The How Many Species Are there in Hawaii Web Page

Featuring: "How many species are there in Hawaii?" (below) From the "Records of the Hawaii Biological Survey for 1994" as published in the Bishop Museum Occasional Paper volume 41: 3-18.

and Numbers of Hawaiian Species: Supplement 1


How Many Species Are there in Hawaii?


Lucius G. Eldredge and Scott E. Miller (Department of Natural Sciences, Bishop Museum, P.O. Box 19000A, Honolulu, Hawai`i 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 physiographic 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 invertebrates 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in several ways: 1) we expect that some literature has been missed; 2) we know that additional information, including unpublished new species, exist in museum collections; and 3) further fieldwork will produce additional records.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.

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 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.

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
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.

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 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).


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 endemic 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 freshwater, 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, bacteria, protists, freshwater algae, etc.), still remain.


Acknowledgements

The authors thank the following people and institutions for their assistance and support 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. 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 (International 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. Norenberg 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

Alvariño, 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 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.

---. 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.

Erséus, C. 1990. Marine Oligochaeta of Hong Kong, p. 259-335. In: Morton, B., ed., 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 zoogeography 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 scientific 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. Helgoländ. 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 descriptions 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.

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 dorotocephala (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 invertebrates. 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 sediments. 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. 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 collections 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 (Coelenterata: 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 Onuphidae (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 phylum Placozoa in Hawaii. Pac. Sci. 43(2): 117-21.

Pettibone, M.H. 1986. Review of the Iphioninae (Polychaeta: Polynoidae) and revision of Iphione cimex Quatrefages, Gattyana deludens Fauvel, and Harmothoe iphionelloides 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 chaetognaths. 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 microorganisms, 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, University 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 Commission 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. Cephalopods 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 Association 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 (Tunicata, Thaliacea), with descriptions of two new species. Beaufortia 22(288): 17-55.

---. 1974b. A revision of the genera Salpa Forskål, 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 (Ectoprocta), 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 Cladoniaceae. 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.

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. Second 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.

Vávra, 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, Nidallia 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: biological 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.


Numbers of Hawaiian Species: Supplement 1


[Originally published in Bishop Museum Occasional Papers 45: 8-17 (1996).]

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 currently 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 HawaiiĞs 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       4464

Endemic 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 include 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 surveyed, 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.

Free-living marine protozoa: 39 species (Phillips 1977, Larsen & Patterson 1990), in addition to over 1000 species of Forminifera (Chave 1987).
Free-living freshwater protozoa: 61 species, including 59 testaceous rhizopods (Van Winkle 1927), Paramecium primaurelia Sonneborn 1975, and Tetrahymena tropicalis Nanney & McCoy 1976 (= T. pyriformis variety 9 of Elliott et al. 1964).
Free-living soil protozoa: At least 31 ciliates, 9 flagellates, 6 Sarcodina, and 10 Testacea recorded from various habitats on Hawaii Island (Bamforth & Eggler 1973, Foissner 1993, 1994).
Parasitic protozoa of terrestrial vertebrates including humans: 60 species. Alicata (1969) detailed the species known from humans and domestic animals. Other key references include: Colgrove (1976 ¦ dog), Metcalf (1923 ¦ amphibian), Upton et al. (1994 ¦ reptiles), Wallace & Frenkel (1975 ¦ cat), Van Hoven et al. (1992 ¦ deer), and Van Riper (1991 ¦ birds).
Parasitic protozoa of invertebrates: 11 species symbionts in 3 species of termites (Kirby 1941-1946, Lai et al. 1983), at least 9 species from insects (Fujii & Tamashiro 1967, Tamashiro 1968, Tanabe & Tamashiro 1967, Tanada 1957), 2 species from marine crabs (Ball 1963).
Parasitic protozoa of marine fishes: 8 species (Basson et al. 1990, Lom 1970, Moser & Noble 1977a, b).

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 intermingling 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 pressİ 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).

