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Carlton & Eldredge — Marine Bioinvasions of Hawai‘i 33 As early as 1933, C.H. Edmondson speculated that some species of hydroids may have been transported to the islands in ship fouling (Edmondson, 1933). Cooke (1977a) remarked on the presence of many cosmopolitan species in the Hawaiian hydroid fauna, stating, “It should be noted ... that many of the forms reported were collected from artificial habitats such as floats, pilings, and so forth, or from disturbed areas such as Kāne‘ohe Bay. This would also tend to favor the establishment of colonizing (in the ecological, not morphological sense) species, which in general are quite widely distributed.” Cooke (1975) earlier noted, relative to the Enewetak hydroid biota, that the “many hundreds of ships and barges [that] visited Enewetak Atoll in the later part of World War II and during the period of atomic bomb testing” provided “a ready method ... for the introduction of so many cosmopolitan species.” While occasionally recognizing ships as a transport mechanism for harbor-dwelling hydroids, the hydrozoan systematic and biogeographic literature, in general, does not attempt to identify or suggest which species in subtropical and tropical harbor biotas, or in adjacent habitats in the open ocean, may be introduced, presumably in deference to the largely undocumented (and often unlikely) possibility of natural dispersal as oceanic drifters. We treat several species here as introduced, but most taxa are of necessity cryptogenic. Chu & Cutress (1955) reported Syncoryne mirabilis (Agassiz, 1862) (now known as Sarsia tubulosa [(M. Sars, 1835); Calder (1988)] as causing dermatitis in swimmers in Hawai‘i. There are no further reports of this or any similar hydroid from Hawai‘i, and D. Calder (pers. comm., July 2003) considers the report of Chu & Cutress to be a misidentification. We do not further consider it. As with a number of other groups, we note that many more species of shallow-water hydroids from the islands could be considered cryptogenic. Thus, the species listed here are examples of cryptogens, and our treatment is not meant to imply that we consider all other hydroids known from the islands to be native. Corydendriidae Cordylophora caspia (Pallas, 1771) Introduced = Cordylophora lacustris Allman, 1844 Cooke (1977a) reported this species in an anchialine (18 to 32 ‰) pond (Hālua Pond) at Cape Kinau, Maui, growing on the alga Caulerpa serrulata, collected by Diana Wong and John Maciolek. A Ponto-Caspian species native to the Black and Caspian Seas, Cordylophora has been widely distributed by ships in the Atlantic and Pacific Oceans. Wong (1975) notes the record; her material was collected in the summers of 1974 and 1975. Turritopsis nutricula McCrady, 1857 Introduced Cooke (1977a) found Turritopsis growing on the hydroid Halocordyle (= Pennaria) stems, “in harbors such as Ala Wai Yacht Harbor” in 1972 (BPBM-D456). Cooke further suggested that “it is likely that the medusa figured by Edmondson (1946a); [see Fig. 1b (sic, lapsus for 13b)] represents a maturing T. nutricula medusa.” Edmondson’s figure is of a medusa “occasionally taken in the tow net on Waikiki reef and in Pearl Harbor, Oahu” (Edmondson,1946a; the same figure and statement appears in Edmondson, 1933). However, Edmondson’s medusa is not Turritopsis nutricula (C. Mills, pers. comm., June 2003), and it may or may not be a campanulariid, as Edmondson suggested. Grovhoug & Rastetter (1980) reported T. nutricula from 1976–1977 collections in Pearl Harbor and Kāne‘ohe Bay. It still occurs in Kāne‘ohe Bay (Coles et al., 2002a) and doubtless still occurs in Pearl Harbor as well. This species, possibly North Atlantic in origin, now also occurs on the Pacific coast of America, and in South Africa, Papua New Guinea, and New Zealand (Schuchert, 1996), which we suggest is due to ship-borne transport. We thus also consider it introduced to Hawai‘i. It is often cited as being described in 1859 (e.g., Calder, 1988), but the date of publication was 1857 (D. Calder, pers. comm., June 2003).
