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How many species are there in Hawaii?1
LUCIUS G. ELDREDGE2 and SCOTT E. MILLER (Department of Natural Sciences, Bishop
Museum, P.O. Box 19000A, Honolulu, Hawaii 96817, USA)
Introduction
The Hawaiian Islands are the most isolated island group in the world. The 8 main
southeastern islands, with their sequentially younger geological ages, great physiograph-
ic and climactic variation, are ideal natural laboratories for evolutionary and ecological
research (see Howarth 1990, Kay 1994 for further background). Although the Hawaiian
biota has been subjected to systematic study since shortly after the founding of Linnaean
nomenclature, these results are scattered through publications and collections around the
world. Moreover, many gaps remain in the knowledge of Hawaii?s biota, especially for the
smaller organisms such as viruses, bacteria, freshwater algae, protists, and soil inverte-
brates other than insects and mollusks. This paper is a first approximation by Bishop
Museum?s Hawaii Biological Survey to answer the frequently asked question: ?How
many species are there in Hawaii??
In compiling this list, we have collected and assembled data and wisdom of the staff
of Bishop Museum, libraries of the Bishop Museum and the University of Hawaii, and
many colleagues in Hawaii and around the world. The marine portions draw heavily from
the ongoing project Reef and shore fauna of Hawaii. The terrestrial invertebrate data
derive primarily from an invertebrate survey pilot project supported by the John D. and
Catherine T. MacArthur Foundation in the late 1980s. An early version of the terrestrial
and freshwater portions of the list was circulated at the Hawaii Conservation Biology
Initiative annual meeting in 1992. The interest of that group prompted us to continue the
effort. We know that the present list is incomplete in several ways: 1) we expect that some
literature has been missed; 2) we know that additional information, including unpublished
new species, exist in museum collections; and 3) further fieldwork will produce addition-
al records. The Hawaii Biological Survey is creating comprehensive databases for basic
taxonomic, distributional, and biological information on all the organisms occurring in
Hawaii, so we welcome additions and corrections to this list.
Classification of animals generally follows Parker (1982), with the use of kingdoms
in the sense of Margulis & Schwartz (1988). Some artificial categories of convenience are
used (for examples, marine mollusks). Freshwater categories generally include brackish
water dwellers. Viruses and bacteria are not included because too few data are available,
although the plant parasitic species were reviewed by Raabe et al. (1981). Human para-
sites are also not included. Unless noted, all numbers represent described species report-
ed in the literature and often under-represent the eventual total which can be expected
based on further exploration. On the other hand, there may be many more names in the
literature for some groups because the taxa have not been reviewed or revised to eliminate
the synonyms. The numbers from the Reef and shore fauna manuscripts include previ-
ously unpublished records.
The term endemic is problematic. In the terrestrial environment it is easy to deter-
mine the boundaries of the Hawaiian Islands, but the fact that a species is known only
from Hawaii may mean only that it has not yet been reported elsewhere. This is especial-
3
1.  Contribution No. 1995?002 to the Hawaii Biological Survey.
2.  Executive Secretary, Pacific Science Association, P.O. Box 17801, Honolulu, Hawaii 96817, USA
ly a problem in groups that require specialized sampling techniques and that may have
been better studied in Hawaii than in many other places (e.g., planktonic crustaceans, soil
mites). In the marine environment this problem is worse because the ocean provides a
medium for movement throughout the Indo-west Pacific. Moreover, it is difficult to deter-
mine boundaries. Thus, we expect that most of the taxa currently listed as endemic to
Hawaii?s marine environment will ultimately prove to be more widely distributed.
Definitions
For the marine environment ?Hawaii? includes the coastal areas to the deepest
waters, including all the waters surrounding all of the islands of the archipelago.
Nonindigenous species (NIS) are those species that do not naturally occur in the
Hawaiian Islands and have arrived either accidentally or intentionally through biological
control efforts, aquaculture imports, etc. These species have also been referred to as alien,
exotic, adventive, or introduced species (Frank & McCoy 1990).
Indigenous species are those species which naturally occur in the islands but are not
endemic to the islands (i.e., they also occur naturally elsewhere).
For birds, visitors are nonresident species; migratory birds follow a repeatable pat-
tern of occurrence in the islands.
Subfossil/fossil species in Hawaii are used in the context of birds and bats that
became extinct prior to the arrival of Captain Cook. Invertebrate fossils discovered
through core sampling, archaeological digs, and other techniques have not been included
in the data tabulated below.
Extinct species are those forms that no longer exist in their natural habitat. It is dif-
ficult at best to attempt a tabulation of how many of the species originally described from
the Hawaiian Islands have become extinct through natural processes or with the advent of
humans and the concomitant destruction of native habitats and associated species through
time. Data on known extinct vertebrates are included in the notes, but not in Table 1.
Table 1. Estimates of Numbers of Species of the Hawaiian Biota
Taxon Total Endemic NIS Reference(s)
PROTOCTISTA
Algae
freshwater 120+ 2 ? Vis et al. 1994; I.A. Abbott, pers. comm.
marine 470 ? 5 I.A. Abbott, pers. comm.
Myxomycota 101 0 ? Eliasson, 1991
Protozoa
Foraminifera 1000+ ? 0 Phillips, 1977; Chave, 1987
Other protozoa 42 2? 0 Ball, 1963; Phillips, 1977; Larsen & 
Patterson, 1990; Boyko, 1994
FUNGI
Fungi 1300 ? ? Baker & Goos, 1972; Goos, 1977; 
Kohlmeyer, 1969
Lichens 723 240 ? Smith, 1991, 1993
PLANTAE
Bryophytes
mosses 244 112 11 Hoe, 1979
liverworts 220 ? ? Miller & Whittier, 1990
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Taxon Total Endemic NIS Reference(s)
Fern Allies 24 7 2 W.H. & F. Wagner, pers. comm.
Ferns 205 124 27 W.H. & F. Wagner, pers. comm.
Gymnosperms 10? 0 10? W. Appleby, pers. comm.
Angiosperms 1894 850 861 Wagner et al., 1990; HBS Records for
1994*
ANIMALIA
Phagocytellozoa
Placozoa 1 0 0 Pearse, 1989
Parazoa
Porifera 
marine 84 24 4+ Bergquist, 1977; Chave & Jones, 1991
freshwater 1 0 1 Svihla, 1941
Eumetazoa
Cnidaria
Hydroida 
marine 23 4 1 Cooke, 1977
deepwater 51 31 0 Nutting, 1905
freshwater 2 0 2 Matthews, 1966
Siphonophora 1 0 0 Eldredge & Devaney, 1977
Chondrophora 2 0 0 Eldredge & Devaney, 1977
Scyphozoa 13 0 5 Devaney & Eldredge, 1977; Cooke, 
1984; Burch & Burch, 1995
Octocorallia 105 17 1 Devaney, 1977b; Grigg & Bayer, 
1976; Muzik, 1978; Versevelt & 
Bayer, 1988; Bayer, 1990; Chave & 
Jones, 1991
Zoantharia 23 5 1 Cutress, 1977
Zoanthiniara 7 4 0 Walsh & Bowers, 1977
Scleractinia 99 12 0 Maragos, 1977, 1995; Grigg et al., 1981;  
Cairns, 1984
Antipatharia 15 1 0 Grigg & Opresko, 1977; Chave & Jones, 
1991
Ctenophora 10 0 1 Devaney, 1977a; R. Galt, pers. comm
Platyhelminthes
marine 41 20 0 Poulter, 1987
freshwater/terrestrial 8 4 4 Hyman, 1939; Kawakatsu & Mitchell, 
1984; Kawakatsu et al., 1984 
parasitic 519 371+ 0 Yamaguti, 1968a, 1968b, 1970; Kaneko et
al., 1988; Dailey et al., 1992
Nemertinea
marine 26 5? 0 Coe, 1947; Devaney & Eldredge,
1987; J. Norenburg, pers. comm.
freshwater/terrestrial 2 0 2 Howarth & Moore, 1983;
J. Norenburg, pers. comm.
Gnathostomulida 8 4 0 Sterrer, 1991
Rotifera
marine 3 0 0 Hope, 1987
freshwater 27 ? 0 P.N. Turner, pers. comm
Nematoda
marine 54 23 0 Hope, 1987
plant/soil 127 ? ? Olveira, 1940; E.M. Noffsinger & E.P. 