Insecta: 7862 total species, including 5237 endemic and 2527 nonindigenous;
Araneae: 200 total species, including 126 endemic and 70 nonindigenous;
Acari: 529 total species, including 103 endemic and 365 nonidigenous.
Symphyla: Has 5 nonindigenous species (Nishida 1994), was overlooked in the original list.

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 assisted 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.

Literature Cited

Aeby, G.S. 1991. Behavioral and ecological relationships of a parasite and its hosts within a coral reef system. Pac. Sci. 45: 263-69.

Alicata, J.E. 1969. Parasites of man and animals in Hawaii. S. Karger, Basel. 190 p.

Allison, A., S.E. Miller & G.M. Nishida. 1995. Hawaii Biological Survey: a model for the Pacific Region, p. 349-55 In Maragos, J.E., M.N.A. Peterson, L.G. Eldredge, J.E. Bardach & H.F. Takeuchi, eds., Marine and coastal biodiversity in the tropical island Pacific region. Volume 1. Species systematics and information management priorities. East-West Center, Honolulu. xxii + 424 p.

Anderson, K.I. 1987. A redescription of Diphyllobothrium stemmacephalum Cobbold, 1858 with comments on other marine species of Diphyllobothrium Cobbold, 1958. J. Nat. Hist. 21: 411-27.

Asquith, A. 1994. Revision of the endemic Hawaiian genus Sarona Kirkaldy (Heteroptera: Miridae: Orthotylinae). Bishop Mus. Occas. Pap. 40: 1-81.

Bailey-Brock, J.H. & P.J. Hayward. 1984. A freshwater bryozoan, Hyalinella vaihiriae Hastings (1929), from Hawaiian prawn farms. Pac. Sci. 38: 199-204.

Ball, G.H. 1963. Cephaloidophora carpilodei n. sp. and C. pinguis n. sp., gregarines of a xanthid crab in Hawaii. J. Protozool. 10: 321-27.

Bamforth, S.S. & W.A. Eggler. 1973. Protozoa from pumice soils of Kilauea Iki Volcano, Hawaii. J. Protozool. 20: 496. [abstract]

Basson, L., J.G. Van As & L. Fishelson. 1990. A new species of Trichodina (Ciliophora: Peritrichia) from the intestine of the surgeonfish Acanthurus xanthopterus. Int. J. Parasitol. 20: 785-87.

Beveridge, I. & R. A. Campbell. 1993. A revision of Dasyrhynchus Pinter (Cestoda: Trypanorhyncha), parasitic in elasmobranch and teleost fishes. Syst. Parasitol. 24: 129-57.

Brill, R.W., R. Bourke, J. A. Brock & M. D. Dailey. 1987. Prevalence and effects of infection of the dorsal aorta in yellowfin tuna, Thunnus albacares, by the larval cestode, Dasyrhynchus talismani. Fish. Bull. 85: 767-76.

Bruce, N.L. 1994. New records of marine isopod crustaceans (Sphaeromatidae) from Hawaiian waters. Crustaceana 67: 381-86.

-----, R.D. Adlard & L.R.G. Cannon. 1994. Synoptic checklist of ascaridoid parasites (Nematoda) from fish hosts. Invertebr. Taxon. 8: 583-674.

Carvajal, J., R.A. Campbell & E.M. Cornford. 1976. Some tyrypanorhynch cestodes from Hawaiian fishes with descriptions fo new species. J. Parasitol. 62: 70-77.

Chimonides, P.J. & P.L. Cook. 1994. Notes on the genus Cranosina (Bryozoa, Cheilostomida). Zool. Scripta 23: 43-49.

Cid del Prado, Vera, I., A.R. Maggenti & C. van Riper, III. 1985. New species of Spiruridae (Nemata: Spirurida) from endemic Hawaiian honeycreepers (Passeriformes: Drepanididae), the Japanese white-eye (Passeriformes: Zosteropidae) and a new species of Acuariidae (Nemata: Spirurida) from the Japanese white-eye collected on the island of Hawaii. Proc. Helminthol. Soc. Wash. 52: 247-59.

