Phylogeny of Capitata and Corynidae (Cnidaria, Hydrozoa) in light of mitochondrial 16S rDNA data

New sequences of the partial rDNA gene coding for the mitochondrial large ribosomal subunit, 16S, are derived from 47 diverse hydrozoan species and used to investigate phylogenetic relationships among families of the group Capitata and among species of the capitate family Corynidae. Our analyses identify a well-supported clade, herein named Aplanulata, of capitate hydrozoans that are united by the synapomorphy of undergoing direct development without the ciliated planula stage that is typical of cnidarians. Aplanulata includes the important model organisms of the group Hydridae, as well as species of Candelabridae, Corymorphidae, and Tubulariidae. The hypothesis that Hydridae is closely related to brackish water species of Moerisiidae is strongly controverted by 16S rDNA data, as has been shown for nuclear 18S rDNA data. The consistent phylogenetic signal derived from 16S and 18S data suggest that both markers would be useful for broad-scale multimarker analyses of hydrozoan relationships. Corynidae is revealed as paraphyletic with respect to Polyorchidae, a group for which information about the hydroid stage is lacking. Bicorona , which has been classified both within and outside of Corynidae, is shown to have a close relationship with all but one sampled species of Coryne . The corynid genera Coryne , Dipurena , and Sarsia are not revealed as monophyletic, further calling into question the morphological criteria used to classify them. The attached gonophores of the corynid species Sarsia lovenii are confirmed as being derived from an ancestral state of liberated medusae. Our results indicate that the 16S rDNA marker could be useful for a DNA-based identification system for Cnidaria, for which it has been shown that the commonly used cytochrome c oxidase subunit 1 gene does not work.     

Evolution and taxonomy in capitate hydroids and medusae (Cnidaria: Hydrozoa)

A cladistic analysis of Capitata groups the families in four suborders based on medusa characters (such as manubrium morphology, position of gonads, and position and number of marginal tentacles) and hydroid characters (such as presence or absence of an oral tentacle whorl, and the different development of the tentacles of the oral and aboral whorls). On the family and generic levels, the revision results in changes which unite the separate hydroid and medusa taxonomic systems, defining genera which are not based on characters solely relating to the reduction of medusae to fixed gonophores. In those families where the reduction of the medusa can be analysed, it is shown that the reduction occurred after all synapomorphies defining the genera had evolved and usually affected individual species within a genus rather than the original species from which the other species in the genus evolved. This supports the view that medusa reduction is not in itself a valid generic character. A discussion of the theories of 'inconsistent' or 'mosaic' evolution concludes that no difference in evolutionary rate or degree of specialization can be demonstrated among taxa with free medusae and taxa with gonophores.     

Cladistic analysis of Medusozoa and cnidarian evolution

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.     

Phylogenetic analysis of the Cnidaria

The recent members of the phylum Cnidaria were analyzed with phylogenetic methodology and the help of the PAUP Computer program. The Cnidaria are established as a monophylum by their cnidocysts, planula larva, and a polyp stage. The Ctenophora were seen as the most probable sister group of the Cnidaria. Arguments for the monophyly of the cnidarian classes Anthozoa, Scyphozoa, Cubozoa, and Hydrozoa were providea. For the ground plan of the Cnidaria the following characters were postulated: triphasic life cycle consisting of a planula larva, a benthic polyp stage, and a sexually propagating medusa like stage. For the polyp a radial symmetry, lack of septae, and hollow tentacles were assumed. The original medusa probably was tetraradial and developed from the polyp stage by a total metamorphosis. Twelve polarized characters were used to generate cladograms. The most parsimonious one showed the Anthozoa as the first offshoot of the tree with the united Scyphozoa, Cubozoa and Hydrozoa forming its sister group. Within this sister group the Scyphozoa and Cubozoa were seen as sistergroups to each other. Both groups united are then the sistergroup of the Hydrozoa. A bootstrap analysis yielded the same tree with high probabilities for the internal nodes. Despite assuming a planktonic origin of the Cnidaria in this investigation, the resulting cladogram is also compatible with an evolution of the medusa stage within the Cnidaria after the splitting-off of the Anthozoa. The possible loss of the medusa stage in the Anthozoa is discussed.     