Caswell (unpubl. 1989)
zooparasitic 33+ ? ? Chapin, 1925; Deardorff et al., 1982; 
Deardorff, 1987; McKenzie & 
Davidson, 1989
RECORDS OF THE HAWAII BIOLOGICAL SURVEY FOR 1994 5
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Taxon Total Endemic NIS Reference(s)
Priapulida 1 0 0 J. Bailey-Brock, pers. comm..
Mollusca
Gastropoda
marine 572 119 4 Kay, 1979; Kay & Palumbi, 1987
terrestrial 831? 759 50? R.H. Cowie et al., in press 
freshwater 37? 7 30? R.H. Cowie, pers. comm.
Bivalvia
marine 129 66 1 Kay, 1979; Kay & Palumbi, 1987
freshwater 1 0 1 Burch, 1978; HBS Records for 1994
Polyplacophora 4 3 0 Kay, 1979
Scaphopoda 3 0 0 Kay, 1979
Cephalopoda 73 2 0 Kay, 1979; Berry, 1914; Roper & 
Young, 1975; Roper et al., 1984; 
Young, 1991
Aplacophora 6 0 0 Kay, 1979
Annelida
Oligochaeta
marine 26 11 3? Bailey-Brock, 1987; Ers?us & Davis, 
1989; Ers?us, 1990
terrestrial 20 0 20 Nakamura, 1992; S.W. James, unpubl.
Archiannelida 2 0 0 Bailey-Brock, 1987
Hirudinea
marine 3 0 0 Bailey-Brock, 1987; Choy et al., 1989
terrestrial 3 0 3 Moore, 1946
Polychaeta
marine 250+ 69+ 4 Pettibone, 1986; Paxton & Bailey-
Brock, 1986; Bailey-Brock & 
Hartman, 1987; Bailey-Brock, 1990, 
1991; Fauchald, 1992
freshwater 1 0 0 Van Zwaluwenburg, 1948
Pogonophora 2 2 0 Southward, 1980
Echiura 6 0 0 Edmonds, 1987 
Sipuncula 14 0 0 Edmonds, 1987
Arthropoda
Insecta (nonmarine)
Blattaria 19 0 19 Nishida, 1994
Coleoptera 1987 1358 597 Nishida, 1994; HBS Records for 1994
Collembola 168 95 71 Nishida, 1994
Dermaptera 24 9 14 Nishida, 1994
Diptera 1426 1048 631 Nishida, 1994
Heteroptera 316 214 96 Nishida, 1994; HBS Records for 1994
Homoptera 689 385 299 Nishida, 1994; HBS Records for 1994
Hymenoptera 1283 654 605 Nishida, 1994; HBS Records for 1994
Lepidoptera 1153 950 192 Nishida, 1994; HBS Records for 1994
Mallophaga 102 5 44 Nishida, 1994
Neuroptera 60 51 8 Nishida, 1994
Odonata 41 31 8 Nishida, 1994; HBS Records for 1994
Orthoptera 290 263 26 Nishida, 1994; Otte, 1994
Psocoptera 135 89 42 Nishida, 1994
Thysanoptera 147 29 116 Nishida, 1994
Remaining orders 62 6 45 Nishida, 1994
Insecta (marine) 4 1 0 Herring, 1961, 1965; N.L. Evenhuis, 
pers. comm.
Araneae 198 119 71 Nishida, 1994; HBS Records for 1994
Chilopoda 24 12 12 Nishida, 1994
Diplopoda 26 16 9 Nishida, 1994
Palpigrada 1 0 1 Nishida, 1994
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Taxon Total Endemic NIS Reference(s)
Pauropoda 2 0 1 Nishida, 1994
Pseudoscorpionida 12 12 0 Nishida, 1994
Shizomida 1 0 1 Nishida, 1994
Scorpionida 1 0 1 Nishida, 1994
Crustacea 
Branchiopoda 4 0 0 Ueno, 1936
Ostracoda
marine 31 ? 0 J.C. Holden, MS; Kornicker, 1976; 
Danielopol & Hartmann, 1986; 
Hartmann, 1991
freshwater 1 ? ? V?vra, 1906
Copepoda 100 ? ? L.G. Eldredge, MS; Motoda, 1963; 
Lewis, 1966, 1967; Humes, 1976; 
Wells, 1986; Kunz, 1993; HBS 
Records for 1994
Cirripedia 58 ? 0 W.A. Newman, MS; Pilsbry, 1907; 
Boschma, 1953
Stomatopoda 17 11 1 L.G. Eldredge, MS
Mysidacea 14 ? 0 L.G. Eldredge, MS; Ortmann, 1906
Tanaidacea 8 7 0 M. Miller, MS
Isopoda
marine 27 ? ? M. Miller, MS; Richardson, 1906
terrestrial 51 15 26 Nishida, 1994
Amphipoda
marine 172 ? 8 Barnard, 1970; Brusca, 1973, 1978
terrestrial 9 6 3 Nishida, 1994
Euphausiacea 27 ? 0 Ortmann, 1906; Brinton, 1962
Decapoda
Penaeidae 20 0 0 L.G. Eldredge, MS
Sergestidae 10 0 0 L.G. Eldredge, MS
Stenopodidea 5 4 1 L.G. Eldredge, MS
Caridea 149 ? 1 L.G. Eldredge, MS
Astacidea 1 0 1 Eldredge, 1994
Thalassinidea 8 ? 1 L.G. Eldredge, MS
Palinuridea 13 2 0 L.G. Eldredge, MS
Anomura 43 ? 0 L.G. Eldredge, MS
Brachyura 189 ? 7 L.G. Eldredge, MS; HBS Records for 1994
Acari
marine 19 17 2 Nishida, 1994
terrestrial 517 97 337 Nishida, 1994; HBS Records for 1994
Pycnogonida 12 ? 0 J.H. Stock, MS; Child, 1972
Tardigrada 22 1 19 Nishida, 1994; McInnes, 1994
Phoronida 3 0 0 Emig & Bailey-Brock, 1987
Bryozoa 150+ ? ? Soule et al., 1987
Entoprocta ? ? ? Soule et al., 1987
Brachiopoda 3 0 0 Emig, 1987; B.L. Burch, pers. comm.
Chaetognatha 8 1 0 Alvari?o, 1978; Pierrot-Bults & Nair, 1991
Echinodermata
Crinoida 16 15 0 L.G. Eldredge, MS; Clark, 1949
Asteroidea 82 53 0 L.G. Eldredge, MS; Clark, 1949
Echinoidea 75 35 0 L.G. Eldredge, MS; Clark, 1949
Holothuroidea 48+ 19 0 L.G. Eldredge, MS; Clark, 1949
Ophiuroidea 57 28 0 D.M. Devaney & A.N. Baker, MS; Clark, 
1949
Hemichordata 4 1 0 L.G. Eldredge, MS; Hadfield & Young, 
1983; 1 undescribed sp. 