34 BISHOP MUSEUM BULLETIN IN CULTURAL AND ENVIRONMENTAL STUDIES 4 (2009) Moerisiidae Moerisia horii (Uchida & Uchida, 1929) Cryptogenic =Ostroumovia horii Cooke (1977a) reported this species, as Ostroumovia horii, otherwise known from Japan, from a brackish pond in a lava flow near Honokohau, island of Hawai‘i (collected in 1972; W. Cooke, pers. comm., July 1996). He noted that the “recording of this species from both Japan and Hawaii is quite interesting, since most species in this family have a severely restricted distribution, unless transported by human activity.” We regard it as cryptogenic, pending further taxonomic work. No specimens from this collection in the Hawaiians Islands exist in the Bishop Museum collections, but Cooke (1977a, fig. 1) provides a figure of Hawaiian material. Bougainvilliidae Garveia sp. Introduced We regard an unidentified Garveia found in O‘ahu fouling communities as introduced by ships. Cooke (1977a) reported it on the hydroid Pennaria at Ala Wai Yacht Harbor, based upon collections in 1972 (W. Cooke, pers. comm., July 1996). Grovhoug & Rastetter (1980) reported it as “Garveia humilis (Allman, 1877)” from Pearl Harbor and Kāne‘ohe Bay based on 1976–1977 collections, an identification we regard as requiring confirmation. We predict that once identified, it will be found to be a species with a disjunct, harbor-based distribution. Bougainvillia muscus (Allman, 1863) Introduced = B. ramosa (van Beneden, 1844), a junior homonym; see Calder, 1988 and Schuchert, 1996). Tusov & Davis (1971) recorded a “Bougainvillia sp.” from Kāne‘ohe Bay, O‘ahu in the summer of 1967. Cooke (1977a) reported B. ramosa from Kāne‘ohe Bay in the form of dense colonies growing on the stems of the hydroid Pennaria and noted that it was “more common in temperate waters, but known to penetrate subtropically to Brazil and Australia.” Bishop Museum collections contain material from Kāne‘ohe Bay collected in October 1972 by T. Gosliner. We regard it as introduced by shipping, and, given the history and pattern of records of the species, as native to the Atlantic Ocean, and introduced to ports in the Pacific, such as in Australia, Papua New Guinea, and New Zealand (Schuchert, 1996). Eudendriidae Eudendrium sp. Cryptogenic Cooke (1977a) reported a Eudendrium species (lacking gonophores and thus not resolvable to species) from Honolulu Harbor and Kāne‘ohe Bay based upon material collected 1972 (W. Cooke, pers. comm., July 1996). We regard it as cryptogenic, as Eudendrium species are also frequently found in natural communities worldwide. Cladonematidae Cladonema radiatum Dujardin, 1843 Introduced This well-known North Atlantic hydroid (Calder, 1988) was first collected in the Pacific Ocean in 1949, when it was described as a new species from Wellington, New Zealand; it was later reported from Japan (Schuchert, 1996). W. Cooke collected it on O‘ahu in June, 1972 (BPBM-D461) who noted that “the medusae are often found on the alga Ulva in relatively quiet waters” (Cooke, 1977a). It has since been introduced to Puget Sound, Washington (Kozloff, 1996; Wrobel & Mills, 1998). Pennariidae Pennaria disticha Goldfuss, 1820 Introduced = Halocordyle disticha (following Gibbons & Ryland, 1989) = Pennaria tiarella Ayres, 1854 (following Vervoort, 1968) = Corydendrium splendidum Boone, 1938 (following Cooke, 1977a) Gibbons & Ryland (1989) have argued for the retention of the name Pennaria, with the genus Halocordyle as a junior synonym; Schuchert (1996) follows this usage. We regard this common foul-
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