Colgrove, D.J. 1976. Canine babesiosis in Hawaii. J. Amer. Vet. Med. Assoc. 168: 591-92.

Cornford, E.M. 1974. Two tetraphyllidean cestodes from Hawaiian stingrays. J. Parasitol. 60: 942-48.

Cowie, R.H., N.L. Evenhuis & C.C. Christensen. 1995. Catalog of the native land and freshwater molluscs of the Hawaiian Islands. Backhuys Publ., Leiden. vi + 248 p.

Dailey, M.D. & W. Vogelbein. 1982. Mixodigmatidae, a new family of cestode (Trypanorhyncha) from a deep sea, planktivorous shark. J. Parasitol. 68: 145-49.

-----, 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: 84-91.

-----, R.V. Santangelo & W. G. Gilmartin. 1988. A coprological survey of helminth parasites of the Hawaiian monk seal from the northwestern Hawaiian Islands. Mar. Mamm. Sci. 4: 125-31.

Deardoff, 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: 28-39.

----- & R.M. Overstreet. 1982. Hysterothylacium pelagicum sp. n. and H. cornutum (Stossich, 1904) (Nematoda: Anisakidae) from marine fishes. Proc. Helminthol. Soc. Wash. 49: 246-51.

-----, 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: 187-201.

-----, R.B. Raybourne & T.E. Mattis. 1984. Infections with trypanorhynch plerocerci (Cestoda) in Hawaiian fishes of commercial importance. Sea Grant Q. 6: 1-6.

Desjardin, D.E., G.J. Wong & D.E. Hemmes. 1992. Agaricales of the Hawaiian Islands. I. Marasmioid fungi: new species, new distributional records, and poorly known taxa. Can. J. Bot. 70: 530-42.

Doty, M.S. & L.-L. Watson. 1981. Terrestrial algae, p. 165-71. In: Mueller-Dombois, D., K.W. Bridges & H.L. Carson, eds., Island ecosystems: biological organization in selected Hawaiian communities. Hutchinson Ross Publ. Co., Stroudsburg, Penn.

Eldredge, L.G. & S.E. Miller. 1995. How many species are there in Hawaii? Bishop Mus. Occas. Pap. 41: 3-18.

Elliott, A.M., M.A. Studier & J.A. Work. 1964. Tetrahymena pyriformis from several Pacific islands and Australia. J. Protozool. 11: 370-78.

Foissner, W. 1993. Corticocolpoda kaneshiroae n. g., n. sp., a new colpodid ciliate (Protozoa, Ciliophora) from the bark of ohia trees in Hawaii. J. Euk. Microbiol. 40: 764-75.

-----. 1994. Bryometopus hawaiiensis sp. n., a new colpodid ciliate from a terrestrial biotope of the Hawaiian archipelago (Protozoa: Ciliophora). Ann. Naturhist. Mus. Wien (B) 96: 19-27.

Font, W.F. & D.C. Tate. 1994. Helminth parasites of native Hawaiian freshwater fishes: an example of extreme ecological isolation. J. Parasitol. 80: 682-88.

Fujii, J.K. & M. Tamashiro. 1972. Nosema tephrititae sp. n., a microsporidian pathogen of the Oriental Fruit Fly, Dacus dorsalis Hendel. Proc. Hawaii. Entomol. Soc. 21: 191-203.

Fungladda, N., I. Kaczmarska & S.R. Rushforth. 1983. A contribution to the freshwater diatom flora of the Hawaiian islands. Biblioth. Diatomol. 2: 1-103.

Gardner, D.E. 1994. The native rust fungi of Hawaii. Can. J. Bot. 72: 976-89.

----- & C.S. Hodges, Jr. 1989. The rust fungi (Uredinales) of Hawaii. Pac. Sci. 43: 41-55.

Gillespie, R.G. 1994. Hawaiian spiders of the genus Tetragnatha: III. Tetragnatha acuta clade. J. Arachnol. 22: 161-68.

Grygier, M.J. 1985. Lauridae: taxonomy and morphology of ascothoracid crustacean parasites of zoanthids. Bull. Mar. Sci. 36: 278-303.