Chordata
Cephalochordata 1 0 0 Eldredge, 1967
RECORDS OF THE HAWAII BIOLOGICAL SURVEY FOR 1994 7
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Taxon Total Endemic NIS Reference(s)
Urochordata (Tunicata)
Thaliacea 24 0 0 Metcalf & Hopkins, 1919; Yount, 1954; 
Soest, 1974a, 1974b
Larvacea 2 0 0 Taguchi, 1982; Kitalong, 1986
Ascidiacea 45 7? 2? D.P. Abbott, MS; Tokioka, 1967
Pisces
marine 1150+ 134+ 33 Eldredge, 1994; B. Mundy, pers. comm.
freshwater 45 5 40 Eldredge, 1994; Maciolek, 1984, Devick 
et al., 1992
Amphibia 4 0 4 Vitousek et al., 1987; C. Kishinami, pers. 
comm.
Reptilia 18 0 18 Vitousek et al., 1987; Eldredge, 1994; C. 
Kishinami, pers. comm.
Aves 274 60 46 R.L. Pyle, pers. comm. 
Mammalia
terrestrial 20 1 19 Tomich, 1986
marine 24 0 0 Tomich, 1986
totals 21,383 8,759 4,532
*  HBS Records for 1994 = articles in Bishop Museum Occasional Papers 41 and 42 of the Records of the
Hawaii Biological Survey for 1994.
Notes
Algae: Marine algae numbers are from an identification manual in preparation by I.A. Abbott
& W. Magruder; numbers for freshwater and terrestrial species are only estimates, except for Vis et.
al. (1994) who reported 34 species; all other existing literature is outdated and unreliable (e.g.,
MacCaughey 1918).
Protozoa: Chave (1987) noted more than 1000 species in Hawaiian waters; Philipps (1977)
reported on more than 400 species from depths less than 100 ft; Larsen & Patterson (1990) reported
31 benthic marine flagellate species, including 2 new species.
Fungi: Baker & Goos (1972), following Parris (1940) recorded 683 species, including
Myxomycota (slime molds) but not lichens; Goos (1977) recorded 900 species of Fungi Imperfecti;
Baker (1977) noted 1591 species of fungi from Hawaii and the central Pacific islands. Undoubtedly,
the true number is much higher because the fungi are poorly studied in Hawaii (see also Goos &
Gowing 1992). Hawksworth (1994) suggested that in a given geographical area there are about 6
times as many fungi in all habitats as there are native and naturalized plant species, which suggests
that there may be 12,000 species of fungi. Endemism is very low (Baker & Goos 1972, Baker 1977,
Goos 1977). Kohlmeyer (1969) recorded 27 species of marine fungi including 2 nonindigenous
species.
Lichens: Smith (1991) recorded 723 species, suggesting that there may be more than 800
species (his 1991 numbers include taxonomic changes published in Smith 1993). He recorded 240
species as supposedly endemic and suggested that the true number may be lower; 1 genus
(Ramalinopsis) is endemic. See also Stenroos & Smith (1993).
Ferns (Polypodiophyta) and fern allies (Psilophyta and Lycophyta): Only native and natu-
ralized (e.g., growing in the wild) species are listed; more than 200 additional species are cultivated
(G. Staples, pers. comm.)
Angiosperms: Only native and naturalized species are listed; on the order of 8000 addition-
al species are cultivated in agricultural settings and home gardens (G. Staples, pers. comm.). 
Porifera: Of 80 species listed, 5 are known only from depths greater than 100 m, 5 recorded
only from the Waikiki Aquarium, and 8 that might be considered doubtful species (Bergquist 1977).
Cnidaria: Octocorallia: Of 105 species listed, 4 are shallow water (Devaney 1977b),
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remaining are 1 species of bamboo coral (Muzik 1978) and 101 species of deep-water gorgonians
(Grigg & Bayer 1976, Versevelt & Bayer1988, Bayer 1990).
Cnidaria: Scleractinia: Of 98 species listed, 42 found in shallow water (Maragos 1977), 49
in deeper water (Cairns 1984), 3 from northwest islands (Grigg et al. 1981), 5 new records in
Maragos (1995); of 91 published species, 50 are exclusively ahermatypic, 37  are exclusively her-
matypic, 4 are facultative species (Cairns 1984).
Platyhelminthes (Parasitic): Of 511 species listed, 147 monogenetic species including 132
species described from 121 species of Hawaiian fishes (Yamaguti 1968b). Of 357 digenetic species,
314 species including 227 species described from Hawaiian fishes. Also 30 species not cited
(Yamaguti 1970), 13 from Yamaguti (1965), 7 species of cestodes (Yamaguti 1968a), and non-
indigenous, marine, cage-cultured, freshwater tilapia with 1 species of marine monogean (Kaneko et
al. 1988).
Nemertinea: Of the 28 species in the table, 6 species from Coe (1947), 7 benthic and 12
pelagic undescribed/unreported species, 2 introduced terrestrial species (J. Norenburg, pers. comm.)
plus 1 parasitic (Humes 1942).
Annelida: Oligochaeta: Of 26 species listed, 23 Tubificidae (Ers?us & Davis 1989); 1 new
species by Ers?us (1990); the other 2 are unidentified species (Bailey-Brock 1987).
Annelida: Polychaeta: Approximately 100 additional species are known, but not yet report-
ed from Hawaiian waters (J. Bailey-Brock, pers. comm.).
Nematoda: zooparasitic: The figure given is very incomplete. Of 33 species, 22 are from
intermingling herds of mammals on Molokai (McKenzie & Davidson 1989); 2 from monk seals
(Chapin 1935), 8 larval forms from fishes (Deardorff et al. 1982), 1 parasitic record from an elas-
mobranch (Deardorff 1987).
Mollusca: Gastropoda: There are 763 nomenclaturally correct native land species; all
endemic except 2?4 indigenous but not endemic species (Cowie et al., in press), approximately 50
nonindigenous species (R.H. Cowie, pers. comm.); 7 endemic freshwater snails, approximately 30
nonindigenous species (R.H. Cowie, pers. comm.).
Mollusca: Cephalopoda: Of 73 species in the table, only 4 considered benthic (Kay 1979;
Roper et al. 1984); for other pelagic species see Roper & Young (1975) and Young (1991).
Crustacea: Branchiopoda: Four species from Mauna Kea at elevations from 6500? 13,700
ft [1980?4175 m] (Ueno 1936).
Crustacea: Ostracoda: Numerous species; J.C. Holden (MS) made identifications to only
genera, he noted more than 100 fossil species; 30 benthic species (Kornicker 1976, Danielopol &
Hartmann 1986, Hartmann, 1991); 1 freshwater species (V?vra 1906).
Crustacea: Copepoda: The 92 species listed include planktonic, benthic, and ectoparasitic
forms; 5 insterstitial species (Wells 1986).
Crustacea: Cirripedia: Of 58 species in the table, 11 are from deep water including 8 new
species (Pilsbry 1907); 46 shallow-water species includes 3 acrothoracican species (W.A. Newman,
MS key); 1 rhizocephalan parasite (Boschma 1953). 
Crustacea: Isopoda: Of 27 marine species, 3 were collected in deep water (Richardson
1906); 6 species are parasitic epicarideans (Danforth 1970); of 51 terrestrial species 15 are endemic
and 25 nonindigenous species.
