and Numbers of Hawaiian Species: Supplement 1
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in several ways: 1) we expect that some literature has been missed; 2) we know that additional information, including unpublished new species, exist in museum collections; and 3) further fieldwork will produce additional records.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 parasites are also not included. Unless noted, all numbers represent described species reported 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 The numbers from the Reef and shore fauna manuscripts include previously unpublished records.
The term endemic is problematic. In the terrestrial environment it is easy to determine 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 especially 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 determine 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.
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 pattern 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 difficult 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.
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
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
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; Erséus & Davis, 1989;
Erséus, 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 361 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
Symphyla 5 0 5 Nishida, 1994
Palpigrada 1 0 1 Nishida, 1994
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 ? ? Vávra, 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 Alvariño, 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
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 21,388 8,759 4,267
* HBS Records for 1994 = articles in Bishop Museum Occasional Papers 41 and 42 of the Records of the Hawaii Biological Survey for 1994.
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 naturalized (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 additional 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), remaining are 1 species of bamboo coral (Muzik 1978) and 101 species of deep-water gorgonians (Grigg & Bayer 1976, Versevelt & Bayer 1988, 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 hermatypic, 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 nonindigenous, 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 (Erséus & Davis 1989); 1 new species by Erséus (1990); the other 2 are unidentified species (Bailey-Brock 1987).
Annelida: Polychaeta: Approximately 100 additional species are known, but not yet reported 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 elasmobranch (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. 1995), 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 (Vávra 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 gammarideans (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 (Alvariño 1978).
Echinodermata: Crinoidea: Of 16 species listed in the table, 11 described as new species by Clark (1908); no shallow-water forms.
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 visitor 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).
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 readings. 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 Sacromastigophora, Gnathostomula, Gastrotricha, Rotifera, and Tardigrada). Am. Mus. Novit. 3106, 44 p.
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.
---. & 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 parasitized 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 rhizostomes. 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. Backhuys Publishers, Leiden. vi + 248 p.
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 (Platyhelminthes: Digenea) parasitic in green turtles, Chelonia mydas (L.) from Hawaii. Bull. South. Calif. Acad. Sci. 91(2): 84-91.
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.
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Scott E. Miller and Lucius G. Eldredge (Department of Natural Sciences, Bishop Museum, 1525 Bernice Street, Honolulu, Hawaii 96817, USA)
This is a supplement to our earlier tabulation of species known from Hawaii (Eldredge & Miller 1995; see also Mlot 1995). The Hawaii Biological Survey is cur rently working on species checklists and bibliographies for many of these taxa, so this should be viewed as an interim report subject to further change. See Eldredge & Miller (1995) for definitions and scope. We have not included bacteria and viruses, but have now included human parasitic protists and helminths. Allison et al. (1995, table 21.3) summarized the numbers of marine species based on Eldredge & Miller (1995).
From literature and unpublished sources, approximately 22,056 species have been recorded from the Hawaiian Islands and surrounding waters. Of these, 8,850 are endemic to the Hawaiian Islands, and 4,464 are nonindigenous species of protists, fungi, plants and animals. These numbers include significant refinements from the previous compilation by Eldredge & Miller (1995), based on continued literature survey and review by specialists, especially among the parasitic Protozoa and helminths. Many gaps in knowledge of HawaiiÐs biota remain and many species of protists, algae, fungi, worms and arthropods remain to be described.
Table 1. Estimates of numbers of species of the Hawaiian biota (based on Eldredge & Miller 1995, as updated by this paper and other papers in this issue of Records of the Hawaii Biological Survey ).
Taxon Total Endemic NIS Algae and other protists 2228 4 5? Fungi and lichens 2023 240 0? Flowering plants 1910 850 877 Other plants 703 243 50 Mollusks 1656 956 75+ Insects 7862 5237 2527 Other arthropods 1795 335 519 Other invertebrates 2338 785 250 Fish 1195 139 73 Amphibians 4 0 4 Reptiles 24 0 20 Birds 274 60 46 Mammals 44 1 19 Total 22056 8850 4464Endemic species are restricted to Hawaii; nonindigenous (= alien, = exotic, includes introduced) species do not naturally occur in Hawaii; indigenous species occur naturally in Hawaii but are not endemic. See Eldredge & Miller 1995 for additional definitions and qualifications.
Freshwater and terrestrial Cyanophyta (blue-green algae)
We have not yet reconciled the many old records (e.g., MacCaughey 1918, Tilden 1901, 1910) with current nomenclature. Some recent records include Doty & Watson 1981, MacEntee & Bold 1978, and Zimmerman & Bergman 1990. Patterson et al. (1991 and included references) included many Hawaiian species in a phytochemical screening program. The recorded flora of freshwater and terrestrial forms likely approaches 100 species.
Freshwater and terrestrial algae
We have not yet reconciled the many old records (e.g., MacCaughey 1918, Tilden 1901) with current nomenclature, but our original estimate of over 120 species was too low. Recent authoritative records in clude over 172 diatoms (Fungladda et al. 1983), 26 desmids (Prescott et al. 1975-1983), and 34 macroalgae in streams (Vis et al. 1994). The total likely exceeds 250 species.
Protozoa
The protists of the Hawaiian islands have never been systematically sur veyed, but we have accumulated records of 207 species (excluding Foraminifera) from a variety of sources. Only authoritative identifications have been included. We expect that the ultimate number of species will be vastly greater.
Fungi
We have not yet tabulated the number of fungi beyond our original estimates, but it is worth pointing to the recent work on yeasts associated with flowering plants (Starmer 1981), especially Opuntia cactus (Starmer et al. 1990); the recent reviews of rust fungi (over 74 species ¦ Gardner & Hodges 1989, Gardner 1994); and Agaricales (101 taxa ¦ Desjardin et al. 1992).