Gulyaev, V.D. & V.D. Korotaeva. 1989. [Redescription of the type species of Glossobothrium Yamaguti, 1952 (Cestoda, Pseudophyllidea.] Vestn. Zool. 1989(6): 8-11. [in Russian]

Holmquist, C. 1979. Mysis costata Holmes, 1900, and its relations (Crustacea, Mysidacea). Zool. Jahrb. (Syst.) 106: 471-99.

Howarth, F.G. 1990. Hawaiian terrestrial arthropods: An overview. Bishop Mus. Occas. Pap. 30: 4-26.

Kaneko, J.J., II, R. Yamada, J.A. Brock & R.M. Nakamura. 1988. Infection of tilapia, Oreochromis mossambicus (Trewavas), by a marine monogenean, Neobenedenia melleni (MacCallum, 1927), 1963 in Kaneohe Bay, Hawaii, USA, and its treatment. J. Fish Diseases 11: 295-300.

Kirby, H. 1941-1946. Devescovinid flagellates of termites. Univ. Calif. Publ. Zool. 45: 1-318.

Kunz, H. 1993. Beitrag zur kenntnis von zwei Psammopsyllus-arten (Copepoda, Harpacticoida, Cylindropsyllidae). Crustaceana 64: 143-54.

Lai, P.Y., M. Tamashiro & J.K. Fujii. 1983. Abundance and distribution of the three species of symbiotic Protozoa in the hindgut of Coptotermes formosanus (Isoptera: Rhinotermitidae). Proc. Hawaii. Entomol. Soc. 24: 271-76.

Langston, N. & N. Hilgarth. 1995. Moult varies with parasites in Laysan Albatross. Proc. R. Soc. Lond. (B) 261: 239-43.

Lebedev, B.I. 1984. [New taxa of gastrocotylid-like monogenans.] Parazity Zhivotnykh i Rastenii [Parasites of animals and plants], p. 17-24. [In Russian with English summary.]

Lewin, V. & J.C. Holmes. 1971. Helminths from exotic game birds of the Puuwaawaa Ranch, Hawaii. Pac. Sci. 25: 372-81.

Lom, J. 1970. Trichodinid ciliates (Peritrichida: Urceolariidae) from some marine fishes. Folia Parasitologica 17: 113-25.

MacCaughey, V. 1918. Algae of the Hawaiian Archipelago. Bot. Gaz. 65: 42-57, 121-49.

MacEntee, F.J. & H.C. Bold. 1978. Some microalgae from sand. Texas J. Sci. 30: 167-73.

Mamaev, Y. 1989. [The species compositon and morphological characteristics of the genus Microcotyle (Microcotylidae, Monogenoidea)], p. 32-38 In: Lebedev, B.I., ed., Parazitologicheskie Issledovaniya [Investigations in parasitology]. [In Russian with English summary.]

Margulis, L., J.O. Corliss, M. Melkonian & D.J. Chapman, eds. 1990. Handbook of Protoctista. Jones and Bartlett, Boston. xli + 914 p.

McKenzie, M.E. & W.R. Davidson. 1989. Helminth parasites of interminglind axis deer, wild swine and domestic cattle from the island of Molokai, Hawaii. J. Wildlife Diseases 25: 252-57.

Metcalf, M.M. 1923. The opalinid ciliate infusorians. U.S. Natl. Mus. Bull. 120: 1-484.

Mlot, C. 1995. Biological surveys in Hawaii, taking inventory of a biological hot spot. Science 269: 322-23.

Moser, M. & E.R. Noble. 1977a. Three genera of Myxosporida (Protozoa) in macrourid fishes. Int. J. Parasitol. 7: 93-96.

----- & E.R. Noble. 1977b. Zschokkella (Protozoa: Myxosporida) in macrourid fishes. Int. J. Parasitol. 7: 97-100.

Murano, M. 1995. New and already known species of the genus Anisomysis (Mysidacea) from Hawaii and the Society Islands. J. Crust. Biol. 15: 355-64.

