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Siphonophorae (Cnidaria) of the Gulf of Mexico, Pp Pugh, P. R. and R. Gasca. 2009. Siphonophorae (Cnidaria) of the Gulf of Mexico, Pp. 395–402 in Felder, D.L. and D.K. Camp (eds.), Gulf of Mexico–Origins, Waters, and Biota. Biodiversity. Texas A&M University Press, College Station, Texas. •20 Siphonophorae (Cnidaria) of the Gulf of Mexico Philip R. Pugh and Rebeca Gasca The siphonophorae are a subclass of the class Hydro- zoa, phylum Cnidaria. They are highly polymorphic and complex animals, and their zooids, both medusoid and polypoid, have become specialized to carry out various functions, such as swimming, feeding, reproduction, and protection. Although phylogenetically they originated as colonies, structurally, physiologically, and embryologi- cally they qualify as an organism (Wilson 1975), and they are sometimes referred to as “superorganisms” (Mackie 1963). They are exclusively marine, and the vast majority of species are pelagic. The only exceptions are a family of physonect siphonophores, the Rhodaliidae, that use their tentacles to attach themselves to the bottom substrate, and the only well- known species, the Portuguese Man o’ War Physalia physalis, which is pleustonic, floating at the sur- face with its tentacles hanging down many tens of meters into the water below. Siphonophores occur in a bewil- Siphonophora. After Brusca and Brusca 2003, modified by dering array of shapes and sizes, and can vary in length F. Moretzsohn. from a few millimeters to several tens of meters; some are undoubtedly the longest animals in the world (Robison asexual swimming bells (nectophores), arranged in a 2004). They can be found throughout the world’s oceans, biserial or spiral series immediately below the float, are and throughout the water column down to a depth of at present (Physonectae) or absent (Cystonectae). How- least 4500 m, although individual species are restricted in ever, recent molecular studies (Dunn, Pugh, and Had- their geographical and depth distributions to a lesser or dock 2005) have shown that although the Cystonectae are greater extent. monophyletic and sister to all other siphonophores, the Classically, siphonophores have been split into 3 Physonectae are paraphyletic and gave rise to the Caly- orders, based first on the presence (Cystonectae and Phy- cophorae. Dunn, Pugh, and Haddock (2005) suggested sonectae) or absence (Calycophorae) of an apical, gas- that these 2 latter groups should be collectively called the filled float (pneumatophore). The orders whose species Codonophora. The consequence of this study is that the possess a pneumatophore are split according to whether traditional taxonomy of the Physonectae will have to be 395 396 ~ Siphonophorae (Cnidaria) substantially altered, particularly with regard to the fami- is largely a reflection of sampling effort, and one would lies Agalmatidae and Athorybiidae. However, because we expect to find all the species throughout the area. Most of do not yet know the phylogenetic relations of all the phy- the 13 species found exclusively in the SE quadrant were sonect genera, for the present review the classical system- recently collected by the submersible Johnson- Sea- Link, atic arrangement, as summarized by Totton (1965), will be and the only quadrant where net sampling took place at maintained, with the addition of the recently established depths greater than 500 m was the SW one, where 9 spe- family Erennidae (Pugh 2001). cies were exclusively found. The best text for the systematics and morphology of siphonophores remains A Synopsis of the Siphonophora Abbreviations by Totton (1965), although 7 of the species presently recorded in the Gulf of Mexico have been described since In the checklist, the families, and the genera and species then. Other useful, well- illustrated texts on the siphono- within each, are listed in alphabetical order for each of phores of certain areas are Bouillon et al. (2004, Mediter- the orders—Cystonectae, Physonectae, and Calycopho- ranean); Gasca (1990b, Gulf of Mexico); Kirkpatrick and rae, respectively. Under the heading of Habitat- Biology, Pugh (1984, British Isles); Pagès and Gili (1992, South each species is considered to be either neritic (ner) or West Africa); and Pugh (1999a, South Atlantic); the last deep- sea (dps), and, for the pelagic species, their depth includes a key to the species occurring in that region, with distribution is categorized as epipelagic (epp = 0–ca. 300 illustrations of all but 13 of the species now known to be m), mesopelagic (mep = ca. 300–1000 m), and / or bathy- present in the Gulf of Mexico. Mackie, Pugh, and Purcell pelagic (bap = >1000 m). The latter categories refer the (1987) have summarized our knowledge regarding their usual depth ranges where the species occur (Pugh, unpub- biology, evolution, distribution, and ecology, and Carré lished data), which may not be reflected in the records and Carré (1995) have considered their anatomy and from the Gulf of Mexico itself. One species is pleustonic reproduction. (ple). Under