Systematics and character evolution of capitate hydrozoans.

Capitate hydrozoans are a morphologically and ecologically diverse hydrozoan suborder, currently including about 200 species. Being grouped in two clades, Corynida and Zancleida, these hydrozoans still show a number of taxonomic uncertainties at the species, genus and family levels. Many Capitata species established symbiotic relationships with other benthic organisms, including bryozoans, other cnidarians, molluscs and poriferans, as well as with planktonic dinoflagellates for mixotrophic relationships and with bacteria for thiotrophic ectosymbioses. Our study aimed at providing an updated and comprehensive phylogeny reconstruction of the suborder, at modelling the evolution of selected morphological and ecological characters, and at testing evolutionary relationships between the symbiotic lifestyle and the other characters, by integrating taxonomic, ecological and evolutionary data. The phylogenetic hypotheses here presented shed light on the evolutionary relationships within Capitata, with most families and genera being recovered as monophyletic. The genus Zanclea and family Zancleidae, however, were divided into four divergent clades, requiring the establishment of the new genus Apatizanclea and the new combinations for species in Zanclea and Halocoryne genera. The ancestral state reconstructions revealed that symbiosis arose multiple times in the evolutionary history of the Capitata, and that homoplasy is a common phenomenon in the group. Correlations were found between the evolution of symbiosis and morphological characters, such as the perisarc. Overall, our results highlighted that the use of genetic data and a complete knowledge of the life cycles are strongly needed to disentangle taxonomic and systematic issues in capitate hydrozoans. Finally, the colonization of tropical habitat appears to have influenced the evolution of a symbiotic lifestyle, playing important roles in the evolution of the group.

Modern alongside traditional taxonomy-Integrative systematics of the genera Gymnangium Hincks, 1874 and Taxella Allman, 1874 (Hydrozoa, Aglaopheniidae).

We studied the diversity within the former genus Gymnangium in the South West Indian Ocean by using an integrative approach of both traditional (morphology-based) and modern molecular taxonomy. Nine species were recorded in the material collected. A total of 97 16S mitochondrial DNA sequences and 54 Calmodulin nuclear sequences from eight Gymnangium/Taxella species were analyzed. We found both morphological and molecular differences in the studied Gymnangium species that make it necessary to split the genus. It is proposed to revalidate the genus Taxella which is currently regarded as a synonym of Gymnangium. Two species of the genus Taxella (T. eximia and T. gracilicaulis), until now regarded as distinct species based on morphological characteristics, cluster together in one phylogenetic clade. Possible explanations are discussed. Two species from Madagascar new to science are herein described and rare species from the Indian Ocean islands are re-described.

The polyps of Oceania armata identified by DNA barcoding (Cnidaria, Hydrozoa).

The polyps of the widely distributed medusa Oceania armata have never been found in nature and only the primary polyp is known from breeding experiments. The fully developed colony is so far unknown. This report shows that DNA sequence data of the medusa stage of O. armata permits to identify several hydroid colonies from different geographic origins as the most likely polyp stage of this medusa. These hydroids had previously been misidentified as Turritopsis species, a closely related genus which also produces medusae resembling Oceania armata. It is concluded that most Turritopsis hydroids are not reliably identifiable to species level using morphological traits only. However, DNA barcodes, particularly 16S sequences, are an excellent tool to identify the species, although we still lack information on a few nominal species and the identities of some sequence-delimited clades need to be corroborated by the addition of topotype samples.

High genetic diversity in the hydroid Plumularia setacea: a multitude of cryptic species or extensive population subdivision?

The marine hydroid Plumularia setacea has a near-cosmopolitan distribution. As in other sessile invertebrates with limited dispersal abilities, the wide distribution could also be a taxonomic artefact and the species might in fact be a complex of sibling species. To investigate this, a set of worldwide samples of P. setacea and several closely related species was examined using the mitochondrial markers 16S and COI, as well as the nuclear marker ITS. The results suggest an even higher degree of genetic diversity than expected. Almost all sampled regions had only private haplotypes and the resulting trees split into a multitude of geographically delimited lineages, this both for the mitochondrial and nuclear markers. In the framework of a genealogical species concept, these lineages would qualify as cryptic species. Using alternative species concepts, the results could be reconciled with traditional taxonomy by regarding P. setacea as a single species with an extensive population subdivision. A rapid molecular clock, limited dispersal abilities, and localized clonal propagation are likely the factors that explain the high but dispersed genetic diversity within this species.