Nanney, D.L. & J.W. McCoy. 1976. Characterization of the species of the Tetrahymena pyriformis complex. Trans. Am. Microscop. Soc. 95: 664-82.

Nishida, G.M., ed. 1994. Hawaiian terrestrial arthropod checklist. Second edition. Bishop Mus. Tech. Rep. 4: 1-287.

Noble, E. R. 1966. A new camallanid nematode from Hawaii. Pac. Sci. 20: 360-66.

O'Leary, M.J., J.T. Berthiaume & V. Sakbun. 1985. Treatment of Clonorchis sinensis in Hawaii's Laotian population--experience with praziquantel. Hawaii Med. J. 44: 63-64.

Oliver, G. 1983. Diplectanum yamaguti sp. n. (Monogenea, Monopisthocotylea, Diplectanidae), parasite de Kyphosus cinerascens (Forsskål) û Hawaii. Zool. Scripta 12: 91-93.

Olsen, O.W. & W.R. Haas. 1976. A new record of Spirometra mansoni, a zoonotic tapeworm, from naturally infected cats and dogs in Hawaii. Hawaii Med. J. 35: 261-63.

Ooishi, S. 1994. Haplostomides hawaiiensis, new species (Copepoda: Cyclopoida: Ascidicolidae), associated with the ascidian Polyclinum constellatum at Honolulu, Hawaii. Hydrobiologia 292/293: 89-96.

Patterson, G.M.L., C.L. Baldwin, C.M. Bolis, F.R. Caplan, H. Karuso, L.K. Larsen, I.A. Levine, R.E. Moore, C.S. Nelson, K.D. Tschappat, G.D. Tuang, E. Furusawa, S. Furusawa, T.R. Norton & R.B. Raybourne. 1991. Antineoplastic activity of cultured blue-green algae (Cyanophyta). J. Phycol. 27: 530-36.

Pollock, L.W. 1995. New marine tardigrades from Hawaiian beach sand and phylogeny of the family Halechiniscidae. Invert. Biol. 114: 220-35.

Prescott, G.W. et al. 1975-1983. A synopsis of North American desmids. Part II. Desmidiaceae: Placodermae. Sections 1-5. University of Nebraska Press, Lincoln. [coauthors vary]

Schaefer, P.W. 1970. Periplanta americana (L.) as intermediate host of Moniliformis moniliformis (Bremser) in Honolulu, Hawaii. Proc. Helminthol. Soc. Wash. 37: 204-06.

Schmidt, G.D. & R.E. Kuntz. 1977. Revision of Mediorhynchus Van Cleave 1916 (Acanthocephala) with a key to species. J. Parasitol. 63: 500-07.

Schottelius, J. 1989. Contribution to the characterization of South American Trypanosomatidae: I. The importance of lectins, neuraminic acid and neuraminidase for the differentiation of trypanosomes and leishmania from the New World. Zool. Anz. 223: 67-81.

Smith, H.E. 1973. Syngamus trachea: a first record in Hawaii. Elepaio 33(9): 95-96.

Solovjeva, G.F. & S.E. Pozdnyakov. 1984. [A new nematode species, Hysterothylacium hospitum n. sp. (Nematoda, Anisakidae), found in the spiral valve of a blue shark.] Parazitologia 18(1): 66-68. [In Russian with English summary.]

Sonneborn, T.M. 1975. The Paramecium aurelia complex of fourteen sibling species. Trans. Am. Microscop. Soc. 94: 155-78.

Speare, R. 1990. A review of the diseases of the cane toad, Bufo marinus, with comments on biological control. Aust. Wildl. Res. 17: 387-410.

Starmer, W.T. 1981. A comparison of Drosophila habitats according to the physiological attributes of the associated yeast communities. Evolution 35: 38-52.

-----, M.-A. Lachance, H.J. Phaff & W.B. Heed. 1990. The biogeography of yeasts associated with decaying cactus tissue in North America, the Caribbean, and northern Venezuela. Evol. Biol. 24: 253-96.