the heading Overall Geographic Range, the Phillips (1972) considered that, for the original list geographical distribution of a species is considered in 3 of siphonophores from the Gulf of Mexico, Sears (1954) ways. First, whether the species has a widespread distribu- “presents a hypothetical list of siphonophores, none based tion in the world’s oceans (world) or is restricted to either on actual records.” In actuality this is not the case, as her the Atlantic (Atl), Indian (Ind), or Pacific (Pac) Oceans, or list of 24 species appears to be almost exclusively based to the Mediterranean (Med) Sea. For rarer species, where on Bigelow (1918), although some of the species Bigelow there is no clear- cut pattern of distribution, they are con- listed were actually caught in the western Atlantic and not sidered to have sporadic occurrences. Second, within in the Gulf of Mexico. Nonetheless, there were some spe- these distributional zones, the waters where the species cies that had previously been found in the Gulf of Mexico is most abundant, when known, are considered broadly that Sears did not include. In recent years, there have been as either tropical (trop), subtropical (subtrop), temperate many studies on the siphonophores of the Gulf of Mexico, (temp), or boreal. It should be recognized, however, that as summarized by Gasca (2002). The vast majority of these these are very coarse categories and that their boundaries have reported on net collections in the top 200–500 m of vary spatially. Third, the relative abundance of each spe- the water column, and there are very few records from cies is indicated by the categories: v. common, common, greater depths (Stepanjants 1975). However, judging by uncommon, rare, and v. rare. the number of species new to the Gulf of Mexico that have The specimens from which the new records for the Gulf been collected recently by submersibles (e.g., Pugh 2001), of Mexico are derived are currently housed in the National there is probably still a large number to add to the present Oceanography Centre Collections (NOCC). The numbers list of 82. This number represents almost half of the total refer to the Johnson- Sea- Link dives during which the spec- number of about 175 siphonophore species that are cur- imens were collected. At some time in the future, once the rently recognized as valid. None of the species found in specimens have been fully examined and described, it is the Gulf are endemic to the area. intended to transfer them to the U.S. National Museum of Although only 24 siphonophore species were found Natural History (Smithsonian Institution). in all 4 quadrants of the Gulf of Mexico, with the largest Abbreviations used in the Habitat- Biology column of number of species being found in the 2 southern ones, this the checklist include bap = bathypelagic; dps = deep sea; Pugh and Gasca ~ 397 epp = epibenthic / planktonic; mep = mesopelagic; ner = (Cnidaria: Hydrozoa) en la region sur del Golfo de México. neritic; ple = pleustonic; plg = pelagic. Caribbean Journal of Science 29: 220–225. 13. Gasca, R. 1998. Siphonophore communities in the southern Gulf of Mexico during April–May, 1986. References Proceedings of the Second International Congress on 1. Alvariño, A. 1972. Zooplancton del Caribe, Golfo de Pelagic Biogeography. IOC / UNESCO Workshop Report México y regiones adyacentes del Pacífico. Memorias. IV 142: 120–126. Congreso Nacional de Oceanografía, pp. 223–247. 14. Gasca, R. 1999. Siphonophores (Cnidaria) and summer 2. Bigelow, H. B. 1918. Some Medusae and Siphonophorae mesoscale features in the Gulf of Mexico. Bulletin of from the Western Atlantic. Bulletin of the Museum of Marine Science 65: 75–89. Comparative Zoölogy at Harvard College 62: 363–442, 8 pls. 15. Gasca, R. 2002. Lista faunística y bibliografía comentados 3. Biggs, D. C., D. E. Smith, R. R. Bidigare, and M. A. John- de los sifonóforos (Cnidaria: Hydrozoa) de México. Anales son. 1984. In situ estimation of the population density of del Instituto de Biología, Universidad Nacional Autónoma gelatinous planktivores in Gulf of Mexico surface waters. de México Serie Zoología 73: 123–143. Memorial University of Newfoundland Occasional Papers 16. Gasca, R., and E. Suárez. 1989. Nota acerca de los sifonó- in Biology 9: 17–34. foros (Cnidaria: Siphonophorae) del Canal de Yucatán 4. Bouillon, J., M. D. Medel, F. Pagès, J. M. Gili, F. Boero, and (mayo–junio 1984). Caribbean Journal of Science 25: C. Gravili. 2004. Fauna of the Mediterranean Hydrozoa. 66–70. Scientia Marina 68(Supl. 2): 5–438. 17. Gasca, R., and E. Suárez. 1993. Primer registro de Lensia 5. Burke, W. D. 1975. Pelagic Cnidaria of Mississippi Sound canopusi Stepanjants (Cnidaria: Siphonophora) en el and adjacent waters. Gulf Research Reports 5: 23–38. Océano Atlántico. Caribbean Journal of Science 29: 6. Carré, C., and D. Carré. 1995. Ordre des Siphonophores. 126–127. Pp. 523–596 in D. Doumenc, ed. Traité de Zoologie. 18. Juárez- Fernández, M. 1965. Lista preliminary de los Anatomie, Systématique, Biologie.
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