Platyhelminthes (zooparasitic)
Of 583 species, 56 are cestodes (tapeworms) including 20 from marine fishes (Cornford 1974, Carvajal et al. 1976, Beveridge & Campbell 1993, Brill et al. 1987, Deardoff et al. 1984, Gulyaev & Korotaeva 1989, Yamaguti 1968a), 1 from freshwater fishes (Font & Tate 1994), 21 from terrestrial vertebrates (including human) (Alicata 1969, Olsen & Haas 1976, Lewis & Holmes 1971), of 2 species from intermingling mammals 1 is new record (McKenzie & Davidson), 7 from green turtles (Dailey et al. 1992), 1 from porpoise (Testa & Dailey 1977), 4 from Hawaiian monk seal (Anderson 1987), and 1 from "megamouth" shark (Dailey & Vogelhein 1982); the remaining 527 species are trematodes (flukes), 507 from marine fishes (Lebedev 1984, Mamaev 1989, Oliver 1983, Yamaguti 1965, 1968b, 1970), 16 from terrestrial vertebrates (including human) (Alicata 1969, O'Leary et al. 1985), 1 unidentified species (larva) from freshwater fish (Font & Tate 1994), 1 from nonindigenous, marine, cage-cultured tilapia (Kaneko et al. 1988), 1 species from the cane toad (Speare 1990), 1 species with secondary intermediate stage in coral Porites compressa (Aeby 1991).
Acanthocephala
Three species not reported in 1995: Moniliformis moniliformis from rats (Alicata 1969) and cockroach (Schaefer 1970); Corynosoma rauschi from Hawaiian monk seal (Dailey et al. 1988); and Mediorhynchus orientalis from birds (Schmidt & Kuntz 1977); also unidentified species from freshwater fish (Font & Tate 1994).
Nematoda (zooparasitic)
Of 131 zooparasitic species, 93 reported from terrestrial vertebrates (including man) (Alicata 1969), of 22 species reported from inter mingling mammals 12 are new records (McKenzie & Davidson 1989), 1 species from Hawaiian monk seal (Dailey 1988), 1 from albatross (Langston & Hillgarth 1995), 3 from endemic birds (Cid del Prado et al. 1985), 3 from gamebirds (Lewin & Holmes 1971, Smith 1973), 18 from marine fishes (Bruce et al. 1994, Deardoff & Overstreet 1982, Deardoff et al. 1982, 1983, Deardoff 1987, Noble 1966, Solovjeva & Pozdnyakov 1984), and 1 from freshwater fish (Font & Tate 1994).
Tardigrada Pollock (1995) published the first record of 2 species of marine tardigrades--Dipodarctus borrori and D. anaholiensis--from Anahola Bay, Kauai.
Bryozoa
Two Hawaiian freshwater bryozoan species were overlooked in our earlier report (Bailey-Brock & Hayward 1984); one new deepwater, marine species described (Chimonides & Cook 1994).
Mollusca
Cowie et al. (1995) previously cited as “in pressÝ has been published; the 75 non-indigenous species in the table represents non-marine mollusks only¦ numbers of non-indigenous marine mollusks have not yet been ascertained (R.H. Cowie, pers. comm.).
Annelida: Hirudinea
Two leeches collected in association with freshwater fishes to be added to species list: Myzobdella lugubris and Cystobranchus sp. (Font & Tate 1994).
Arthropoda: Crustacea
Recent additions to the crustacean fauna of the Hawaiian Islands include: cyclopoid copepod Haplostomides hawaiiensis [n.sp.](Ooishi 1994); harpacticoid copepod Psammopsyllus stri (Kunz 1993); ascothoracid cirriped Laura bicornuta [n.sp.] and Zoanthoecus cerebroides [n.sp.](Grygier 1985); mysids Anisomysis hawaiiensis [n.sp.], A. extranea [n.sp.], A. xenops (Murano 1995), Holmesimysis costata (Holmquist 1979), and Gnathophausia longispina (Wilson & Boehlert 1993); marine isopods Neonaesa rugosa and Cymodocella hawaiiensis [n.sp.](Bruce 1994); deep-water caridean shrimp Opaepele loihi [n.gen. & n.sp.] (Williams & Dobbs 1995).
Terrestrial and freshwater Arthropoda
The numbers for some groups have changed because of continued refinement of synonymies and distribution status in the database (G.M. Nishida, pers. comm.), as well as recent publications (Gillespie 1994¦3 new endemic spiders; Asquith 1994¦39 new endemic bugs). Many additional endemic species od arthropods remain to be described (e.g., Howarth 1990). Also, the original figure for nonindigenous species of the insect order Diptera was a typographical error and should read 361 rather than 631 (this did not impact the total numbers).
Chordata: Reptilia
Our original list omitted 3 indigenous species of sea turtles and 3 nonindigenous species of freshwater turtles (C. Kishinami, pers. comm.).
Acknowledgements
Our literature survey was improved by access to the CAB Abstracts and AGRICOLA databases (CAB International and National Agriculture Library, U.S. Dept. of Agriculture, respectively). We thank the libraries of Bishop Museum, Santa Barbara Museum of Natural History, University of California at Santa Barbara, and University of Hawaii at Manoa for access to literature. Various specialists have as sisted in literature search and analysis, or submitted reprints. The specimen database for the U.S. National Animal Parasite Collection, U.S. Department of Agriculture, Beltsville, Maryland (E.P. Hoberg & P.A. Pilitt) also helped our literature search. This is an interim product from a project supported by the John D. and Catherine T. MacArthur Foundation and National Biological Service.
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