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Synopsis of the Families and Genera of the Hydromedusae of the World, with a List of the Worldwide Species

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Phylogeny and Classification of Hydroidomedusae SYNOPSIS OF THE FAMILIES AND GENERA OF THE HYDROMEDUSAE OF THE WORLD, WITH A LIST OF THE WORLDWIDE SPECIES. Jean Bouillon (1) and Ferdinando Boero (2) (1) Laboratoire de Biologie Marine, Université Libre de Bruxelles, 50 Ave F. D. Roosevelt, 1050 Bruxelles, Belgium. (2) Dipartimento di Biologia, Stazione di Biologia Marina, Università di Lecce, 73100 Lecce, Italy. Abstract: This report provides a systematic review of the pelagic Hydrozoa, Siphonophores excluded; diagnoses and keys are given for the different families and genera with a short description of their hydroid stage where known; a list of the world-wide hydromedusae species is established. Key words: Hydrozoa, Automedusae, Hydroidomedusae, systematics, diagnosis A: INTRODUCTION: The hydromedusae are on the whole one of the best known groups of all the Hydrozoa, three great monographs covering the world-wide described species having been dedicated to them, the first by Haeckel (1879-1880), the second by Mayer (1910) and the last by Kramp (1961). A generic revision has been done by Bouillon, 1985, 1995 and several large surveys covering various 47 Thalassia Salentina n. 24/2000 geographical regions have been published in recent times, more particularly, those by Kramp, 1959 the “Atlantic and adjacent waters”, 1968 “Pacific and Indian Ocean”, Arai and Brinckmann-Voss, 1980 “British Columbia and Puget Sound”; Bouillon, 1999 “South Atlantic”; Bouillon and Barnett, 1999 “New- Zealand”; Boero and Bouillon, 1993 and Bouillon et al, (in preparation) “ Mediterranean”; they all largely improved our knowledge about systematics and hydromedusan biodiversity. The present work is a compilation of all the genera and species of hydromedusae known, built up from literature since Kramp’s 1961 synopsis to a few months before publication. Keys and diagnoses are given for the families and genera; where known a brief description is given of the hydroid stage. Such work, although as extensive as possible, can never been considered complete, presenting both errors and omissions and of is course essentially not definitive needing continuous updates. It may nevertheless be a useful tool and assist students of the group. In all, 1086 hydromedusae species have been recorded from modern literature; 244 of them are however considered either as doubtful species, or as conspecific, or as being insufficiently or improperly described or known only as medusa buds, in consequence 842 hydromedusae species may been estimated as valid. Two hundred sixty two genera of Hydrozoan medusae have been recorded, 41 belong to the Automedusae, 222 to the Hydroidomedusae. Less than half of the 222 Hydroidomedusae genera have one or some species with their cycle known. A great effort has thus to be done by Hydrozoan specialist to study Hydroidomedusae life cycles, the knowledge of which is an essential condition for establishing valid species definition and a sound phylogenetic classification of those animals. The exclusion of species with no medusa stage is due to an even more complex taxonomic situation than that of the hydromedusae. A host of thecate species, for instance, has been described on the basis of tiny skeletal details that might be simply due to environment-induced variations. Careful revisions are badly needed to “clean” a possible list of all Hydroidomedusae from synonymies. 48 Phylogeny and Classification of Hydroidomedusae B: OUTLINE CLASSIFICATION: Subclass Actinulidae Swedmark and Teissier, 1959 Family Halammohydridae Remane 1927 Family Otohydridae Swedmark and Teissier, 1958 Subclass Anthomedusae Haeckel, 1879 Order Filifera Kühn, 1913 Suborder Margelina Haeckel, 1879 Family Australomedusidae Russell, 1971 Family Bougainvillidae Lütken, 1850 Family Clavidae McCrady, 1859 Family Cytaeididae L. Agassiz, 1862 Family Eucodoniidae Schuchert, 1996 Family Hydractiniidae L. Agassiz, 1862 Family Ptilocodiidae Coward, 1909 Family Rathkeidae Russell, 1953 Family Trichydridae Hincks, 1868 Suborder Pandeida Haeckel,1879 Family Bythotiaridae Maas, 1905. ( = Calycopsidae;) Family Niobiidae Petersen, 1979 Family Pandeidae Haeckel, 1879 Family Proboscidactylidae Hand and Hendrickson, 1950 Family Protiaridae, Haeckel 1879 Family Russelliidae Kramp, 1957 Order Capitata Khün, 1913 Suborder Moerisiida Poche, 1914 Family Moerisiidae Poche, 1914 Family Halimedusidae Arai and Brinckmann-Voss, 1980 Family Polyorchidae Agassiz, 1862 Family Urashimeidae Mills 2000 (in press) Suborder Sphaerocorynida Petersen, 1990 49 Thalassia Salentina n. 24/2000 Family Hydrocorynidae Rees, 1957 Family Sphaerocorynidae Prévot, 1959 Family Zancleopsidae Bouillon, 1978 Suborder Tubulariida, Fleming, 1828 