Reconstruction of the phylogeny of the five extant classes of the phylum Echinodermata has proven difficult. Results concerning higher-level taxonomic relationships among echinoderms are sensitive to the choice of analytical parameters and methods. Moreover, the proposal of a putative sixth class based on a small enigmatic disc-shaped echinoderm, Xyloplax, from the deep seas of the Bahamas and New Zealand in the 1980s further complicated the problem. Although clearly an echinoderm, Xyloplax did not have clear affinity among known groups. Using molecular sequence and developmental data from recently collected Xyloplax adults and embryos, we show that rather than representing an ancient distinct lineage as implied by its status as a class, Xyloplax is simply a starfish that is closely related to the asteroid family Pterasteridae. Many members of the Pterasteridae and all Xyloplax inhabit deep or polar seas and brood young. Brooding pterasterids and Xyloplax hold their young in specialized adult chambers until the young reach an advanced juvenile stage after which they are released as free-living individuals. We hypothesize that the unique morphology of Xyloplax evolved via progenesis--the truncation of somatic growth at a juvenile body plan but with gonadal growth to maturity. Although the overall phylogeny of extant echinoderms remains sensitive to the choice of analytical methods, the placement of Xyloplax as sister to pterasterid asteroids is unequivocal. Based on this, we argue that the proposed class and infraclass status of Xyloplax should be suppressed.

Many questions in evolutionary biology are best addressed by comparing traits in different species. Often such studies involve mapping characters on phylogenetic trees. Mapping characters on trees allows the nature, number, and timing of the transformations to be identified. The parsimony method is the only method available for mapping morphological characters on phylogenies. Although the parsimony method often makes reasonable reconstructions of the history of a character, it has a number of limitations. These limitations include the inability to consider more than a single change along a branch on a tree and the uncoupling of evolutionary time from amount of character change. We extended a method described by Nielsen (2002, Syst. Biol. 51:729-739) to the mapping of morphological characters under continuous-time Markov models and demonstrate here the utility of the method for mapping characters on trees and for identifying character correlation.

The sea star, Asterina minor, was surveyed for Hox genes using the method of PCR and subsequent sequence determination. Seven different Hox-type homeobox fragments and homeobox fragments of two other types, the Gbx-type and the Xlox-type, were identified. The results of comparative analysis with known homeobox sequences suggest that the sea star has only one Hox gene cluster including two genes of the anterior group, four genes of the medial group, and one gene of the posterior group. The existence of a gene of the cognate group 1 has not been known in echinoderm species. Each of the other fragments indicated a definite relationship with one of sea urchin homeoboxes. The hypothetical cluster in the sea star is consistent with the results published for another class of echinoderm, sea urchins, in the putative number of cluster. The present result provides strong evidence that a single Hox cluster is common to echinoderms and its structure in the anterior region is more similar to other deuterostomes than previously thought.

To help elucidate the cluster organization of Hox genes in echinoderms, we amplified a homeobox region by polymerase chain reaction (PCR) and cloned and sequenced the PCR products for the comatulid crinoid Oxycomanthus japonicus and the ophiuroid Stegophiura sladeni. The crinoid had at least three anterior, four medial, and four posterior genes, and the ophiuroid had at least one anterior, three medial, and six (one of which being a possible trans-paralog) posterior genes. The survey of the crinoid detected all three anterior complements (PG1-3 genes). It was inferred that the Hox genes of each species are organized into a single cluster and that a novel cognate group of echinoderm posterior genes (tentatively termed HboxP9) exists among echinoderms in addition to the known posterior genes Hbox4, Hbox7, and Hbox10. The results, combined with the data of other echinoderm classes, strongly suggest that the presence of a single Hox gene cluster is a common feature among echinoderms and that the cluster has the general features of the deuterostome Hox gene cluster, i.e., the anterior conservation and posterior expansion. The results of the ophiuroid imply that the posterior genes in this class diversified after the phylum Echinodermata had been established.

Asteroids display four distinct modes of developmental patterns: the indirect mode, the nonbrachiolarian mode, the direct mode, and the mode with a barrel-shaped larva. Among them the former two are planktotrophic, whereas the latter two are lecithotrophic. The direct mode and the mode with a barrel-shaped larva are thought to have evolved from the more primitive planktotrophic mode, the nonbrachiolarian and the indirect mode, respectively. However, whether the nonbrachiolarian mode or the indirect mode is the more primitive in asteroids is unresolved, despite discussion since early this century. A key aspect of this problem is the phylogenetic status of paxillosidans, since the nonbrachiolarian mode and the mode with a barrel-shaped larva are seen only in paxillosidans. To resolve this problem, we performed a molecular phylogenetic study of asteroids, based on the nucleotide sequences of mitochondrial rDNAs. Phylogenetic trees support a close relationship between the Asterinidae and the Solasteridae. We suggest that the paxillosidans are not a monophyletic group; rather, the Luidiidae (one family of Paxillosida) is a sister group to the rest of the asteroids. Although some aspects of our results contradict a recent study by Lafay et al.(1995, Syst. Biol. 44: 190-208) based on 28S rRNA sequences, both studies agree on a paraphyletic nature for the paxillosidans. We conclude that characters shared by paxillosidans are primitive; hence the primitive mode of development in asteroids is the nonbrachiolarian mode.