Phylogeny of Medusozoa and the evolution of cnidarian life cycles

To investigate the evolution of cnidarian life cycles, data from the small subunit of the ribosome are used to derive a phylogenetic hypothesis for Medusozoa. These data indicate that Cnidaria is monophyletic and composed of Anthozoa and Medusozoa. While Cubozoa and Hydrozoa are well supported clades, Scyphozoa appears to be paraphyletic. Stauromedusae is possibly the sister group of either Cubozoa or all other medusozoans. The phylogenetic results suggest that: the polyp probably preceded the medusa in the evolution of Cnidaria; within Hydrozoa, medusa development involving the entocodon is ancestral; within Trachylina, the polyp was lost and subsequently regained in the parasitic narcomedusans; within Siphonophorae, the float originated prior to swimming bells; stauromedusans are not likely to be descended from ancestors that produced medusae by strobilation; and cubozoan polyps are simplified from those of their ancestors, which possessed polyps with gastric septa and four mesogleal muscle bands and peristomial pits.     

Phylogeny of the Plumularioidea (Hydrozoa, Leptothecata): evolution of colonial organisation and life cycle

The Plumularioidea (Cnidaria, Hydrozoa, Leptothecata) are the most species rich superfamily of the class Hydrozoa. They display a complex and diversified colonial organisation and their life cycle comprises either a reduced free-living, pelagic generation (medusoid), alternating with the benthic colonial form or in most species, no pelagic generation. In order to understand the evolution of colonial and life cycle characters among Plumularioidea, we have reconstructed their phylogeny. Partial mitochondrial 16S rRNA sequences and 64 morphological characters were analysed separately and in combination. The morphological data included not only characters of the individual polyps and medusae, but also characters describing the organisation of colonies, for which we propose general principles applying to character coding in modular organisms. The phylogenetic analyses supported the monophyly of Plumularioidea and of the four plumularioid families (Aglaopheniidae, Halopterididae, Kirchenpaueriidae and Plumulariidae). Most genera were paraphyletic or polyphyletic. This study highlights multiple morphological simplifications of the colonial organisation during the evolution of Plumularioidea and the convergence of the defensive polyps — the dactylozooids — of Plumularioidea with those of others Leptothecata ( Hydrodendron ) or Anthoathecata ( Hydractinia ). Concerning the evolution of the life cycle, the phylogeny supports a provocative scenario, where the medusa was lost in an ancestor of the Plumularioidea, and then re-acquired four times independently within this group, in the form of simple medusoids.     

Evolutionary relationships among scyphozoan jellyfish families based on complete taxon sampling and phylogenetic analyses of 18S and 28S ribosomal DNA

A stable phylogenetic hypothesis for families within jellyfish class Scyphozoa has been elusive. Reasons for the lack of resolution of scyphozoan familial relationships include a dearth of morphological characters that reliably distinguish taxa and incomplete taxonomic sampling in molecular studies. Here, we address the latter issue by using maximum likelihood and Bayesian methods to reconstruct the phylogenetic relationships among all 19 currently valid scyphozoan families, using sequence data from two nuclear genes: 18S and 28S rDNA. Consistent with prior morphological hypotheses, we find strong evidence for monophyly of subclass Discomedusae, order Coronatae, rhizostome suborder Kolpophorae and superfamilies Actinomyariae, Kampylomyariae, Krikomyariae, and Scapulatae. Eleven of the 19 currently recognized scyphozoan families are robustly monophyletic, and we suggest recognition of two new families pending further analyses. In contrast to long-standing morphological hypotheses, the phylogeny shows coronate family Nausithoidae, semaeostome family Cyaneidae, and rhizostome suborder Daktyliophorae to be nonmonophyletic. Our analyses neither strongly support nor strongly refute monophyly of order Rhizostomeae, superfamily Inscapulatae, and families Ulmaridae, Catostylidae, Lychnorhizidae, and Rhizostomatidae. These taxa, as well as familial relationships within Coronatae and within rhizostome superfamily Inscapulatae, remain unclear and may be resolved by additional genomic and taxonomic sampling. In addition to clarifying some historically difficult taxonomic questions and highlighting nodes in particular need of further attention, the molecular phylogeny presented here will facilitate more robust study of phenotypic evolution in the Scyphozoa, including the evolution characters associated with mass occurrences of jellyfish.     

