A newly compiled data set of nearly complete sequences of the large subunit of the nuclear ribosome (LSU or 28S) sampled from 31 diverse medusozoans greatly clarifies the phylogenetic history of Cnidaria. These data have substantial power to discern among many of the competing hypotheses of relationship derived from prior work. Moreover, LSU data provide strong support at key nodes that were equivocal based on other molecular markers. Combining LSU sequences with those of the small subunit of the nuclear ribosome (SSU or 18S), we present a detailed working hypothesis of medusozoan relationships and discuss character evolution within this diverse clade. Stauromedusae, comprising the benthic, so-called stalked jellyfish, appears to be the sister group of all other medusozoans, implying that the free-swimming medusa stage, the motor nerve net, and statocysts of ecto-endodermal origin are features derived within Medusozoa. Cubozoans, which have had uncertain phylogenetic affinities since the elucidation of their life cycles, form a clade-named Acraspeda-with the scyphozoan groups Coronatae, Rhizostomeae, and Semaeostomeae. The polyps of both cubozoans and hydrozoans appear to be secondarily simplified. Hydrozoa is comprised by two well-supported clades, Trachylina and Hydroidolina. The position of Limnomedusae within Trachylina indicates that the ancestral hydrozoan had a biphasic life cycle and that the medusa was formed via an entocodon. Recently hypothesized homologies between the entocodon and bilaterian mesoderm are therefore suspect. Laingiomedusae, which has often been viewed as a close ally of the trachyline group Narcomedusae, is instead shown to be unambiguously a member of Hydroidolina. The important model organisms of the Hydra species complex are part of a clade, Aplanulata, with other hydrozoans possessing direct development not involving a ciliated planula stage. Finally, applying phylogenetic mixture models to our data proved to be of little additional value over a more traditional phylogenetic approach involving explicit hypothesis testing and bootstrap analyses under multiple optimality criteria. [18S; 28S; Cubozoa; Hydrozoa; medusa; molecular systematics; polyp; Scyphozoa; Staurozoa.].

Abstract. A cladistic analysis of 87 morphological and life history characters of medusozoan cnidarians, rooted with Anthozoa, results in the phylogenetic hypothesis (Anthozoa (Hydrozoa (Scyphozoa (Staurozoa, Cubozoa)))). Staurozoa is a new class of Cnidaria consisting of Stauromedusae and the fossil group Conulatae. Scyphozoa is redefined as including those medusozoans characterized by strobilation and ephyrae (Coronatae, Semaeostomeae, and Rhizostomeae). Within Hydrozoa, Limnomedusae is identified as either the earliest diverging hydrozoan lineage or as the basal group of either Trachylina (Actinulida (Trachymedusae (Narcomedusae, Laingiomedusae))) or Hydroidolina (Leptothecata (Siphonophorae, Anthoathecata)). Cladistic results are highly congruent with recently published phylogenetic analyses based on 18S molecular characters. We propose a phylogenetic classification of Medusozoa that is consistent with phylogenetic hypotheses based on our cladistic results, as well as those derived from 18S analyses. Optimization of the characters presented in this analysis are used to discuss evolutionary scenarios. The ancestral cnidarian probably had a sessile biradial polyp as an adult form. The medusa is inferred to be a synapomorphy of Medusozoa. However, the ancestral process (metamorphosis of the apical region of the polyp or lateral budding involving an entocodon) could not be inferred unequivocally. Similarly, character states for sense organs and nervous systems could not be inferred for the ancestral medusoid of Medusozoa.

Staurozoan classification is highly subjective, based on phylogeny-free inferences, and suborders, families, and genera are commonly defined by homoplasies. Additionally, many characters used in the taxonomy of the group have ontogenetic and intraspecific variation, and demand new and consistent assessments to establish their correct homologies. Consequently, Staurozoa is in need of a thorough systematic revision. The aim of this study is to propose a comprehensive phylogenetic hypothesis for Staurozoa, providing the first phylogenetic classification for the group. According to our working hypothesis based on a combined set of molecular data (mitochondrial markers COI and 16S, and nuclear markers ITS, 18S, and 28S), the traditional suborders Cleistocarpida (animals with claustrum) and Eleutherocarpida (animals without claustrum) are not monophyletic. Instead, our results show that staurozoans are divided into two groups, herein named Amyostaurida and Myostaurida, which can be distinguished by the absence/presence of interradial longitudinal muscles in the peduncle, respectively. We propose a taxonomic revision at the family and genus levels that preserves the monophyly of taxa. We provide a key for staurozoan genera and discuss the evolution of the main characters used in staurozoan taxonomy.

Hydroidolina is a group of hydrozoans that includes Anthoathecata, Leptothecata and Siphonophorae. Previous phylogenetic analyses show strong support for Hydroidolina monophyly, but the relationships between and within its subgroups remain uncertain. In an effort to further clarify hydroidolinan relationships, we performed phylogenetic analyses on 97 hydroidolinan taxa, using DNA sequences from partial mitochondrial 16S rDNA, nearly complete nuclear 18S rDNA and nearly complete nuclear 28S rDNA. Our findings are consistent with previous analyses that support monophyly of Siphonophorae and Leptothecata and do not support monophyly of Anthoathecata nor its component subgroups, Filifera and Capitata. Instead, within Anthoathecata, we find support for four separate filiferan clades and two separate capitate clades (Aplanulata and Capitata sensu stricto). Our data however, lack any substantive support for discerning relationships between these eight distinct hydroidolinan clades.

