Hydroidolina
SUBCLASS HYDROIDOLINA Hydroidolina Collins, 2000 includes Anthoathecata, Leptothecata, and Siphonophorae (see Marques & Collins 2004; Collins et al. 2006 a). The monophyly of Hydroidolina is well supported by phylogenetic analyses of molecular (Bridge et al. 1995; Collins 2002; Collins et al. 20...
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| Summary: | SUBCLASS HYDROIDOLINA Hydroidolina Collins, 2000 includes Anthoathecata, Leptothecata, and Siphonophorae (see Marques & Collins 2004; Collins et al. 2006 a). The monophyly of Hydroidolina is well supported by phylogenetic analyses of molecular (Bridge et al. 1995; Collins 2002; Collins et al. 2006 a) and morphological (Bouillon & Boero 2000 a; Marques & Collins 2004) data. The statocysts of Hydroidolina, when present, are ectodermal in origin. Hydroidolina polyps may be solitary or colonial, and the colonies may be polymorphic, a state that is only present in one genus ( Monobrachium ) of Trachylina. The relationships between major groups of Hydroidolina (Leptothecata, Anthoathecata, Siphonophorae) are uncertain (Collins 2002; Collins et al. 2006 a). Order Anthoathecata Anthoathecata Cornelius, 1992 comprises two suborders, Filifera and Capitata, and approximately 1,140 valid species (Bouillon et al. 2006). The polyps do not have a skeletal covering and can be solitary or colonial. The medusae do not have statocysts and the gametogenic tissue is confined to the manubrium. Molecular phylogenetic studies do not support monophyly of Anthoathecata, suggesting instead that Anthoathecata is a paraphyletic assemblage that gave rise to one or more of the other suborders of Hydroidolina (Collins et al. 2006 a). Suborder Filifera Filifera comprises 22 families (Schuchert 2007 a) and approximately 765 valid species (Bouillon et al. 2006). Although the suborder has the putative morphological synapomorphies of filiferan tentacles on the feeding polyps and desmoneme and eurytele nematocysts, molecular phylogenetic analyses do not support its monophyly (Collins 2002; Collins et al. 2005, 2006a). Included families Australomedusidae Russell 1971 comprises three genera (Schuchert 2007 a) and approximately five valid species (Bouillon et al. 2006). No representatives have been included in molecular phylogenetic analyses and its members have never been the subject of a phylogenetic analysis. This family is distinguished by polyps with large extensible hypostomes and medusae with (usually) four radial canals and four clusters of tentacles at the perradii of the bell margin. Balellidae Stechow 1922 is a monospecific family (Bouillon et al. 2006) whose sole species has not been included in molecular phylogenetic analyses. This family is distinguished by polyps with two widely separated whorls of filiform tentacles. Bougainvilliidae Lütken, 1850 comprises 13 genera (Schuchert 2007 a) and approximately 100 valid species (Bouillon et al. 2006). Bougainvilliidae includes Rhizorhagium , which is not found within Schuchert's (2007 b) classification (2007 a). Molecular phylogenetic analyses that include two species from two genera were ambiguous with regard to monophyly (Collins et al. 2006 a) and the group awaits a thorough phylogenetic investigation. Bougainvilliidae lacks morphological synapomorphies and shares many features with other families (Calder 1988; Schuchert 2007 c). Some genera classified elsewhere by Schuchert (2007 a), e.g., Lizzia in Rathkeidae, have been considered part of Bougainvilliidae (Bouillon et al. 2006). Bythotiaridae Maas, 1905 comprises nine genera (Schuchert 2007 a) and approximately 25 valid species (Bouillon et al. 2006). No representatives have been included in molecular phylogenetic analyses and the group has not been the subject of a phylogenetic analysis. Hydroids from this family can be distinguished by their habit of living inside the prebranchial cavity of ascidians. Medusae are recognized by marginal tentacles that have tiny or absent basal bulbs and terminate in a cluster of cnidae, but these features are present in Eucodoniidae. Clathrozoellidae Peña Cantero, Vervoort & Watson, 2003 is a monogeneric family (Schuchert 2007 a) with four valid species (Bouillon et al. 2006). No representatives have been included in molecular phylogenetic analyses. This family is distinguished by a pseudohydrotheca covering the polyps. Cytaeididae Agassiz, 1862 comprises three genera (Schuchert 2007 a) and approximately 20 valid species (Bouillon et al. 2006). No representatives have been included in molecular phylogenetic analyses and the group has not been the subject of an explicit phylogenetic analysis. There are no known morphological synapomorphies for this group and the validity of one of the genera ( Perarella ) is questionable (Bouillon et al. 2006; Schuchert 2007 c). Eucodoniidae Schuchert, 1996 is a monospecific family (Schuchert 1996) that has not been included in molecular phylogenetic analyses. This family is distinguished by four