Phylogeny of the millipede order Spirobolida (Arthropoda: Diplopoda: Helminthomorpha)

This study examines relationships within the millipede order Spirobolida using an exemplar approach, sampling within families to maximize geographical and morphological diversity; due to lack of available material, Allopocockiidae and Hoffmanobolidae were not included in analyses. The focus of this study was to test monophyly of the order, the suborders, and the families of Spirobolida and to propose interfamilial relationships using morphological and molecular data in a total‐evidence approach. Both maximum‐parsimony analyses and Bayesian inference were employed to analyse two datasets consisting of combined morphological and molecular data, one aligned using progressive alignment methods and the second aligned by secondary structure models. Rhinocricidae was recovered sister to all remaining spirobolidan millipedes and is elevated to suborder status as suborder Rhinocricidea. Trigoniulidea was recovered as monophyletic as was Spirobolidea excluding Rhinocricidae; Spirobolidea is redefined to reflect this change. All previously recognized families were recovered, with the exception of Spirobolidae; in all instances, this family was paraphyletic or part of a polytomy that lacked sufficient resolution to assess its monophyly. The results reaffirm much of the existing taxonomic foundation within Spirobolida. This study provides the first phylogenetic test of higher‐level relationships within Spirobolida and will serve as a foundation for future work in this group at finer levels. © The Willi Hennig Society 2010.     

Phylogenetic analysis with multiple markers indicates repeated loss of the adult medusa stage in Campanulariidae (Hydrozoa, Cnidaria)

The Campanulariidae is a group of leptomedusan hydroids (Hydrozoa, Cnidaria) that exhibit a diverse array of life cycles ranging from species with a free medusa stage to those with a reduced or absent medusa stage. Perhaps the best-known member of the taxon is Obelia which is often used as a textbook model of hydrozoan life history. However, Obelia medusae have several unique features leading to a hypothesis that Obelia arose, in a saltational fashion, from an ancestor that lacked a medusa, possibly representing an example of a rare evolutionary reversal. To address the evolution of adult sexual stages in Campanulariidae, a molecular phylogenetic approach was employed using two nuclear (18S rDNA and calmodulin) and two mitochondrial (16S rDNA and cytochrome c oxidase subunit I) genes. Prior to the main analysis, we conducted a preliminary analysis of leptomedusan taxa which suggests that Campanulariidae as presently considered needs to be redefined. Campanulariid analyses are consistent with morphological understanding in that three major clades are recovered. However, several recognized genera are not monophyletic calling into question some "diagnostic" features. Furthermore, ancestral states were reconstructed using parsimony, and a sensitivity analysis was conducted to investigate possible evolutionary transitions in life-history stages. The results indicate that life-cycle transitions have occurred multiple times, and that Obelia might be derived from an ancestor with Clytia-like features.     

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.].     

Taxonomic classification of the reef coral family Mussidae (Cnidaria: Anthozoa: Scleractinia)

Molecular analyses are transforming our understanding of the evolution of scleractinian corals and conflict with traditional classification, which is based on skeletal morphology. A new classification system, which integrates molecular and morphological data, is essential for documenting patterns of biodiversity and establishing priorities for marine conservation, as well as providing the morphological characters needed for linking present‐day corals with fossil species. The present monograph is the first in a series whose goal is to develop such an integrated system. It addresses the taxonomic relationships of 55 Recent zooxanthellate genera (one new) in seven families (one new), which were previously assigned to the suborder Faviina (eight genera are transferred to incertae sedis). The present monograph has two objectives. First, we introduce the higher‐level classification system for the 46 genera whose relationships are clear. Second, we formally revise the taxonomy of those corals belonging to the newly discovered family‐level clade (restricted today to the western Atlantic and Caribbean regions); this revised family Mussidae consists of ten genera (one of which is new) and 26 species that were previously assigned to the ‘traditional’ families Faviidae and Mussidae. To guide in discovering morphologic characters diagnostic of higher‐level taxa, we mapped a total of 38 morphologic characters [19 macromorphology, eight micromorphology, 11 microstructure] onto a molecular tree consisting of 67 species [22 Indo‐Pacific and seven Atlantic species in the traditional family Faviidae; 13 Indo‐Pacific and ten Atlantic species in the traditional family Mussidae; 13 species in the traditional families Merulinidae (5), Pectiniidae (7), and Trachyphylliidae (1); two Atlantic species of traditional Montastraea], and trace character histories using parsimony. To evaluate the overall effectiveness of morphological data in phylogeny reconstruction, we performed morphology‐based phylogenetic analyses using 27 (80 states) of the 38 characters, and compared morphological trees with molecular trees. The results of the ancestral state reconstructions revealed extensive homoplasy in almost all morphological characters. Family‐ and subfamily‐level molecular clades [previously identified as XVII?XXI] are best distinguished on the basis of the shapes of septal teeth and corresponding microstructure. The newly revised family Mussidae (XXI) has septal teeth with regular pointed tips (a symplesiomorphy) and a stout blocky appearance. It has two subfamilies, Mussinae and Faviinae. The subfamily Mussinae is distinguished by spine‐shaped teeth and widely spaced costoseptal clusters of calcification centres. The subfamily Faviinae is distinguished by blocky, pointed tricorne or paddle‐shaped teeth with elliptical bases, transverse structures such as carinae that cross the septal plane, and well‐developed aligned granules. Defining diagnostic characters for the broader data set is more challenging. In analyses of taxonomic subsets of the data set that were defined by clade, morphological phylogenetic analyses clearly distinguished the families Mussidae (XXI) and Lobophylliidae (XIX), as well as the two subfamilies of Mussidae (Mussinae, Faviinae), with one exception (Homophyllia australis). However, analyses of the entire 67‐species data set distinguished the family Lobophylliidae (XIX), but not the Merulinidae (XVII) and not the newly defined Mussidae (XXI), although the subfamily Mussinae was recovered as monophyletic. Some lower‐level relationships within the Merulinidae (XVII) agree with molecular results, but this particular family is especially problematic and requires additional molecular and morphological study. Future work including fossils will not only allow estimation of divergence times but also facilitate examination of the relationship between these divergences and changes in the environment and biogeography. Published 2012. This article is a U.S. Government work and is in the public domain in the USA. Zoological Journal of the Linnean Society, 2012, 166 , 465–529.     