Family Boeromedusidae Bouillon, 1985 Family Cladonematidae Gegenbaur, 1857 Family Corynidae Johnston, 1836 Family Corymorphidae Allman, 1872 Family Dicyclocorynidae Petersen, 1979 Family Eleutheriidae Russell, 1953 Family Euphysidae Haeckel, 1879 Family Margelopsidae Uchida, 1927 Family Pennariidae McCrady, 1859 Family Tubulariidae Fleming, 1828 Suborder Zancleida Russell, 1953 Family Asyncorynidae Kramp, 1949 Family Cladocorynidae Allman, 1872 Family Porpitidae Goldfuss, 1818 Family Rosalindidae Bouillon, 1985 Family Teissieridae Bouillon, 1974 Family Zancleidae Russell, 1953 Subclass Laingiomedusae Bouillon, 1978 Family Laingiidae Bouillon, 1978 Subclass Leptomedusae Haeckel, 1866 (1879) Order Conica Broch, 1910 Family Aequoreidae Eschscholtz, 1829 Family Aglaopheniidae Marktanner-Turneretscher, 1890 Family Barcinidae Gili, Bouillon, Pagès, Palanques and Puig, 1999 Family Blackfordiidae Bouillon, 1984 Family Cirrholoveniidae Bouillon, 1984 Family Clathrozoidae Hirohito, 1967 Family Dipleurosomatidae Russell, 1953 50 Phylogeny and Classification of Hydroidomedusae Family Eirenidae Haeckel, 1879 Family Haleciidae Hincks, 1868 Family Lafoeidae Hincks, 1868 Family Laodiceidae Agassiz, 1862 Family Lovenellidae Russell, 1953 Family Malagazziidae Bouillon, 1984 Family Melicertidae Agassiz, 1862 Family Mitrocomidae Haeckel, 1879 (part); Torrey, 1909 Family Octocannoidae Bouillon, Seghers and Boero, 1991 Family Orchistomatidae Bouillon, 1984 Family Parateclaiidae Bouillon, Pages and Gili, 2000 Family Phialellidae Russell, 1953 Family Plumulariidae Agassiz, 1862 (Hincks, 1868) Family Sertulariidae Lamouroux, 1812 Family Sugiuridae Bouillon, 1984 Family Teclaiidae Bouillon, Pagès, Gili, Palanques, Puig and Heusner, 1999 Family Tiarannidae Russell, 1940 Family Tiaropsidae Boero, Bouillon and Danovaro, 1987 Order Proboscoida Broch, 1910 Family Campanulariidae Jonhston, 1836 Family Phialucidae Bouillon, 1984 Subclass Limnomedusae Kramp, 1938 Family Armorhydridae Swedmark and Teissier, 1958 Family Olindiidae Haeckel, 1879 Subclass Narcomedusae Haeckel, 1879 Family Aeginidae Gegenbaur, 1857, emend. Maas, 1904 Family Cuninidae Bigelow, 1913 Family Solmarisidae Haeckel, 1879 Subclass Polypodiozoae Raikova, 1988 Family Polypodiidae Poche, 1914 Subclass Siphonophorae Eschscholtz, 1829 not treated here Subclass Trachymedusae Haeckel, 1866 (1879) 51 Thalassia Salentina n. 24/2000 Family Geryoniidae Eschscholtz, 1829 Family Halicreatidae Fewkes, 1886 Family Petasidae Haeckel, 1879 Family Ptychogastriidae Mayer, 1910 Family Rhopalonematidae Russell, 1953 C: SIMPLIFIED KEY FOR SUB-CLASSES IDENTIFICATION: A- without statocysts: A1-gonads on manubrium occasionally on manubrium and extending for a short distance along most basal region of radial canals; marginal tentacles solid or hollow, peripheral on exumbrellar rim; exumbrella entire, not divided into lobes; usually with marginal tentacular bulbs; with radial canals and circular canal; sense organs, where present, ocelli; medusa typically bell-shaped; life cycle with hydroid stage: Anthomedusae. A2-gonads exclusively on manubrium; marginal tentacles solid, leaving umbrella some distance above exumbrella margin; with or without marginal tentacular bulbs; with umbrella divided by peronial grooves or similar structures so that umbrellar margin is more or less lobed; with radial canals, but without typical circular canal but a solid core of endodermal cells around umbrellar margin; anatomically visible sense organs apparently missing; umbrella roughly hemispherical; hydroid stage unknown: Laingiidae. B-statocysts, where present, formed exclusively by the velar ectoderm, open or enclosed; gonads on radial canals, exceptionally contiguous with base of manubrium; marginal tentacles usually hollow, peripheral on exumbrellar rim; with marginal tentacular bulbs; exumbrella entire, not divided into lobes; with radial and 52 Phylogeny and Classification of Hydroidomedusae circular canal; sometimes cordyli, rarely ocelli; umbrella usually flattened; life cycle with hydroid stage: Leptomedusae. C-statocysts, where present, as ecto-endodermal statocysts with endodermal axis issued from ring canal: C1- pelagic hydromedusae: C1a- gonads only on radial canals; marginal tentacles solid (rarely, Geronyidae, with a mixture of solid and hollow tentacles), tentacles peripheral on exumbrella margin; exumbrella entire, not divided into lobes; with an exumbrellar marginal cnidocyst ring; without marginal tentacular bulbs; with radial canals and circular canal; umbrella tall to hemispherical; usually with free, more rarely enclosed, statocysts usually growing out of umbrella margin; without ocelli; without hydroid stage: Trachymedusae. C1b- gonads only on manubrium or on manubrial pouches; tentacles solid, living exumbrella at some distance above exumbrellar margin; without marginal tentacular
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