clusters of embedded nematocysts around the mouth margin of medusae (Schuchert 1996). Eudendriidae Agassiz, 1862 comprises two genera (Schuchert 2007 a) and approximately 85 valid species (Marques 1996). Only a single representative has been included in molecular phylogenetic analyses (Collins et al. 2006 a). Possible synapomorphies for this group include the absence of desmoneme nematocysts, a styloid gonophore, and trumpet-shaped hypostome (Marques 1996). Hydractiniidae Agassiz, 1862 comprises seven genera (Schuchert 2007 a) and approximately 100 valid species (Bouillon et al. 2006). Eleven species representing two genera have been included in molecular phylogenetic analyses, which supported monophyly of the group (Cunningham & Buss 1993). However, taxon sampling in this analysis was not broad. The hydractiniid genus Clava is sometimes classified together with genera of Oceanidae in the family Clavidae (see Bouillon et al. 2006), suggesting that its phylogenetic status is uncertain. Hydroids of the family are distinguished by stolonal, polymorphic colonies that may bear spines. Laingiidae Bouillon, 1978 comprises three genera and four valid species (Bouillon 1978; Bouillon et al. 2006). One species has been included in molecular phylogenetic analyses (Collins et al. 2006 a). This group was originally classified as its own subclass by Bouillon (1978) but molecular phylogenetic analyses have shown that at least one member of the group, Fabienna sphaerica , is nested within Hydroidolina and closely related to Proboscidactylidae (see Collins et al. 2006 a). This hypothesis is supported by morphological data, including a solid radial canal and macrobasic euryteles (Schuchert 1996). Niobiidae Peterson, 1979 is a monospecific family (Bouillon et al. 2006) whose sole species has not been included in molecular phylogenetic analyses. The hydroid stage is unknown and the medusae are distinguished by marginal tentacle bulbs that develop into medusae buds (Petersen 1979). Oceanidae Eschscholtz, 1829 comprises eight genera (Schuchert 2004) and approximately 25 valid species (Bouillon et al. 2006). Representatives of the family have not been included in any phylogenetic analyses. The family is distinguished by scattered filiform tentacles on the polyps, but this character is not specific to this group (Calder 1988; Schuchert 2004). Several genera of this family are sometimes classified with Clava in the family Clavidae (see Bouillon et al. 2006), suggesting that its phylogenetic status is uncertain. Pandeidae Haeckel, 1879 comprises 23 genera (Schuchert 2007 a) and approximately 75 valid species (Bouillon et al. 2006). One species has been included in molecular phylogenetic analyses (Collins et al. 2006 a), but no explicit analyses of its phylogeny have been attempted. There are no known synapomorphies for this group and it is probably not monophyletic, as it encompasses a diverse assemblage of genera (Calder 1988). Proboscidactylidae Hand & Hendrickson, 1950 is a monogeneric family (Schuchert 2007 a) with six valid species (Bouillon et al. 2006). One species has been included in molecular phylogenetic analyses (Collins et al. 2006 a). Hydroids of this family are distinguished by polymorphic stolonal colonies bearing gastrozooid polyps with two filiform tentacles. Protiaridae Haeckel, 1879 comprises five genera (Schuchert 2007 a) and approximately eight valid species (Bouillon et al. 2006). No representatives have been included in molecular phylogenetic analyses. Medusae of this family are distinguished by large, hollow tentacular bulbs and four well-developed tentacles that are often interspersed with short, solid tentacles. Ptilocodiidae Coward, 1909 comprises six genera (Schuchert 2007 a) and eight valid species (Bouillon et al. 2006). The group has not been the subject of a phylogenetic analysis. Hydroids of this family are distinguished by the absence of tentacles on the feeding polyps. Rathkeidae Russell, 1954 comprises six genera (Schuchert 2007 a) and approximately 20 valid species (Bouillon et al. 2006). Three species representing three genera have been included in molecular phylogenetic analyses and in phylogenetic analyses, and these support familial monophyly (Schuchert 2007 a). Species of this family are distinguished by primary medusae buds arising from stolons and secondary medusae buds arising interradially from the medusa manubrium. Rhysiidae Brinckmann, 1965 is a monogeneric family (Schuchert 2007 a) with three valid species (Bouillon et al. 2006). No representatives have been included in molecular phylogenetic analyses. This family is distinguished by dactylozooids covered with perisarc to the capitate apical tip and female gonozooids that transform into a sporosac-like structure. Russelliidae Kramp, 1957 is a monospecific family (Bouillon et al. 2006) whose sole species has not been included in molecular phylogenetic analyses. The hydroid stage is unknown and the medusa possesses marginal tentacles in groups