Christopher L. Mah

Global Diversity and Phylogeny of the Asteroidea (Echinodermata)

Members of the Asteroidea (phylum Echinodermata), popularly known as starfish or sea stars, are ecologically important and diverse members of marine ecosystems in all of the worlds oceans. We present a comprehensive overview of diversity and phylogeny as they have figured into the evolution of the Asteroidea from Paleozoic to the living fauna. Living post-Paleozoic asteroids, the Neoasteroidea, are morphologically separate from those in the Paleozoic. Early Paleozoic asteroid faunas were diverse and displayed morphology that foreshadowed later living taxa. Preservation presents significant difficulties, but fossil occurrence and current accounts suggests a diverse Paleozoic fauna, which underwent extinction around the Permian-Triassic interval was followed by re-diversification of at least one surviving lineage. Ongoing phylogenetic classification debates include the status of the Paxillosida and the Concentricycloidea. Fossil and molecular evidence has been and continues to be part of the ongoing evolution of asteroid phylogenetic research. The modern lineages of asteroids include the Valvatacea, the Forcipulatacea, the Spinlosida, and the Velatida. We present an overview of diversity in these taxa, as well as brief notes on broader significance, ecology, and functional morphology of each. Although much asteroid taxonomy is stable, many new taxa remain to be discovered with many new species currently awaiting description. The Goniasteridae is currently one of the most diverse families within the Asteroidea. New data from molecular phylogenetics and the advent of global biodiversity databases, such as the World Asteroidea Database (http://www.marinespecies.org/Asteroidea/) present important new springboards for understanding the global biodiversity and evolution of asteroids.

Biodiversity and phylogeography of Arctic marine fauna: insights from molecular tools

The last decade has seen an increase in the frequency and breadth of application of molecular tools, many of which are beginning to shed light on long-standing questions in biogeography and evolutionary history of marine fauna. We explore new developments with respect to Arctic marine invertebrates, focusing on molecular taxonomy and phylogeography—two areas that have seen the most progress in the time-frame of the Census of Marine Life. International efforts to generate genetic ‘barcodes’ have yielded new taxonomic insights and applications ranging from diet analysis to identification of larval forms. Increasing availability of genetic data in public databases is also facilitating exploration of large-scale patterns in Arctic marine populations. We present new case-studies in meta-population analysis of barcode data from polychaetes and echinoderms that demonstrate such phylogeographic applications. Emerging patterns from ours and other published studies include influences of a complex climatic and glacial history on genetic diversity and evolution in the Arctic, and contrasting patterns of both high gene flow and persistent biogeographic boundaries in contemporary populations.

The Mg-Calcite Composition of Antarctic Echinoderms: Important Implications for Predicting the Impacts of Ocean Acidification

The Southern Ocean is considered to be the canary in the coal mine with respect to the first effects of ocean acidification (OA). This vulnerability is due to naturally low carbonate ion concentrations that result from the effect of low temperature on acid-base dissociation coefficients, from the high solubility of CO2 at low temperature, and from ocean mixing. Consequently, the two calcium carbonate polymorphs, aragonite and calcite, are expected to become undersaturated in the Southern Ocean within 50 and 100 years, respectively. Marine invertebrates such as echinoderms, whose skeletons are classified as high-magnesium carbonate (>4% mol MgCO3), are even more vulnerable to OA than organisms whose skeletons consist primarily of aragonite or calcite, with respect to both increased susceptibility to skeletal dissolution and further challenge to their production of skeletal elements. Currently, despite their critical importance to predicting the effects of OA, there is almost no information on the Mg-calcite composition of Antarctic echinoderms, a group known to be a major contributor to the global marine carbon cycle. Here we report the Mg-calcite compositions of 26 species of Antarctic echinoderms, representing four classes. As seen in tropical and temperate echinoderms, Mg-calcite levels varied with taxonomic class, with sea stars generally having the highest levels. When combined with published data for echinoderms from primarily temperate and tropical latitudes, our findings support the hypothesis that Mg-calcite level varies inversely with latitude. Sea stars and brittle stars, key players in Antarctic benthic communities, are likely to be the first echinoderms to be challenged by near-term OA.

Systematics, phylogeny and historical biogeography of the Pentagonaster clade (Asteroidea: Valvatida: Goniasteridae)

Morphology-based phylogenetic hypotheses developed for living and fossil goniasterid asteroids have provided several unique opportunities to study bathymetric and biogeographic shifts for an ecologically important group of prominent, megafaunal invertebrates. A cladistic analysis of 18 ingroup taxa employing 65 morphological characters resulted in a single most parsimonious tree. The tree supports assignment of the Atlantic Tosia parva (Perrier, 1881) and the Pacific Tosia queenslandensis Livingstone, 1932 to new, separate genera. The phylogenetic tree supports offshore to onshore bathymetric shifts between basal and derived taxa. The phylogeny is also consistent with historical events surrounding the separation of Antarctica from Australia and South Africa. Buterminaster Blake & Zinsmeister, 1988 from the Eocene La Meseta Formation, Antarctic Peninsula, was included in the phylogenetic analysis and is now supported as the only fossil species in the genus Pentagonaster Gray, 1840. Pentagonaster stibarus H. L. Clark, 1914 is separated from synonymy with P. dubeni Gray, 1847 and resurrected as a valid species. The new genus, Akelbaster, gen. nov., shows unusual new structures that resemble cribiform organs, although their function has not been determined. One specific ingroup lineage, including Tosia and Pentagonaster, attains a much larger adult size than those of its sister-taxa, suggesting that Cope’s rule may apply to asteroids within this clade. Pentagonaster and related genera are revised. Descriptions of four new genera and three new species are presented, including: Akelbaster novaecaledoniae, gen. nov., sp. nov., Ryukuaster onnae, gen. nov., sp. nov., Eknomiaster beccae, sp. nov., Pawsonaster parvus, gen. nov., comb. nov. and Anchitosia queenslandensis, gen. nov., comb. nov.