SYNOPSIS Fossil taxa of uncertain phylogenetic affinities can play a crucial role in the analysis of character evolution within major extant groups. Marques & Collins (2004) concluded that conulariids (?Ediacaran-Triassic) are an extinct group of medusozoan cnidarians most closely related to Stauromedusae. However, only six of the 87 characters used by these authors can be observed in conulariid fossils. Rescoring the character states of conulariids in a conservative manner yields a new hypothesis for the phylogenetic position of conulariids, namely that they are the sister group of the scyphozoan order Coronatae rather than Stauromedusae, which is revealed as the earliest diverging lineage of Medusozoa. This new hypothesis also implies several different sequences of character evolution within Cnidaria. Specifically, the presence of a periderm completely covering the polyp in conulariids and coronates appears to be derived within Scyphozoa. Strobilation appears to be a synapomorphy uniting conulariids, Coronatae, Rhizostomeae and Semaeostomeae. This result supports the controversial interpretation of one exceptionally preserved conulariid that potentially shows that these animals produced ephyrae by strobilation. Finally, the pelagic adult medusa stage and the giant fibre nerve net appear to be features that are derived within Medusozoa.

ResumoLista dos Cnidaria Medusozoa do Brasil Uma lista dos Cnidaria Medusozoa marinhos do Brasil foi composta a partir de registros de ocorrência disponíveis na literatura. Até o momento, há um total de 373 espécies registradas para o Brazil: 347 de Hydrozoa, 3 de Cubozoa e 23 de Scyphozoa.http://www.biotaneotropica.org.br AbstractChecklist of the Cnidaria Medusozoa from Brazil Literature records were reviewed to compile a list of species of the marine taxa of Cnidaria MedusozoaThe total number of species of medusozoans so far recorded for Brazil is 373: 347 Hydrozoa, 3 Cubozoa: 23 Scyphozoa.recorded for the Brazilian coast.

Revision of the scyphozoan genus Chrysaora Péron & Lesueur, 1810 was undertaken from observations on museum material (Brazil, Europe, and USA), on living specimens in nature, and on life-cycles of some species cultured under laboratory conditions. A total of 168 museum lots, some of them having many medusae, were inspected. Included amongst these were nine type specimens. The genus comprises 13 valid species (Chrysaora achlyos, C. chinensis, C. colorata, C. fulgida, C. fuscescens, C. hysoscella, C. lactea, C. melanaster, C. pacifica, C. pentastoma, C. plocamia, and C. quinquecirrha), one species inquirenda (Chrysaora caliparea), and two doubtful species (C. kynthia and C. wurlerra). Differentiation of species is based mostly on tentacle number, shape of radial septa, order of tentacle development, colouration, and measurements of nematocysts. We resurrect C. chinensis for specimens from southeast Asia. Chrysaora pacifica is considered valid and distinct from C. melanaster based on tentacle number and nematocyst complement. Mediterranean specimens assigned to C. hysoscella are hermaphroditic and thereby considered distinct from those of C. fulgida from west Africa. Chrysaora achlyos (northeast Pacific) and C. plocamia (southeast Pacific and southwest Atlantic) are geographically isolated but morphologically identical, being distinguished only by colour pattern. The recently described C. southcotti is considered a junior synonym of C. pentastoma. The Australian C. kynthia and C. wurlerra, here considered nomina dubia, merit further study. Our phylogenetic hypothesis indicates that the genus Chrysaora forms a monophyletic group, with C. colorata, C. plocamia, and C. achlyos having a basal position in the phylogeny. Species with more than 24 tentacles (formerly assigned to the genus Dactylometra) form a clade with a derived position.

Diploblastic eumetazoans of the phylum Cnidaria originated during the Neoproterozoic Era, possibly during the Cryogenian Period. The oldest known fossil cnidarians occur in strata of Ediacaran age and consist of polypoid forms that were either nonbiomineralizing or weakly so. The oldest possible anthozoans, including the genus Ramitubus, may be related to tabulate corals and occur in the Doushantuo Lagerstätte (upper Doushantuo Formation, South China), the age of which is poorly constrained (approximately 585 Ma?). Conulariid scyphozoans may first appear as early as 635–577 Ma (Lantian Formation, South China). A definite conulariid, most similar to Palaeozoic species assigned to the genus Paraconularia, occurs in association with the possible scyphozoan, Corumbella werneri, in the latest Ediacaran (c. 543 Ma) Tamengo Formation of Brazil. Basal Cambrian (c. 540 Ma) phosphorites in the upper Kuanchuanpu Formation (South China) yield solitary polyps of the oldest probable anthozoan (Eolympia pediculata), which appears to have been a stem hexacorallian. This same formation contains fossils interpreted by some authors as pentaradial cubozoan polyps; however, both the oldest known cubozoans and the oldest hydrozoans, all medusae, may actually occur in the Cambrian (Series 3, c. 505 Ma) Marjum Formation (Utah, USA). Although these recently published palaeontological data tend to corroborate the hypothesis that Cnidaria has a relatively deep Neoproterozoic history, the timing of major internal branching events remains poorly constrained, with, for example, the results of some molecular clock analyses indicating that the two cnidarian subphyla (Anthozoaria and Medusozoa) may have originated as many as one billion years ago. Further progress towards elucidating the evolution and early fossil record of cnidarians may accrue from: (1) an intensive search for phosphatized soft parts in possible anthozoans from the Ediacaran Doushantuo Formation; (2) an expanded search for Ediacaran conulariids; and (3) additional detailed analyses of the taphonomy and preservation of Ediacaran and Cambrian cnidarians, including possible pentaradial cubozoan polyps from the Fortunian upper Kuanchuanpu Formation.