of three: one large and two small hollow tentacles. The large tentacles are sunk into the umbrella margin, forming a furrow. Stylasteridae Gray, 1847 comprises 26 genera (Schuchert 2007 a) and approximately 260 valid species (Bouillon et al. 2006). Cairns (1984 b) published a cladistic analysis of the genera of this diverse group, but its monophyly was not tested because only one outgroup was considered. Nevertheless, species of the group are readily distinguished by a massive calcareous exoskeleton, often brightly pigmented, and they very likely form a clade. Trichydridae Hincks, 1868 is a monospecific family (Bouillon et al. 2006) whose sole species has not been included in molecular phylogenetic analyses. No putative synapomorphies are known in the hydroid stage, but medusae are distinguished by the possession of many tiny, anastomosing centripetal canals. Tubiclavoididae Moura, Cunha & Schuchert, 2007 is a monospecific family (Moura et al. 2007) whose sole species has been included in molecular phylogenetic analyses (Moura et al. 2007), the results of which have not been published. The species is characterized by elongate polyps with scattered filiform tentacles and hydrocauli covered with striated perisarc. Suborder Capitata Capitata comprises 26 families (Schuchert 2007 a) and approximately 375 valid species (Bouillon et al. 2006). The putative synapormorphies are stenotele nematocysts and capitate tentacles on the polyps or filiform tentacles in separated whorls (Bouillon & Boero 2000 b). Molecular phylogenetic analyses suggest that Capitata is a paraphyletic assemblage containing two clades, Aplanulata and the other capitates (Collins 2002; Collins et al. 2005, 2006a). The synapomorphy for Aplanulata is the absence of a ciliated planula larva (Petersen 1990). Four families, Tubulariidae, Corymorphidae, Candelabridae, and Hydridae have been sampled in molecular phylogenetic analyses that support monophyly of Aplanulata (Collins et al. 2005; Collins et al. 2006 a), but it is likely that other families are part of this group (Petersen 1990; Collins et al. 2006 a). Mosaics of features, very few of which appear to be unique to any particular family, distinguish the medusa stages of capitate families. Included families Acaulidae Fraser, 1924 comprises three genera and five valid species (Bouillon et al. 2006). No representatives have been included in molecular phylogenetic analyses and the group has not been the subject of an explicit phylogenetic analysis. Members of this family are distinguished by scattered capitate tentacles on the distal portions of their solitary polyps. Boeromedusidae Bouillon, 1985 is a monospecific family (Bouillon et al. 2006) whose sole species has not been included in molecular phylogenetic analyses. Hydroids are unknown; medusae have an apical projection, four tentacles terminating in nematocyst clusters, and four perradial pouches bearing gametes hanging from the manubrium. Boreohydridae Wesblad, 1947 comprises two genera (Bouillon 1985) and two valid species (Bouillon et al. 2006). No representatives have been included in molecular phylogenetic analyses and the group has not been the subject of an explicit phylogenetic analysis. This family is distinguished by small solitary polyps that possess a whorl of three to five diminutive tentacles (Schuchert 2006). Candelabridae Stechow, 1921 comprises three genera (Schuchert 2006) and approximately 20 valid species (Bouillon et al. 2006). Just a single representative has been included in molecular phylogenetic analyses (Collins et al. 2005) and no explicit tests of the group’s monophyly have been conducted. The family is distinguished by its solitary or pseudo-colonial polyps that are relatively large and bear numerous scattered capitate tentacles (Schuchert 2006). Cladocorynidae Allman, 1872 comprises two genera (Schuchert 2006) and seven valid species (Bouillon et al. 2006). Only a single representative has been included in molecular phylogenetic analyses (Collins et al. 2005) and no explicit tests of the group’s monophyly have been conducted. The putative synapomorphy for the family is patches of macrobasic euryteles on the body wall of the polyp (Petersen 1990). Cladonematidae Gegenbaur, 1856 comprises four genera (Schuchert 2006) and approximately 20 valid species (Bouillon et al. 2006). Phylogenetic analyses including three species representing two genera support monophyly of the group (Collins et al. 2005). The family is distinguished by benthic medusae with branched tentacles and adhesive structures at the tips of the medusae tentacles (Petersen 1990). Corymorphidae Allman, 1872 comprises 10 genera (Schuchert 2007 a) and approximately 45 valid species (Bouillon et al. 2006). Two species from one genus were included in molecular phylogenetic analyses, the results of which contradicted monophyly (Collins et al. 2005). There are no known synapomorphies for this group (Petersen 1990). Corynidae Johnston, 1836 comprises seven genera