Crustacea: Amphipoda: Of 172 species listed, 121 gammarideans (Barnard 1970), 4 gam-
marideans (Brusca 1973), 7 caprellids, 40 hyperiids (Brusca 1973, 1978).
Entoprocta: Soule et al. (1987) provided only unidentified generic descriptions; 55 species
reported from Kaneohe Bay (Dade & Honkalehto 1986).
Chaetognatha: Seven pelagic species from distribution maps (Pierrot-Bults & Nair 1991); 1
benthic species described from Kure Atoll, Hawaii (Alvari?o 1978).
Echinodermata: Crinoidea: Of 16 species listed in the table, 11 described as new species
by Clark (1908); no shallow-water forms.
RECORDS OF THE HAWAII BIOLOGICAL SURVEY FOR 1994 9
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Echinodermata: Asteroidea: Of 82 species listed, 18 found in shallow water and 64 in
deeper water; 1 species and 2 subspecies of shallow-water forms considered endemic (Devaney &
Baker MS); 50 species endemic to deeper waters described as new (Fisher 1906).
Echinodermata: Echinoidea: Of 75 species, 21 found in shallow water (Eldredge MS), 53
in deeper water; Cylpeaster eurypetalus known only from Pearl and Hermes Reef, and Midway and
Kure atolls; 35 new species described from Albatross material.
Echinodermata: Holothuroidea: Of 48+ species in the table, 24 found in shallow water
(Eldredge, MS), 24 in deeper water; 3 shallow-water and 1 deep-water for considered endemic (Clark
& Rowe 1971); Fisher (1907) described 19 new species out of 46 reported; reports (unpublished) of
additional species are known.
Echinodermata: Ophiuroidea: Of 57 species, 19 found in shallow water (Devaney &
Baker MS), 38 in deeper water; 6 shallow-water and 22-deep-water forms endemic.
Chordata: Urochordata: Thaliacea: 19 species of Salpidae?cosmopolitan oceanic
planktonic organisms in the circumglobal warm water zone (Yount 1958); 1 species of Pyrosoma
(Metcalf & Hopkins 1919). 
Chordata: Pisces: The 1150+ marine species listed include all species from all depths to 200
miles from shore?no freshwater, no undescribed, and no doubtful species are included; 1285 species
when all conditions provided (B. Mundy, pers. comm.); Randall (1992) listed 536 species of shore
fishes, with 25% endemicity.
Chordata: Aves: The 274 species listed is an all-inclusive contemporary list: 60 resident
native species, 46 resident nonindigenous species, 13 breeding visitor species, 155 nonbreeding vis-
itor species. There are also 16 species extinct since Captain Cook, 35+ extinct before Captain Cook
(subfossil), and 150? nonindigenous species not established bringing the total number of species
known to occur or be sighted in Hawaii to 475? (R.L. Pyle, pers. comm). [Pyle (1992) listed 131 bird
species; 53 nonindigenous species.] Thirty-two fossil species have been described by Olson & James
(1991) and James & Olson (1991).
Chordata: Mammalia: Of 20 species of terrestrial mammals 19 are free-ranging species
(the horse is no longer feral) and 1 species of bat (an additional undescribed subfossil species of bat
is known; F.G. Howarth, pers. comm.); of 24 marine species, 2 are littoral marine (seals) and 22 are
pelagic (whales and dolphins) (Tomich 1986).
Summary
From literature and unpublished sources, approximately 21,383 species have been
recorded from the Hawaiian Islands and surrounding waters. Of these, 8,759 are endem-
ic to the Hawaiian Islands, and 4,532 are nonindigenous species of protists, fungi, plants,
and animals. Of these approximately 15,000 species are terrestrial, 300 are found in fresh-
water, and 5,500 are marine-inhabiting. Endemism varies from very low (e.g., lower
plants and many marine organisms) to very high (e.g., insects and snails). However, many
gaps in the knowledge of Hawaii?s biota (especially the smaller organisms?viruses, bac-
teria, protists, freshwater algae, etc.), still remain.
Acknowledgements
The authors thank the following people and institutions for their assistance and sup-
port in compiling the data in this article: A. Allison, W. Appleby, B.L. Burch, T.A. Burch,
R.H. Cowie, N.L. Evenhuis, D.R. Herbst, W.J. Hoe, F.G. Howarth, C. Imada, C.
Kishinami, G.M. Nishida, D.A. Polhemus, R.L. Pyle, G. Staples, and A. Suzumoto (all
from Bishop Museum, Honolulu, USA); I.A. Abbott, J. Bailey-Brock, M. Hadfield, C.W.
Smith, and G. Wong (all from University of Hawai at Manoa, Honolulu, USA); E.P.
BISHOP MUSEUM OCCASIONAL PAPERS:  NO. 41, 199510
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Caswell, E.M. Noffsinger (University of California, Davis, California, USA), R. Galt
(California State University, Long Beach, California, USA), S.W. James (Maharishi
International University, Fairfield, Iowa, USA), D.L. Hawksworth, P.M. Kirk (Inter-
national Mycological Institute, Egham, Surrey, UK), G. Mason (University of Otago,
Dunedin, New Zealand), D. Farr (U.S. Department of Agriculture, Beltsville, Maryland,
USA), B. Mundy (National Marine Fisheries Service, Honolulu, Hawaii, USA), J. Nor-
enberg and W.L. Wagner (Smithsonian Institution, Washington, D.C., USA), P.N. Turner
(Christopher Newport University, Newport News, Virginia, USA); the staff of the libraries
at the Bishop Museum and the University of Hawaii at Manoa; the National Science
Foundation; and the John D. and Catherine T. MacArthur Foundation.
References
Alvari?o, A. 1978. Spadella gaetanoi, a new benthic chaetognath from Hawaii. Proc.
Biol. Soc. Wash. 91(3): 650?57.
Bailey-Brock, J.H. 1987. Phylum Annelida, p. 213?16. In Devaney, D.M. & L.G.
Eldredge, eds., Reef and shore fauna of Hawaii. Section 2: Platyhelminthes through
Phoronida and Section 3: Sipuncula through Annelida. Bishop Museum Press,
Honolulu.
???. 1990. Polydora nuchalis (Polychaeta: Spionidae), a new Hawaiian record from
aquaculture ponds. Pac. Sci. 44(1): 81?87.
???. 1991. Tubeworms (Serpulidae, Polychaeta) collected from sewage outfalls, coral
reefs and deep water off the Hawaiian Islands. Bull. Mar. Sci. 48(2): 198?207.
???. & O. Hartman. 1987. Class Polychaeta, p. 216?454. In Devaney, D.M. & L.G.
Eldredge, eds., Reef and shore fauna of Hawaii. Section 2: Platyhelminthes through
Phoronida and Section 3: Sipuncula through Annelida. Bishop Museum Press,
Honolulu.
Baker, G.E. 1977. The prospect for mycology in the Central Pacific. Harold L. Lyon
Arboretum Lecture 8, 51 p.
???. & R.D. Goos. 1972. Endemism and evolution in the Hawaiian biota: Fungi, p.
409?31. In E. A. Kay, ed., A natural history of the Hawaiian Islands: selected read-
ings. University of Hawai Press, Honolulu.
Ball, C.H. 1963. Cephaloidophora carpilodei n. sp. and C. pinquis n. sp., gregarines of a
xanthid crab in Hawaii. J. Protozool. 10(3): 321?27.
Barnard, J.L. 1970. Sublittoral Gammaridea (Amphipoda) of the Hawaiian Islands.
Smithson. Contrib. Zool. 3, 286 p.