DNA analyses reveal abundant homoplasy in taxonomically important morphological characters of Eusiroidea (Crustacea,Amphipoda)

Eusiroidea is one of the 20 amphipod superfamilies that were erected to subdivide the very large and controversial suborder Gammaridea. Yet, the definition of the superfamily is not based on synapomorphies, but on a combination of diagnostic phenetic similarities that hold more or less consistently across families. Moreover, many of the characters used to define eusiroid families are suspected to show convergent evolution. The current classification of the Eusiroidea may therefore not reflect evolutionary relationships accurately. Here, we present a molecular phylogenetic re‐analysis of the Eusiroidea based on a comparison of 18S and 28S rDNA sequences of 73 species, representing 47 genera and 16 families that potentially belong to the superfamily. The results suggest that at least species belonging to 14 of these traditional families would be part of a eusiroid clade, increasing by more than twofold the species and generic richness of the group. However, most of the eusiroid families surveyed do not appear monophyletic. Finally, the analyses show that several important morphological characteristics, traditionally used in eusiroid taxonomy, are homoplastic.     

Molecular systematics of the bark lice infraorder Caeciliusetae (Insecta: Psocodea)

The phylogenetic relationships of bark lice and parasitic lice (Insecta: Psocodea) have been studied in a number of recent molecular phylogenetic analyses based on DNA sequences. Many of these studies have focused on the position of parasitic lice within the free‐living bark lice. However, fewer such studies have examined the relationships among major groups of free‐living bark lice and their implications for classification. In this study we focus on the infraorder Caeciliusetae, a large group of bark lice (?1000 species) within the suborder Psocomorpha. Using sequences of two mitochondrial and two nuclear genes, we estimated the phylogeny for relationships among the five recognized families within the infraorder Caeciliusetae. Based on the results, the sister‐group relationship and respective monophyly of Stenopsocidae and Dasydemellidae is strongly supported. Monophyly of the larger families Amphipsocidae and Caeciliusidae was not supported, although the causes of this were the placement of two distinct subfamilies (Paracaeciliinae and Calocaeciliinae). The monophyly of Asiopsocidae could not be tested because it was sampled only by one species. Based on these results and consideration of morphological characters, we propose a new classification for Caeciliusetae, recognizing six families: Amphipsocidae, Stenopsocidae, Dasydemellidae, Asiopsocidae, Paracaeciliidae and Caeciliusidae. We expect that this new classification will stabilize the higher‐level taxonomy of this group and help to identify groups in need of further work among these insects.     

Phylogeny of the suborder Psocomorpha: congruence and incongruence between morphology and molecular data (Insecta: Psocodea: ‘Psocoptera’)

The largest suborder of bark lice (Insecta: Psocodea: ‘Psocoptera’) is Psocomorpha, which includes over 3600 described species. We estimated the phylogeny of this major group with family‐level taxon sampling using multiple gene markers, including both nuclear and mitochondrial ribosomal RNA and protein‐coding genes. Monophyly of the suborder was strongly supported, and monophyly of three of four previously recognized infraorders (Caeciliusetae, Epipsocetae, and Psocetae) was also strongly supported. In contrast, monophyly of the infraorder Homilopsocidea was not supported. Based on the phylogeny, we divided Homilopsocidea into three independent infraorders: Archipsocetae, Philotarsetae, and Homilopsocidea. Except for a few cases, previously recognized families were recovered as monophyletic. To establish a classification more congruent with the phylogeny, we synonymized the families Bryopsocidae (with Zelandopsocinae of Pseudocaeciliidae), Calopsocidae (with Pseudocaeciliidae), and Neurostigmatidae (with Epipsocidae). Monophyly of Elipsocidae, Lachesillidae, and Mesopsocidae was not supported, but the monophyly of these families could not be rejected statistically, so they are tentatively maintained as valid families. The molecular tree was compared with a morphological phylogeny estimated previously. Sources of congruence and incongruence exist and the utility of the morphological data for phylogenetic estimation is evaluated. © 2014 The Linnean Society of London     