The organization of the peptidergic system in the larvae of Patiriella species with divergent ontogenies was compared to determine which aspects of neurogenesis are conserved and which are altered in the evolution of development in these sea stars. P. regularis has ancestral-type feeding bipinnaria and brachiolaria larvae and the organization of the nervous system, in association with feeding structures, paralleled the bilateral larval body plan. P. calcar and P. exigua have non-feeding planktonic and benthic brachiolariae, respectively, and there was no trace of the neuronal architecture involved with feeding. The nervous system in the attachment stage brachiolaria was similar in all three species and neuronal organization reflected larval symmetry. Delayed expression of peptidergic lineages to the brachiolaria stage in the lecithotrophs indicates heterochronic change in the timing of neurogenesis or deletion of the ancestral early neurogenic program. The bipinnarial program is suggested to be a developmental module autonomous from the brachiolar one. With a divergence time of less than 10 Ma, the evolution of development in Patiriella has resulted in extensive reduction in the complexity of the larval nervous system in parallel with simplification in larval form. There is, however, strong conservation in the morphology and neuronal architecture of structures involved with settlement.

Oocyte maturation (meiosis reinitiation) in starfish is induced by the natural hormone 1-methyladenine. In some species (group 2) oocyte maturation can be induced by micromolar concentrations of a few fatty acids such as arachidonic and eicosapentaenoic acids or by nanomolar concentrations of hydroxyeicosatetraenoic acid. Complete maturation is triggered: increased protein phosphorylation, appearance of the cytoplasmic "maturation-promoting factor", germinal vesicle breakdown, emission of the two polar bodies and formation of the female pronucleus. In other species (group 1), however, no maturation can be induced by the fatty acids active in the species of group 2, despite a large variety of experimental conditions.

Reconstruction of the phylogeny of the five extant classes of the phylum Echinodermata has proven difficult. Results concerning higher-level taxonomic relationships among echinoderms are sensitive to the choice of analytical parameters and methods. Moreover, the proposal of a putative sixth class based on a small enigmatic disc-shaped echinoderm, Xyloplax, from the deep seas of the Bahamas and New Zealand in the 1980s further complicated the problem. Although clearly an echinoderm, Xyloplax did not have clear affinity among known groups. Using molecular sequence and developmental data from recently collected Xyloplax adults and embryos, we show that rather than representing an ancient distinct lineage as implied by its status as a class, Xyloplax is simply a starfish that is closely related to the asteroid family Pterasteridae. Many members of the Pterasteridae and all Xyloplax inhabit deep or polar seas and brood young. Brooding pterasterids and Xyloplax hold their young in specialized adult chambers until the young reach an advanced juvenile stage after which they are released as free-living individuals. We hypothesize that the unique morphology of Xyloplax evolved via progenesis--the truncation of somatic growth at a juvenile body plan but with gonadal growth to maturity. Although the overall phylogeny of extant echinoderms remains sensitive to the choice of analytical methods, the placement of Xyloplax as sister to pterasterid asteroids is unequivocal. Based on this, we argue that the proposed class and infraclass status of Xyloplax should be suppressed.

A 551-bp region of a PCR product containing the putative mitochondrial control region and flanking sequences was analyzed for sequence variation among 19 sea stars representing 10 previously described PCR-RFLP haplotypes within a cryptic species complex (Leptasterias spp.). Most (97%) of the sequence variation was interhaplotypic rather than intrahaplotypic, which greatly reduced the utility of sequence polymorphisms in this mtDNA region as markers of intrahaplotypic population structure and gene flow. The estimated number of transition and transversion substitutions per nucleotide site, corrected for multiple hits, was 0.0364 and 0.0158, respectively. Most of the sequence variation occurred in the first half of the putative control region. Phylogenetic analysis (both maximum parsimony and maximum likelihood) revealed three well-supported clades, but the position of two PCR-RFLP haplotypes was not completely resolved. Low intraspecific mtDNA sequence divergence over large geographic distances may be a general pattern for echinoderm species.

Although the sex ratios of many groups conform to Fisher's (1930) prediction that parents should invest equally in daughters and sons, a number of taxa are characterized by excesses of one gender. A variety of mechanisms may lead to sex ratio biases, but in organisms that reproduce clonally as well as sexually, gender differences in the rate of cloning could drive the development of sex-ratio biases. In this study, I demonstrate that males of the clonal brittle star Ophiactis savignyi were significantly more likely to divide than females and that the magnitude of this difference was sufficient to explain the consistent and significant excess of males in natural populations. Females were significantly more likely to lose sexual reproductive capabilities following division, and this greater cost associated with division may explain why females are less likely to divide. Gender differences in mortality rates are unlikely to explain the excess of males in this species. Because of their potential influence on the operational sex ratio, gender differences in division rates may have important ecological and evolutionary implications including effects on the direction and strength of selection.