Bayer, F.M. 1990. A new isidid octocoral (Anthozoa: Gorgonacea) from New Caledonia,
with descriptions of other new species from elsewhere in the Pacific Ocean. Proc.
Biol. Soc. Wash. 103(1): 205?28.
Bergquist, P.R. 1977. Porifera, p. 53?69. In Devaney, D.M. & L.G. Eldredge, eds., Reef
and shore fauna of Hawaii. Section 1: Protozoa through Ctenophora. Bishop
Museum Press, Honolulu.
Berry, S. 1914. The Cephalopoda of the Hawaiian Islands. Bull. Bur. Fish. 23: 255?362.
Boschma, H. 1953. The Rhizocephala of the Pacific. Zool. Meded. 32(17): 185?210.
Boyko, C.B. 1994. Catalog of recent type specimens in the Department of Invertebrates,
American Museum of Natural History. I. Micro-invertebrates (phyla Sacromastigo-
phora, Gnathostomula, Gastrotricha, Rotifera, and Tardigrada). Am. Mus. Novit.
3106, 44 p.
RECORDS OF THE HAWAII BIOLOGICAL SURVEY FOR 1994 11
Electronic version via HBS web site: http://www.bishop.hawaii.org/bishop/HBS/ ?1995,1997 HBS
Brinton, E. 1962. The distribution of Pacific euphausiids. Bull. Scripps Inst. Oceanogr.
8(2): 51?279.
Brusca, G.J. 1973. Pelagic Amphipoda from the waters near Oahu, Hawaii, excluding the
family Scinidae. Pac. Sci. 27(1): 8?27.
???. 1978. Contribution to the knowledge of hyperiid amphipods of the family
Scinidae from near Hawaii, with a description of a new species, Scina hawaiensis.
Pac. Sci. 32(3): 281?92.
Burch, B.L. 1978. Asian clam, Corbicula, threatens Hawaii. The Nautilus 92(1): 54?55.
Burch, B.L. & T.A. Burch. 1995. New Hawaiian records for Stephanoscyphus simplex
Kirkpatrick (Cnidaria: Scyphozoa). Bishop Mus. Occas. Pap. 42: 53.
Cairns, S.D. 1984. New records of ahermatypic corals (Scleractinia) from the Hawaiian
and Line Islands. Occas. Pap. B. P. Bishop Mus. 25(10): 1?30.
Chapin, E.A. 1925. Descriptions of new internal parasites. Proc. U.S. Natl. Mus.
68(2603): 1?4.
Chave, E.H. 1987. Common living benthic Foraminifera in Mamala Bay, Hawaii, with
descriptions of two new species. Bishop Mus. Occas. Pap. 27: 25?72.
???. & A.T. Jones. 1991. Deep-water megafauna of the Kohala and Haleakala slopes,
Alenuihaha Channel, Hawaii. Deep-Sea Res. 38(7): 781?803.
Child, C.A. 1972. A new species of Anoropallene (Pycnogonida) from the Hawaiian
Islands. Proc. Biol. Soc. Wash. 85(11): 147?50.
Choy, B.K., G.H. Balazs & M. Dailey. 1989. A new therapy for marine turtles para-
sitized by the piscicolid leech, Ozobranchus brachiatus. Herp. Rev. 20(4): 89?90.
Clark, A.H. 1949. Ophiuroidea of the Hawaiian Islands. B.P. Bishop Mus. Bull. 195, 133
p.
Clark, A.M. & F.W.E. Rowe. 1971. Monograph of shallow-water Indo-west Pacific
echinoderms. British Museum (Natural History), London. 234 p.
Coe, W.R. 1947. Nemerteans of the Hawaiian and Marshall Islands. Occas. Pap. B. P.
Bishop Mus. 19(3): 101?06.
Cooke, W.J. 1977. Order Hydroida, p.71?104. In Devaney, D.M. & L.G. Eldredge, eds.,
Reef and shore fauna of Hawaii. Section 1: Protozoa through Ctenophora. Bishop
Museum Press, Honolulu.
???. 1984. New scyphozoan records for Hawaii: Anomalorhiza shawi Light, 1921,
and Thysanostoma loriferum (Ehrenberg, 1835); with notes on several other rhizos-
tomes. Proc. Biol. Soc. Wash. 97(3): 583?88.
Cowie, R.H., N.L. Evenhuis & C.C. Christensen. 1995. Catalog of the native land and
freshwater molluscs of the Hawaiian Islands. Bishop Mus. Bull. Zool. 3: in press.
Cutress, C.E. 1977. Orders Corallimorpharia, Actiniaria, Ceriantharia, p. 130?147. In
Devaney, D.M. & L.G. Eldredge, eds., Reef and shore fauna of Hawaii. Section 1.
Protozoa through Ctenophora. Bishop Museum Press, Honolulu.
Dade, W.B. & T. Honkalehto. 1986. Common ectoproct bryozoans of Kaneohe Bay, p.
52?65. In: Jokiel, P.L., R.H. Richmond & R.A. Rogers, eds., Coral reef population
biology. Hawaii Institute of Marine Biology Technical Report 37.
Dailey, M.D., M.L. Fast & G.H. Balazs. 1992. A survey of the Trematoda (Platyhel-
minthes: Digenea) parasitic in green turtles, Chelonia mydas (L.) from Hawaii. Bull.
South. Calif. Acad. Sci. 91(2): 84?91.
BISHOP MUSEUM OCCASIONAL PAPERS:  NO. 41, 199512
Electronic version via HBS web site: http://www.bishop.hawaii.org/bishop/HBS/ ?1995,1997 HBS
Danforth, C.G. 1970. Epicaridea (Isopoda) of Hawaii. Bull. South. Calif. Acad. Sci.
69(1): 27?31.
Danielopol, D.L. & G. Hartmann. 1986. Ostracoda. Part 1: stygobiont Ostracoda from
inland subterranean waters, p. 265?94. In: Botosaneanu, L., ed., Stygofauna mundi.
E.J. Brill, Leiden.
Deardorff, T.L. 1987. Redescription of Pulchrascaris chiloscylli (Johnston and Mawson,
1951) (Nematoda: Anisakidae), with comments on species in Pulchrascaris and
Terranova. Proc. Helminthol. Soc. Wash. 54(1): 28?39.
???. M.M. Kliks, M.E. Rosenfeld, R.A. Rychlinski & R.S. Desowitz. 1982. Larval
ascaridoid nematodes from fishes near the Hawaiian Islands, with comments on
pathogenicity experiments. Pac. Sci. 36(2): 187?201.
Devaney, D.M. 1977a. Ctenophora, p. 262?69. In Devaney, D.M. & L.G. Eldredge, eds.,
Reef and shore fauna of Hawaii. Section 1: Protozoa through Ctenophora. Bishop
Museum Press, Honolulu.
???. 1977b. Subclass Octocorallia, p. 119?29. In Devaney, D.M. & L.G. Eldredge,
eds., Reef and shore fauna of Hawaii. Section 1: Protozoa through Ctenophora.
Bishop Museum Press, Honolulu.
Devaney, D.M. & L.G. Eldredge. 1977. Class Scyphozoa, p. 108?18. In Devaney, D.M.
& L.G. Eldredge, eds., Reef and shore fauna of Hawaii. Section 1: Protozoa through
Ctenophora. Bishop Museum Press, Honolulu.
???. 1987. Phylum Nemertea (Rhynchocoela), p. 59?69. In Devaney, D.M. & L.G.
Eldredge, eds., Reef and shore fauna of Hawaii. Section 2: Platyhelminthes through
Phoronida and Section 3: Sipuncula through Annelida. Bishop Museum Press,
Honolulu. 