Polycladida phylogeny and evolution: integrating evidence from 28S rDNA and morphology

Polyclad flatworms have a troubled classification history, with two contradicting systems in use. They both rely on a ventral adhesive structure to define the suborders Acotylea and Cotylea, but superfamilies were defined according to eyespot arrangement (Prudhoe’s system) or prostatic vesicle characters (Faubel’s system). Molecular data available cover a very limited part of the known polyclad family diversity and have not allowed testing morphology-based classification systems on Polycladida yet. We thus sampled a suitable marker, partial 28S ribosomal DNA (rDNA), from Polycladida (19 families and 32 genera), generating 136 new sequences and the first comprehensive genetic dataset on polyclads. Our maximum likelihood (ML) analyses recovered Polycladida, but the traditional suborders were not monophyletic, as the supposedly acotyleans Cestoplana and Theama were nested within Cotylea; we suggest that these genera should be included in Cotylea. The partial 28S rDNA trees were generally well supported and robust but in conflict with both Faubel’s and Prudhoe’s superfamilies. Therefore, we compiled morphological and anatomical characters for all taxa used and examined their distribution on our molecular tree. Combining morphological and molecular evidence, we redefined polyclad superfamilies. Acotylea contain tentaculated and atentaculated groups and is now divided in three superfamilies. The suborder Cotylea can be divided in five superfamilies. In general, there is a trait of anteriorization of sensory structures, from the plesiomorphic acotylean body plan to the cotylean gross morphology. Traditionally used characters, such as prostatic vesicle, eyespot distribution, and type of pharynx, are all homoplastic and likely have misled polyclad systematics so far.     

Phylogeny and morphological evolution of the amblystegiaceae (Bryopsida)

To circumscribe the moss family Amblystegiaceae, we performed a broad-scale analysis of trnL-trnF spacer sequence data for 168 species of the Hypnales and 11 species of the Hookeriales and additional analyses of trnL-trnF and atpB-rbcL (chloroplast DNA), one nuclear region, the internal transcribed spacers of 18S-26S rDNA, and 68 morphological characters for a reduced data set of 54 species of Hypnales. The traditionally circumscribed Amblystegiaceae are polyphyletic and include the Amblystegiaceae s. str. and the Calliergonaceae fam. nov., plus several taxa closely related to other Hypnalean families. Generic relationships within the redefined Amblystegiaceae were investigated by analyzing data from the three DNA regions and morphology as used in the broader analysis. Reconstruction of morphological evolution was evaluated using maximum-parsimony and maximum-likelihood. Numerous independent character-state transitions implied by the phylogeny suggest that morphological characters that have traditionally been used to delineate the Amblystegiaceae are homoplastic. Sporophytic traits, which are generally given primacy over gametophytic traits in moss classification, are more labile than previously thought, and many characters that are related to sporophyte specializations are strongly correlated with habitat conditions. The evolution of several gametophyte features previously thought to be reliable for delineating the family are also strongly correlated with habitat. These observations help to explain the instability of the Amblystegiaceae in previous taxonomic and phylogenetic analyses based on morphology.     

Character evolution in Hydrozoa (phylum Cnidaria)

The diversity of hydrozoan life cycles, as manifested in the wide range of polyp, colony, and medusa morphologies, has been appreciated for centuries. Unraveling the complex history of characters involved in this diversity is critical for understanding the processes driving hydrozoan evolution. In this study, we use a phylogenetic approach to investigate the evolution of morphological characters in Hydrozoa. A molecular phylogeny is reconstructed using ribosomal DNA sequence data. Several characters involving polyp, colony, and medusa morphology are coded in the terminal taxa. These characters are mapped onto the phylogeny and then the ancestral character states are reconstructed. This study confirms the complex evolutionary history of hydrozoan morphological characters. Many of the characters involving polyp, colony, and medusa morphology appear as synapomorphies for major hydrozoan clades, yet homoplasy is commonplace.