and approximately 90 valid species (Schuchert 2001). A molecular phylogenetic analysis sampling 13 species from four genera strongly contradicted monophyly of the group, with some species being closely allied to members of Polyorchidae (Collins et al. 2005). Not surprisingly, no morphological synapomorphies have been identified for Corynidae (Schuchert 2001). Halimedusidae Arai & Brinckmann-Voss, 1980 comprises three genera (Mills 2000; Schuchert 2007 a), each with a single valid species (Bouillon et al. 2006). No representatives have been included in molecular phylogenetic analyses and the group has not been the subject of an explicit phylogenetic analysis. Species of this family have small solitary polyps that give rise to medusae with distinct interradial peaks in jelly above the manubrium base, a feature also present in medusae of Boeromedusidae (Mills 2000). Hydridae Linnaeus, 1758 is a monogeneric family (Schuchert 2007 a) with approximately 30 valid species (Bouillon et al. 2006). A molecular phylogenetic analysis sampling three species representing both the green and brown hydra groups supported monophyly of the family (Collins et al. 2006 a). Hemmrich et al. (2007) sampled additional taxa (mainly focused on laboratory strains) and also found the group to be monophyletic, although this study aimed at elucidating relationships within the group. Hydridae is distinguished by the absence of medusae, its freshwater habitat, and lateral budding of polyps. Hydrocorynidae Rees, 1957 comprises two genera (Schuchert 2007 a) and three valid species (Bouillon et al. 2006). No representatives have been included in molecular phylogenetic analyses and the group has not been the subject of an explicit phylogenetic analysis. Hydroids of the group are colonial, with hydranths arising from a chitinized hydrorhizal plate. No putative synapomorphies have been identified for the medusa stage. Margelopsidae Uchida, 1927 comprises three genera (Schuchert 2006) and six valid species. No representatives have been included in molecular phylogenetic analyses and the group has not been the subject of an explicit phylogenetic analysis. The family is distinguished by its small, pelagic, solitary polyps. Milleporidae Fleming, 1828 is a monogeneric family with approximately seven valid species (Razak & Hoeksema 2003). One species has been sampled for molecular analysis (Collins et al. 2006 a). This family is distinguished by colonies that build massive calcareous skeletons, polyps with capitate tentacles, and dimorphism with gastrozooids and dactylozooids. Moerisiidae Poche, 1914 comprises three genera (Schuchert 2007 a) and fewer than 10 valid species (Bouillon et al. 2006). One representative has been included in molecular phylogenetic analyses (Collins et al. 2005), but the family has never been the subject of an explicit phylogenetic analysis. Polyp stages of this family, where known, are solitary with scattered filiform tentacles. Medusae are recognized by a manubrium with radial lobes that extend toward and connect with the radial canals. Paracorynidae Picard, 1957 is a monospecific family (Bouillon et al. 2006). No representatives have been included in molecular phylogenetic analyses. This family is distinguished by a flat, highly organized colony, with polymorphic zooids. Bouillon (1974) suggested that Paracoryne could be interpreted as an individual flattened polyp, rather than as a colony. Pennariidae McCrady, 1859 is a monogeneric family with two valid species (Schuchert 2006). A single, widespread representative has been included in molecular phylogenetic analyses and no explicit tests of the group’s monophyly have been conducted. The family is distinguished by a pinnate hydroid colony with polyps that contain an aboral whorl of filiform tentacles and capitate tentacles scattered towards the oral end. Polyorchidae Agassiz, 1862 comprises three genera (Schuchert 2007 a) and five valid species (Bouillon et al. 2006). Three species from two genera have been included in m : Published as part of Daly, Marymegan, Brugler, Mercer R., Cartwright, Paulyn, Collins, Allen G., Dawson, Michael N., Fautin, Daphne G., France, Scott C., Mcfadden, Catherine S., Opresko, Dennis M., Rodriguez, Estefania, Romano, Sandra L. & Stake, Joel L., 2007, The phylum Cnidaria: A review of phylogenetic patterns and diversity 300 years after Linnaeus *, pp. 127-182 in Zootaxa 1668 on pages 153-168, DOI: 10.5281/zenodo.180149 : {"references": ["Marques, A. C. & Collins, A. G. (2004) Cladistic analysis of Medusozoa and cnidarian evolution. Invertebrate Biology, 123 (1), 23 - 42.", "Collins, A. G., Schuchert, P., Marques, A. C., Jankowski, T., Medina, M. & Schierwater, B. (2006 a) Medusozoan phylogeny and character evolution clarified by new large and small subunit rDNA data and an assessment of the utility of phylogenetic mixture models. Systematic Biology, 55 (1), 97 - 115.", "Bridge, D., Cunningham, C. 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