Devick, W.S., J.M. Fitzsimons & R.T. Nishimoto. 1992. Conservation of Hawaiian
freshwater fishes. State of Hawaii Division of Aquatic Resources. 26 p.
Edmonds, S.J. 1987. Phyla Sipuncula and Echiura, p. 185?212. In Devaney, D.M. & L.G.
Eldredge, eds., Reef and shore fauna of Hawaii. Section 2: Platyhelminthes through
Phoronida and Section 3: Sipuncula through Annelida. Bishop Museum Press,
Honolulu.
Eliasson, U.H. 1991. The myxomycete biota of the Hawaiian Islands. Mycol. Res. 95:
257?67.
Eldredge, L.G. 1967. Record of a lancelet from Hawaii. Pac. Sci. 21(4): 564.
???. 1994. Introductions of commercially significant aquatic organisms to the Pacific
islands. South Pacific Commission, Noumea. 127 p.
Eldredge, L.G. & D.M. Devaney. 1977. Other hydrozoans, p. 105?07. In Devaney, D.M.
& L.G. Eldredge, eds., Reef and shore fauna of Hawaii. Section 1: Protozoa through
Ctenophora. Bishop Museum Press, Honolulu.
Emig, C.C. 1987. Phylum Brachiopoda, p. 167?70. In Devaney, D.M. & L.G. Eldredge,
eds., Reef and shore fauna of Hawaii. Section 2: Platyhelminthes through Phoronida
and Section 3: Sipuncula through Annelida. Bishop Museum Press, Honolulu.
???. & J.H. Bailey-Brock. 1987. Phylum Phoronida, p. 171?81. In Devaney, D.M. &
L.G. Eldredge, eds., Reef and shore fauna of Hawaii. Section 2: Platyhelminthes
through Phoronida and Section 3: Sipuncula through Annelida. Bishop Museum
Press, Honolulu.
Ers?us, C. 1990. Marine Oligochaeta of Hong Kong, p. 259?335. In: Morton, B., ed., The
RECORDS OF THE HAWAII BIOLOGICAL SURVEY FOR 1994 13
Electronic version via HBS web site: http://www.bishop.hawaii.org/bishop/HBS/ ?1995,1997 HBS
marine flora and fauna of Hong Kong and southern China 2. Vol. 1. Introduction and
taxonomy. Hong Kong University Press, Hong Kong.
???. & D. Davis. 1989. The marine Tubificidae (Oligochaeta) of Hawaii. Asian Mar.
Biol. 6: 73?100.
Fauchald, K. 1992. A review of the genus Eunice (Polychaeta: Eunicidae) based upon
type material. Smithson. Contrib. Zool. 523, 422 p.
Fisher, W.K. 1906. The starfishes of the Hawaiian Islands. Bull. U.S. Fish Commn. 23(3):
987?1130.
???. 1907. The holothurians of the Hawaiian Islands. Proc. U.S. Natl. Mus. 32(1555):
637?744.
Frank, J.H. & E.D. McCoy. 1990. Endemics and epidemics of shibboleths and other
things causing chaos. Fla. Entomol. 73(1): 1?9.
Goos, R.D. 1977. Comparative observations of the Hyphomycetes of Hawaii and Central
America. Kavaka 6: 31?35.
???. & D.P. Gowing. 1992. Type specimens of fungi maintained at Herbarium
Pacificum, Bernice P. Bishop Museum, Honolulu. Mycotaxon 43: 177?98.
Grigg, R.W. & F.M. Bayer. 1976. Present knowledge of the systematic and zoogeogra-
phy of the Order Gorgonacea in Hawaii. Pac. Sci. 30(2): 167?75.
???. & D. Opresko. 1977. Order Antipatharia, p. 242?61. In Devaney, D.M. & L.G.
Eldredge, eds., Reef and shore fauna of Hawaii. Section 1: Protozoa through
Ctenophora. Bishop Museum Press, Honolulu.
???. J.W. Wells & C. Wallace. 1981. Acropora in Hawaii. Part 1. History of the sci-
entific record, systematics, and ecology. Pac. Sci. 35(1): 1?13.
Hadfield, M.G. & R.E. Young. 1983. Planctosphaera (Hemichordata: Enteropneusta) in
the Pacific Ocean. Mar. Biol. 73(2): 151?53.
Hartmann, G. 1991. Ostracoden von Hawaii, inbesondere aus dem marinen Interstitial.
Helgol?nd. Meeresuntersuch. 45: 165?98.
Hawksworth, D.L. 1994. The tropical fungal biota: census, pertinence, prophylaxis, and
prognosis, p. 265?93 In: Issac, S., J.C. Frankland, A.J. Whalley & R. Watling, eds.,
Tropical mycology. Cambridge University Press, Cambridge.
Hering, J.L. 1961. The genus Halobates (Hemiptera: Gerridae). Pac. Insects 3(2?3):
223?305.
???. 1965. Hermatobates, a new generic record for the Atlantic Ocean, with descrip-
tions of new species (Hemiptera: Gerridae). Proc. U.S. Natl. Mus. 117(3510):
123?30. 
Hoe, W.J. 1974. Annotated checklist of Hawaiian mosses. Lyonia 1: 1?45.
???. 1979. The phytogeographical relationships of Hawaiian mosses. Unpubl. Ph.D.
Dissertation, University of Hawaii, Honolulu. 357 p.
Hope, W.D. 1987. The Aschelminth complex, p. 71?81. In Devaney, D.M. & L.G.
Eldredge, eds., Reef and shore fauna of Hawaii. Section 2: Platyhelminthes through
Phoronida and Section 3: Sipuncula through Annelida. Bishop Museum Press,
Honolulu.
Howarth, F.G. 1990. Hawaiian terrestrial arthropods: an overview. Bishop Mus. Occas.
Pap. 30: 4?26.
???. & J. Moore. 1983. The land nermetine Argonemertes dendyi (Dakin) in Hawaii
(Nemertinea: Hoplonemertinea: Prosorhochmidae). Pac. Sci. 37(2): 141?44.
BISHOP MUSEUM OCCASIONAL PAPERS:  NO. 41, 199514
Electronic version via HBS web site: http://www.bishop.hawaii.org/bishop/HBS/ ?1995,1997 HBS
Humes, A.G. 1942. The morphology, taxonomy, and bionomics of the nemertean genus
Carcinonemertes. Ill. Biol. Monogr. 18(4): 3?105.
???. 1976. Distribution and hosts of Stellicola (Copepoda, Cyclopoida) associated
with Linckia (Asteroidea) in the Indo-west Pacific. Beaufortia 25(321): 49?61.
Hyman, L.H. 1939. Land planarians from the Hawaiian Islands. Arch. Zool. Exper. Gen.
80(3): 116?24.
James, H.F. & S.L. Olson. 1991. Descriptions of thirty-two species of birds from the
Hawaiian Islands: Part II. Passeriformes. Ornithol. Monogr.46: 1?88.
Kaneko, J.J., II, R. Yamada, J.A. Block & R. Nakamura. 1988. Infection of tilapia,
Oreochromis mossambicus (Trewavas), by a marine monogenean, Neobenedenia
melleni (MacCallum, 1927) Yamaguti, 1963 in Kaneohe Bay, Hawaii, USA, and its
treatment. J. Fish Diseases 11: 295?300.
Kawakatsu, M. & R.W. Mitchell. 1984. Oahuhawaiiana kazukolinda gen. et sp. nov.