Tamashiro, M. 1968. Microbial control in Hawaii. Proc. Hawaii. Entomol. Soc. 20: 157-67.

Tanabe, A.M. & M. Tamashiro. 1967. The biology and pathogenicity of a microsporidian (Nosema trichoplusiae sp. n.) of the cabbage looper, Trichoplusia ni (Hübner) (Lepidoptera: Noctuidae). J. Invert. Pathol. 9: 188-95.

Tanada, Y. 1957. An annotated list of infectious diseases of insects in Hawaii. Proc. Eighth Pac. Sci. Cong. 3A: 1235-1250.

Testa, J. & M.D. Dailey. 1977. Five new morphotypes of Phyllobothrium delphini (Cestoda: Tetraphyllidea), their relationship to existing morphotypes, and their zoogeography. Bull. South. Calif. Acad. Sci. 76: 99-110.

Tilden, J.E. 1901. Collection of algae from the Hawaiian Islands. Hawaii. Ann. 1902: 106-13.

-----. 1910. Minnesota algae. Volume I. The Myxophyceae of North America and adjacent regions including Central America, Greenland, Bermuda, the West Indies and Hawaii. University of Minnesota, Minneapolis. iv + 328 p.

Upton, S.J., A.A. Marchiondo & R.N. Williams. 1988. New species of Isospora (Apicomplexa: Eimeriidae) from passeriform birds of Hawaii. Syst. Parasitol. 12: 81-85.

-----, K. Hanley & T.J. Case. 1994. Eimeria frenatus n.sp. and Eimeria rochalimai (Apicomplexa: Eimeriidae) from Hemidactylus frenatus (Sauria: Gekkonidae) in Hawaii, U.S.A. Trans. Am. Microscop. Soc. 113: 390-94.

Van Hoven, W., F.M.C. Gilchrist & T.C. Telfer. 1992. A comparison of rumen ciliates in black-tailed deer from California and Hawaii. Calif. Fish Game 78: 124-27.

Van Riper, C., III. 1991. The impact of introduced vectors and avian malaria on insular passeriform bird populations in Hawaii. Bull. Soc. Vector Ecol. 16: 59-83.

Van Winkle, M.E. 1927. Freshwater testaceous rhizopods of Hawaii. Unpubl. M.S. thesis, University of Hawaii, Honolulu. 41 p.

Vis, M.L., R.G. Sheath, J.A. Hambrook & K.M. Cole. 1994. Stream macroalgae of the Hawaiian Islands: a preliminary survey. Pac. Sci. 48: 175-87.

Wallace, G.D. & J.K. Frenkel. 1975. Besnoitia species (Protozoa, Sporozoa, Toxoplasmatidae): recognition of cyclic transmission by cats. Science 188: 369-71.

Williams A.B. & F.C. Dobbs. 1995. A new genus and species of caridean shrimp (Crustacea: Decapods: Bresiliidae) from hydrothermal vents on Loihi Seamount, Hawaii. Proc. Biol. Soc. Wash. 108: 228-37.

Wilson, C.D. & G.W. Boehlert. 1993. Population biology of Gnathophausia longispina (Mysidacea: Lophogastrida) from a central North Pacific seamount. Mar. Biol. 115: 537-43.

Yamaguti, S. 1965. New digenetic trematodes from Hawaiian fishes, I. Pac. Sci. 19: 458-81.

-----. 1968a. Cestode parasites of Hawaiian fishes. Pac. Sci. 22: 21-36.

-----. 1968b. Monogenetic trematodes of Hawaiian fishes. Unviersity of Hawaii Press. 287 p.

-----. 1970. Digenetic trematodes of Hawaiian fishes. Keigaku Publishing Co., Tokyo. 436 p.

Zimmerman, W.J. & B. Bergman. 1990. The Gunnera symbiosis: DNA restriction fragment length polymorphism and protein comparisons of Nostoc symbionts. Microbial Ecol. 19: 291-302.


Go Back to HBS Home Page

Bishop Museum Home Page


These pages are being maintained by Neal L. Evenhuis

and were last modified on 10 January 2002