Phylogenetic placement of Hydra and relationships within Aplanulata (Cnidaria: Hydrozoa).

The model organism Hydra belongs to the hydrozoan clade Aplanulata. Despite being a popular model system for development, little is known about the phylogenetic placement of this taxon or the relationships of its closest relatives. Previous studies have been conflicting regarding sister group relationships and have been unable to resolve deep nodes within the clade. In addition, there are several putative Aplanulata taxa that have never been sampled for molecular data or analyzed using multiple markers. Here, we combine the fast-evolving cytochrome oxidase 1 (CO1) mitochondrial marker with mitochondrial 16S, nuclear small ribosomal subunit (18S, SSU) and large ribosomal subunit (28S, LSU) sequences to examine relationships within the clade Aplanulata. We further discuss the relative contribution of four different molecular markers to resolving phylogenetic relationships within Aplanulata. Lastly, we report morphological synapomorphies for some of the major Aplanulata genera and families, and suggest new taxonomic classifications for two species of Aplanulata, Fukaurahydra anthoformis and Corymorpha intermedia, based on a preponderance of molecular and morphological data that justify the designation of these species to different genera.

The status of Plumularia lagenifera Allman, 1885 (Cnidaria, Hydrozoa) and related species.

The current status of Plumularia lagenifera Allman, 1885, a common thecate hydroid of the west coast of the USA and Canada, is problematic as it is difficult to distinguish from the near cosmopolitan and very variable Plumularia setacea. Type material of P. lagenifera and newly collected material of P. lagenifera and P. setacea from the region of the type locality of the former was used to compare it to P. setacea from the Atlantic. Measurements of a number of morphological traits were made and analysed using principal components analyses. Type material of the Californian Plumularia palmeri Nutting, 1900 was also included in the comparisons and confirmed the view of earlier workers that it is indistinguishable from P. setacea. Additionally, South African material referred to P. lagenifera by Millard (1975) was compared to the material from the NE Pacific. Plumularia lagenifera remains difficult to separate from P. setacea. The convex outer wall of the hydrotheca offers the only operational character to distinguish P. lagenifera from P. setacea, which always has straight or even concave hydrothecae. For morphological and biogeographic reasons, South African P. lagenifera sensu Millard (1975) should be referred to P. gaimardi (Lamouroux, 1924).

Molecular phylogenetics of Thecata (Hydrozoa, Cnidaria) reveals long-term maintenance of life history traits despite high frequency of recent character changes.

Two fundamental life cycle types are recognized among hydrozoan cnidarians, the benthic (generally colonial) polyp stage either producing pelagic sexual medusae or directly releasing gametes elaborated from an attached gonophore. The existence of intermediate forms, with polyps producing simple medusoids, has been classically considered compelling evidence in favor of phyletic gradualism. In order to gain insights about the evolution of hydrozoan life history traits, we inferred phylogenetic relationships of 142 species of Thecata (= Leptothecata, Leptomedusae), the most species-rich hydrozoan group, using 3 different ribosomal RNA markers (16S, 18S, and 28S). In conflict with morphology-derived classifications, most thecate species fell in 2 well-supported clades named here Statocysta and Macrocolonia. We inferred many independent medusa losses among Statocysta. Several instances of secondary regain of medusoids (but not of full medusa) from medusa-less ancestors were supported among Macrocolonia. Furthermore, life cycle character changes were significantly correlated with changes affecting colony shape. For both traits, changes did not reflect graded and progressive loss or gain of complexity. They were concentrated in recent branches, with intermediate character states being relatively short lived at a large evolutionary scale. This punctuational pattern supports the existence of 2 alternative stable evolutionary strategies: simple stolonal colonies with medusae (the ancestral strategy, seen in most Statocysta species) versus large complex colonies with fixed gonophores (the derived strategy, seen in most Macrocolonia species). Hypotheses of species selection are proposed to explain the apparent long-term stability of these life history traits despite a high frequency of character change. Notably, maintenance of the medusa across geological time in Statocysta might be due to higher extinction rates for species that have lost this dispersive stage.

Medusozoan phylogeny and character evolution clarified by new large and small subunit rDNA data and an assessment of the utility of phylogenetic mixture models.

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.].