(Turbellaria, Tricladida, Maricola), a new freshwater planarian from Honolulu, Oahu
Island, Hawaii, U.S.A. Zool. Sci. 1(3): 487?500.
???. R.W. Mitchell, Y. Hirao & I. Tanaka. 1984. Occurrence of Dugesia doroto-
cephala (Woodworth, 1897) (Turbellaria, Tricladida, Paludicola) in Honolulu,
Hawaii. Biol. Mag. (Okinawa) 22: 1?9.
Kay, E.A. 1979. Hawaiian marine shells. Reef and shore fauna of Hawaii. Section 4:
Mollusca. Bishop Museum Press, Honolulu. 652 p.
???. 1994. A natural history of the Hawaiian islands. Selected readings II. University
of Hawaii Press, Honolulu. xi + 520 p.
???. & S.R. Palumbi. 1987. Endemism and evolution in Hawaiian marine inverte-
brates. Trends Ecol. Evol. 2(7): 183?86.
Kitalong, A.E. 1986. A preliminary study on the emergence patterns of microfauna in
Kaneohe Bay, Hawaii, p. 414?23. In: Jokiel, P.L., R.H. Richmond & R.A. Rogers,
eds., Coral reef population biology. Hawaii Institute of Marine Biology Technical
Report 37.
Kohlmeyer, J. 1969. Marine fungi of Hawaii including the new genus Helicascus.
Canad. J. Bot. 47: 1469?1487.
Kornicker, L.S. 1976. Benthic marine Cypridinacea from Hawaii (Ostracoda). Smithson.
Contrib. Zool. 231, 24 p.
Kunz, H. 1993. Beitrag zur Kenntnis von Zwei Psammopsyllus-Arten (Copepoda,
Harpacticoida, Cylindropsyllidae). Crustaceana 64(2): 143?54.
Larsen, J. & D.J. Patterson. 1990. Some flagellates (Protista) from tropical marine sed-
iments. J. Nat. Hist. 24: 801?937.
Lewis, A.G. 1966. Copepod crustaceans parasitic on elasmobranch fishes of the Hawaiian
Islands. Proc. U.S. Natl. Mus. 118(3524): 57?154.
???. 1967. Copepod crustaceans parasitic on teleost fishes of the Hawaiian Islands.
Proc. U.S. Natl. Mus. 121(3574): 1?204.
MacCaughey, V. 1918. Algae of the Hawaiian Archipelago. Bot. Gaz. 65: 42?57, 121?49.
Maciolek, J.A. 1984. Exotic fishes in Hawaii and other islands of Oceania, p. 131?61. In:
Courtenay, W.R., Jr. & J.R. Stauffer, eds., Distribution, biology, and management of
exotic fishes. Johns Hopkins University Press, Baltimore.
Maragos, J.E. 1977. Order Scleractinia, p. 158?241. In Devaney, D.M. & L.G. Eldredge,
eds., Reef and shore fauna of Hawaii. Section 1: Protozoa through Ctenophora.
RECORDS OF THE HAWAII BIOLOGICAL SURVEY FOR 1994 15
Electronic version via HBS web site: http://www.bishop.hawaii.org/bishop/HBS/ ?1995,1997 HBS
Bishop Museum Press, Honolulu.
???. 1995. Revised checklist of extant shallow water stony coral species from Hawaii
(Cnidaria: Anthozoa: Scleractinia). Bishop Mus. Occas. Pap. 42: 53?54.
Margulis, L. & K.V. Schwartz. 1988. Five kingdoms: an illustrated guide to the phyla
of life on earth. Second edition. W.H. Freeman & Co., New York. 376 p.
Matthews, D.C. 1966. A comparative study of Craspedacusta sowerbyi and Calpasoma
dactyloptera life cycles. Pac. Sci. 20(2): 246?59.
McInnes, S.J. 1994. Zoogeographic distribution of terrestrial/freshwater tardigrades from
current literature. J. Nat. Hist. 28(2): 257?352.
McKenzie, M.E. & W.R. Davidson. 1989. Helminth parasites of intermingling axis deer,
wild swine and domestic cattle from the island of Molokai, Hawaii. J. Wildl.
Diseases 25(2): 252?57.
Metcalf, M.M. & H. S. Hopkins. 1919. Pyrosoma?taxonomic study based upon the col-
lections of the United States Bureau of Fisheries and the United States National
Museum. Bull. U.S. Natl. Mus. 100 2(3): 195?272.
Miller, H.A. & H. O. Whittier. 1990. Bryophyte floras of tropical Pacific islands. Trop.
Bryol. 2: 167?75.
Moore, J.P. 1946. Leeches (Hirudinea) from the Hawaiian Islands, and two new species
from the Pacific region in the Bishop Museum collection. Occas. Pap. B. P. Bishop
Mus. 18(11): 171?91.
Motoda, S. 1963. Coryceus and Farranula copepods (Cyclopoida) in Hawaiian waters.
Publ. Seto Mar. Biol. Lab. 11(2): 209?62.
Muzik, K. 1978. A bioluminescent gorgonian, Lepidisis olapa, new species (Coelen-
terata: Octocorallia), from Hawaii. Bull. Mar. Sci. 28(4): 735?41.
Nakamura, M. 1992. An ecological study of earthworms in Papua New Guinea. J. Chuo
Univ. 13: 19?33.
Nishida, G.M., ed. 1994. Hawaiian terrestrial arthropod checklist. Second edition. Bishop
Mus. Tech. Rep. 4, 287 p.
Nutting, C.C. 1905. Hydroids of the Hawaiian Islands collected by the steamer Albatross
in 1902. Bull. U.S. Fish Commn. 23(3): 931?59.
Oliveira, J.M. 1940. Plant-parasitic and free-living nematodes in Hawaii. B. P. Bishop
Mus. Occas. Pap. 15(29): 361?73.
Olson, S.L. & H.F. James. 1991. Descriptions of thirty-two new species of birds from
the Hawaiian Islands: Part I. Non-Passeriformes. Ornithol. Monogr. 45: 1?88.
Ortmann, A.E. 1906. Schizopods of the Hawaiian Islands collected by the steamer
Albatross in 1902. Bull. U.S. Fish Commn. 23(3): 961?73.
Otte, D. 1994. The crickets of Hawaii. Origin, systematics and evolution. Orthopterist?s
Society, Philadelphia. vi + 396 p.
Parker, S.P., ed. 1982. Synopsis and classification of living organisms. 2 vols. McGraw-
Hill Book Co., New York.
Parris, G.K. 1940. A check list of fungi, bacteria, nematodes, and viruses occurring in
Hawaii, and their hosts. Plant Disease Reporter, Supplement 121, 91 p.
Paxton, H. & J.H. Bailey-Brock. 1986. Diopatra dexiognatha, a new species of Onu-
phidae (Polychaeta) from Oahu, Hawaiian Islands. Pac. Sci. 40(1-4): 16.
Pearse, V.B. 1989. Growth and behavior of Trichoplax adhaerens: first record of the phy-
lum Placozoa in Hawaii. Pac. Sci. 43(2): 117?21.
BISHOP MUSEUM OCCASIONAL PAPERS:  NO. 41, 199516
Electronic version via HBS web site: http://www.bishop.hawaii.org/bishop/HBS/ ?1995,1997 HBS
Pettibone, M.H. 1986. Review of the Iphioninae (Polychaeta: Polynoidae) and revision
of Iphione cimex Quatrefages, Gattyana deludens Fauvel, and Harmothoe iphionel-
loides Johnson (Harmothoinae). Smithson. Contrib. Zool. 428, 43 p.
