


201 

recta, Aetea, 130 

rectisetosus, Pectinodrilus, 51, 52 

rehderi, Tanystylum, 124, 125 

reptans, Symplegma, 135, 140 

reticulata, Tedania, 25, 30 

reticulata, Trichocorixa, 116, 118 

reticulata, Ulva, 151, 158 

reticulatus, Amphibalanus, 104, 106 

reticulatus, Balanus, 106 

rhizophorae, Cytospora, 21 

rigida, Ulva, 151, 159 

riisei, Carijoa, 40, 42 

riisei, Telesto, 42 

robusta, Bugula, 131 

rosenbergii, Macrobrachium, 108, 109 

Rotifera, 48 

rubicundus, Limnodriloides, 51, 53 

rupestris, Kuhlia, 143 

ruppelli, Diodora, 68, 69 

sagamiensis, Polyandrocarpa, 135, 140 

salicornia, Gracilaria, 151, 152, 155 

salicornia, Psychoda, 116, 121 

salsamentus, Tropisternus, 116, 119 

salvatoris x mexicana group, Poecilia, 142, 143 

sanctijosephi, Sabellastarte, 59 

sandwicensis, Eleotris, 47 

sandwicensis, Vitularia, 74 

sanguinea, Eulalia, 56 

sanguinea, Eumida, 51, 56 

sapidus, Callinectes, 108, 110 

savignyi, Ophiactis, 134 

saxatilis, Morone, 143 

sayi, Enochrus, 116, 119 

scaura, Caprella, 92 

Scleractinia, 40 

sculpta, Paracerceis, 98, 100 

Scyphozoa, 44 

sebae, Lutjanus, 144 

seculus, Salsuginus, 48, 50 

secundaria, Antennella, 32, 36 

semibaccata, Atriplex, 162, 164 

semicirculatus, Pomacanthus, 144 

semidecussata, Tapes, 84 

semilutea, Paratissa, 123 

semimilitaris, Pileolaria, 60 

seminudus, Glabropilumnus, 107, 112 

senegalensis, Namalycastis, 52, 56 

sericeus, Conocarpus erectus, 165 

serrata, Scylla, 107, 110 

servilia, Crocothemis, 116, 117 

sesere, Actinogeton, 43 

setacea, Plumularia, 32, 36 

sexangula, Bruguiera, 162, 165 

sexradia, Ophiactis, 134 

shawi, Anomalorhiza, 44, 46 

sibogae, Halecium, 32, 35 

simodensis, Botrylloides, 135, 139 

simplex, Ascobius, 23 

sinensis, Eriocheir, 108, 114 

sitiens, Pellina, 27 

sivickisi, Carybdea, 44, 46 

sp., Abra, 78 

sp., Amphiprion, 144 

sp., Apseudes, 98, 99 

sp., Atrichopogon, 116, 121 

sp., Aurelia, 44 

sp., Biemna, 24, 29 

sp., Botrylloides, 135, 139 

sp., Botryllus, 135, 139 

sp., Buchnerillo, 98, 103 

sp., Capitella, 51, 59 

sp., Cassiopea, 22, 44, 45 

sp., Chaetopterus, 58 

sp., chalinid, 28 

sp., Cliona, 24, 26 

sp., Convolutriloba, 49 

sp., Dendronephthya, 41 

sp., Dysidea, 25, 31 

sp., Eudendrium, 34 

sp., Exosphaeroma, 98, 100 

sp., Favonigobius, 142, 150 

sp., Garveia, 34 

sp., Halichondria, 24, 27 

sp., Haliotis, 68 

sp., Isognomon, 66 

sp., Lethrinus, 143 

sp., Malacoceros, 57, 58 

sp., Mesanthura, 98, 100 

sp., Minuspio, 51, 58 

sp., Monanchora, 24, 30 

sp., Nannastacus, 104 

sp., Oceanapia, 24, 27 

sp., Paraleucothoe, 95 

sp., Pione, 24, 26 

sp., Sarcothelia, 40, 42 

sp., Scherocumella, 104 

sp., Symplegma, 135, 140 

sp., Tedania, 31 

sp., Thylaeodus, 68, 71 

sp., Topsentia, 24 

sp., Toxiclona, 24, 28 

sp., Zoothamnium, 23 

sp. cf. avara, Dysidea, 31 

sp. cf. americana, Evalea, 76 

sp. cf. arenaria, Dysidea, 31 

sp. cf. gracilis, Bowerbankia, 128, 129 

sp. cf. imbricata, Bowerbankia, 128, 129 

sp. cf. murphyi, Parathroscinus, 119 

sp. cf. tenella, Culicia, 41 

sp. cf. wulferti, Proalides, 48 

species A, Ascidia, 136, 138 

species B, Ascidia, 135, 138 

speciosa, Sertularella, 30 

spectabilis, Sabellastarte, 51, 59 

sphaerocephala, Lumbrineris, 51, 54 

spicifera, Acanthophora, 151, 152, 153

202 


spinosum, Crucibulum, 68,73 

spinosus, Hyastenus, 107, 114 

Spirorbinae, 63 

splendidum, Corydendrium, 34 

squamosa, Tridacna, 79, 89 

stagnalis, Scatella, 116, 124 

staminea, Protothaca, 89 

staphylaea, Cypraea, 73, 74 

steueri, Janua, 65 

stolonifera, Bugula, 128, 131 

Stomatopoda, 107, 114 

stri, Psammopsyllus, 90 

striata, Martesia, 78, 86 

striatum, Eucheuma, 154 

striatum, Kappaphycus, 151, 152, 154 

strictocarpa, Plumularia, 32, 36 

stultorum, Tivela, 78, 89 

stylirostris, Litopenaeus, 109 

stylirostris, Penaeus, 109 

suberecta, Atriplex, 162 

subovoidea, Watersipora, 133 

subtilis, Sertularia, 32, 39 

subtilis, Sertularia, 39 

subtorquata, Watersipora, 128, 133 

succinea, Neanthes, 51, 55 

sydneiensis, Ascidia, 135, 138 

symphytifolia, Pluchea, 163 

taeniata, Ulva, 151, 159 

tagal, Ceriops, 165 

tahitiensis, Hippopodina, 128, 132 

Tanaidacea, 98 

?tapeinosoma, Pseudochromua, 144 

tarnis, Thetella, 116, 118 

taupou, Chrysiptera, 142, 149 

“taxiformis”, Asparagopsis, 151, 155 

tenuicollis, Ascocotyle, 48, 50 

terebrans, Sphaeroma, 99 

teredini, Taenioplana, 48, 49 

Teredinidae, 86 

thysanius, Parablennius, 142, 149 

tiarella, Pennaria, 34 

tibialis, Mosillus, 116, 123 

tiera, Platax, 144 

tikvahiae, Gracilaria, 151, 156 

timsanus, Pigrogromitus, 124, 125 

tintinnabulum peninsularis, Balanus, 105 

tongensis, Sertularella, 32, 38 

torreyi, Sertularella, 38 

trachis, Chrysallida, 68, 76 

Trematoda, 50 

tribranchiata, Salmacina, 51, 62 

tripunctata, Limnoria, 98, 100 

trulliformis, Teredo, 88 

truncata, Aetea, 128, 130 

truncatus, Olibrinus, 98, 103 

tuberculata, Melanoides, 68, 70 

tuberculata, Pustulostrea, 79, 90 

tubulosa, Sarsia, 33 

tubulosa, Ulva, 158 

tulipa, Eualetes, 72 

turbinata, Tridentata, 32, 39 

turgescens, Montipora, 41 

typica, Teredicola, 90 

uberrima, Vittaticella, 132 

uncinata, Clasiopella, 116, 122 

unguiculata, Neofolitispa, 30 

unicornia, Schizoporella, 132 

urodeta, Cephalopholis, 142 

vaginatum, Paspalum, 162, 164 

vaigiensis, Abudefduf, 142, 149 

valida, Stenothoe, 87 

valida, Stenothoe, 91, 96, 99 

vanicolensis, Mulloidichthys, 147 

vannamei, Litopenaeus, 108, 109 

vannamei, Penaeus, 109 

variopedatus, Chaetopterus, 58 

variseta, Tethina, 120 

vasculosum, Polyclinum, 140 

vastifica, Cliona, 26 

ventosa, Eurystheus, 94 

ventosa, Ruffojassa, 94 

ventosa, Ventojassa, 94 

venusta, Trypostega, 128, 133 

vermicularis, Serpula, 63 

vernicosa, Bulla, 68, 77 

verticillatum, Zoobotryon, 128, 129 

violaceae, Parafolliculina, 23 

virginica, Crassostrea, 23, 78, 80 

viridis, Lagotia, 23 

Viruses, 21 

vittata, Harengula, 145 

vittata, Limia, 142, 146 

vittata, Poecilia, 145 

vittatus, Upeneus, 142 

volans, Mazzaella, 151, 157 

vulgare, Armadillidium, 103 

walkeri, Sphaeroma, 98, 100 

watsoni, Serpula, 51, 57, 63 

websteri, Polydora, 51 

wilberti, Haplophragmoides, 22 

williamsi, Procanace, 116, 120 

willistoni, Tethina, 116, 120 

xanthopunctatus, Apolemichthys, 22, 144 

zebra, Istiblennius, 47 

zeteki, Suberites, 26 

zeteki, Terpios, 26 

zooritensis, Polyandrocarpa, 135, 140

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