Phillips, F.J. 1977. Protozoa, p. 12?52. In Devaney, D.M. & L.G. Eldredge, eds., Reef
and shore fauna of Hawaii. Section 1: Protozoa through Ctenophora. Bishop
Museum Press, Honolulu.
Pierrot-Bults, A. C. & V. R. Nair. 1991. Distribution patterns in Chaetognatha, p.
86?116. In: Bone, Q., H. Kapp & A.C. Pierrot-Bults, eds., The biology of chaetog-
naths. Oxford University Press, New York.
Pilsbry, H. A. 1907. Hawaiian Cirripedia. Bull. Bur. Fish. 26: 179?190.0
Poulter, J. 1987. Phylum Platyhelminthes, p. 13?58. In Devaney, D.M. & L.G. Eldredge,
eds., Reef and shore fauna of Hawaii. Section 2: Platyhelminthes through Phoronida
and Section 3: Sipuncula through Annelida. Bishop Museum Press, Honolulu.
Pyle, R.L. 1992. Checklist of the birds of Hawaii?1992. ?Elepaio 42: 43?62.
Raabe, R.D. 1981. Checklist of plant diseases in Hawaii: including records of micro-
organisms, principally fungi, found in the state. Enlarged and revised by Ibra L.
Conners and Albert R. Martinez. Hawaii Institute of Tropical Agriculture and Human
Resources, College of Tropical Agriculture and Human Resources, Uni-versity of
Hawaii at Manoa. 313 p.
Randall, J.E. 1992. Endemism of fishes in Oceania. UNEP Regional Seas Rep. Stud. 147:
55?67.
Richardson, H. 1906. Isopods collected at the Hawaiian Islands by the U.S. Fish Com-
mission steamer Albatross. Bull. U.S. Fish Commn. 23(3): 819?26.
Roper, C.F.E. & R.E. Young. 1975. Vertical distribution of pelagic cephalopods.
Smithson. Contrib. Zool. 209, 51 p.
???. M.J. Sweeney & C.E. Nauen. 1984. FAO species catalogue. Vol. 3. Cephalo-
pods of the world. An annotated and illustrated catalogue of species of interest to
fisheries. FAO Fish. Synop. 125, 3: 1?277.
Smith, C.W. 1991. Lichen conservation in Hawaii, p. 35?45. In D.J. Galloway, ed.,
Tropical lichens: their systematics, conservation, and ecology. Systematics Associ-
ation Special Volume 43. Clarendon Press, Oxford.
???. 1993. Notes on Hawaiian parmelioid lichens. Bryologist 96: 326?32.
Soest, R.W.M. van. 1974a. Taxonomy of the subfamily Cyclosalpidae Yount, 1954 (Tun-
icata, Thaliacea), with descriptions of two new species. Beaufortia 22(288): 17?55.
???. 1974b. A revision of the genera Salpa Forsk?l, 1775, Pegea Savigny, 1816, and
Ritteriella Metcalf, 1919 (Tunicata, Thaliacea). Beaufortia 22(293): 153?91.
Soule, J.D., D.F. Soule & H.W. Chaney. 1987. Phyla Entoprocta and Bryozoa (Ecto-
procta), p. 83?166. In Devaney, D.M. & L.G. Eldredge, eds., Reef and shore fauna
of Hawaii. Section 2: Platyhelminthes through Phoronida and Section 3: Sipuncula
through Annelida. Bishop Museum Press, Honolulu. 
Southward, E.C. 1980. Two new species of Pogonophora from Hawaii. Pac. Sci. 34(4):
371?78.
Stenroos, S. & C.W. Smith. 1993. Notes on the amphipacific relations of Hawaiian Cla-
doniaceae. Trop. Biol. 8: 275?80.
Sterrer, W. 1991. Gnathostomulida from Hawaii. Zool. Scripta 20(2): 129?36.
Svihla, A. 1941. The occurrence of freshwater sponges in the Hawaiian Islands. Science
94(2438): 278?79.
RECORDS OF THE HAWAII BIOLOGICAL SURVEY FOR 1994 17
Electronic version via HBS web site: http://www.bishop.hawaii.org/bishop/HBS/ ?1995,1997 HBS
Taguchi, S. 1982. Seasonal study of fecal pellets and discarded houses of Appendicularia
in a subtropical inlet, Kaneohe Bay, Hawaii. Estuar. Coast. Shelf Sci. 14: 545?55.
Tokioka, R. 1967. Pacific Tunicata of the United States National Museum. U.S. Natl.
Mus. Bull. 251, 247 p.
Tomich, P.Q. 1986. Mammals in Hawaii. A synopsis and notational bibliography. Sec-
ond edition. Bishop Museum Press, Honolulu. 375 p.
Ueno, M. 1936. Cladocera of Mauna Kea, Hawaii. B. P. Bishop Mus. Occas. Pap. 12(11):
1?9.
Van Zwaluwenburg, R.H. 1948. Lacastris hawaiiensis Johnson. Proc. Hawaii. Entomol.
Soc. 13(2): 210.
V?vra, W. 1906. Ostracoden von Sumatra, Java, Siam, den Sandwich-Inseln und Japan.
Zool. Jahrb. (Syst.) 23: 413?38.
Versevelt, J. & F.M. Bayer. 1988. Revision of the genera Bellonella, Eleutherobia, Ni-
dallia and Nidaliopsis (Octocorallia: Alcyoniidae and Nidalliidae), with descriptions
of two new genera. Zool. Verh. 245: 1?131.
Vis, M.L., R.G. Sheath, J.A. Hambrook & K.M. Cole. 1994. Stream macroalgae of the
Hawaiian Islands: a preliminary study. Pac. Sci. 48: 175?87.
Vitousek, P.M., L.L. Loope & C.P. Stone. 1987. Introduced species in Hawaii: biolog-
ical effects and opportunities for ecological research. Trends Ecol. Evol. 2(7):
224?27.
Wagner, W.L., D.R. Herbst & S.H. Sohmer. 1990. Manual of the flowering plants of
Hawai?i. University of Hawaii Press & Bishop Museum Press, Honolulu. 1853 p.
Walsh, G.E. & R.L. Bowers. 1977. Order Zoanthiniaria, p. 148?57. In Devaney, D.M. &
L.G. Eldredge, eds., Reef and shore fauna of Hawaii. Section 1: Protozoa through 
Ctenophora. Bishop Museum Press, Honolulu.
Wells, J.B.J. 1986. Copepoda: marine-interstitial Harpacticoida, p. 356?81. In:
Botosaneanu, L., ed., Stygofauna mundi. E.J. Brill, Leiden.
Yamaguti, S. 1965. New digenetic trematodes from Hawaiian fishes, I. Pac. Sci. 19(4):
458?81.
???. 1968a. Cestode parasites of Hawaiian fishes. Pac. Sci. 22(1): 21?36.
???. 1968b. Monogenetic trematodes of Hawaiian fishes. University of Hawaii Press,
Honolulu. 287 p.
???. 1970. Digenetic trematodes of Hawaiian fishes. Keigaku Publishing Co., Tokyo.
436 p.
Young, R.E. 1991. Chiroteuthid and related paralarvae from Hawaiian waters. Bull. Mar.
Sci. 49(1?2): 162?85.
Yount, J.L. 1954. The taxonomy of the Salpidae (Tunicata) of the central Pacific Ocean.
Pac. Sci. 8(3): 276?330.
BISHOP MUSEUM OCCASIONAL PAPERS:  NO. 41, 199518
Electronic version via HBS web site: http://www.bishop.hawaii.org/bishop/HBS/ ?1995,1997 HBS