Multiple processes generate productivity-diversity relationships in experimental wood-fall communities.

Energy availability has long been recognized as a predictor of community structure, and changes in both terrestrial and marine productivity under climate change necessitate a deeper understanding of this relationship. The productivity-diversity relationship (PDR) is well explored in both empirical and theoretical work in ecology, but numerous questions remain. Here, we test four different theories for PDRs (More-Individuals Hypothesis, Resource-Ratio Theory, More Specialization Theory, and the Connectivity-Diversity Hypothesis) with experimental deep-sea wood falls. We manipulated productivity by altering wood-fall sizes and measured responses after 5 and 7 years. In November 2006, 32 Acacia sp. logs were deployed at 3203 m in the Northeast Pacific Ocean (Station Deadwood: 36.154098 degrees N, 122.40852 degrees W). Overall, we found a significant increase in diversity with increased wood-fall size for these communities. Increases in diversity with wood-fall size occurred because of the addition of rare species and increases of overall abundance, although individual species responses varied. We also found that limited dispersal helped maintain the positive PDR relationship. Our experiment suggests that multiple interacting mechanisms influence PDRs.

Abyssal fauna of the UK-1 polymetallic nodule exploration claim, Clarion-Clipperton Zone, central Pacific Ocean: Echinodermata

Abstract We present data from a DNA taxonomy register of the abyssal benthic Echinodermata collected as part of the Abyssal Baseline (ABYSSLINE) environmental survey cruise ‘AB01’ to the UK Seabed Resources Ltd (UKSRL) polymetallic-nodule exploration claim ‘UK-1’ in the eastern Clarion-Clipperton Zone (CCZ), central Pacific Ocean abyssal plain. Morphological and genetic data are presented for 17 species (4 Asteroidea, 4 Crinoidea, 2 Holothuroidea and 7 Ophiuroidea) identified by a combination of morphological and genetic data. No taxa matched previously published genetic sequences, but 8 taxa could be assigned to previously-described species based on morphology, although here we have used a precautionary approach in taxon assignments to avoid over-estimating species ranges. The Clarion-Clipperton Zone is a region undergoing intense exploration for potential deep-sea mineral extraction. We present these data to facilitate future taxonomic and environmental impact study by making both data and voucher materials available through curated and accessible biological collections.

From the Arctic to the Antarctic: the major, minor, and trace elemental composition of echinoderm skeletons

Biogenic carbonate production in benthic marine ecosystems is dominated by representatives of the Echinodermata. Carbon and other major, minor, and trace elements are exported to the seabed where they accumulate or dissolve. Preserved carbonates (Mg-calcite) have applications in oceanography and geochemistry and are used to reconstruct various parameters of ancient seawater, such as temperature (from Mg/Ca, Sr/Ca), seawater Mg/Ca (from Mg/Ca), and pH (from B/Ca). In general, the benthos is widely ignored for its role in the global carbon cycle despite the importance of echinoderms as a carbon sink (∼0.1–0.2 Pg C/yr). Echinoderms produce their skeletons from Mg-calcite, which is more soluble than pure calcite and, therefore, more vulnerable to ocean acidification (OA). Minor and trace elements can also destabilize the calcite lattice, increasing the minerals solubility. Little is known about the concentration of such elements in echinoderm tests. Expanding our knowledge on echinoderm skeleton composition will improve our understanding of elemental flux in the oceans. Furthermore, establishing relationships between the physical parameters of seawater and minor/trace elemental ratios within echinoderm Mg-calcite should expand the utility of fossils as geochemical archives. Herein, we present elemental composition data for Asteroidea (n = 108; 9 families, 23 species), Echinoidea (n = 94; 8 families, 12 species), Ophiuroidea (n = 24; 4 families, 5 species), Holothuroidea (n = 7; 3 families, 3 species), and Crinoidea (n = 3; 1 family, 1 species), collected from the Arctic to the Antarctic oceans, from depths ranging from surface waters to 1200 m. The following elements were measured and normalized to [Ca]: Li, Be, Mg, Al, P, S, K, Sc, Ti, V, Cr, Mn, Fe, Co, Ni, Cu, Zn, Ga, Ge, As, Se, Rb, Sr, Y, Zr, Mo, Ag, Cd, Sn, Sb, Te, Ba, La, Ce, Nd, Dy, W, Re, Au, Hg, Tl, Pb, Bi, and U. Data are presented for the whole body, arms (plates), calcareous ossicles, spines, and test plates. Elements were quantified using inductively coupled plasma mass spectrometry. Our study presents the most comprehensive data set to date for a phylum whose skeletons are composed of Mg-calcite.

Comments on "The phylogeny of post-Palaeozoic Asteroidea (Neoasteroidea, Echinodermata)" by A.S. Gale and perspectives on the systematics of the Asteroidea

A new, morphologically based classification of extant asteroids with comments on select fossils was published by A.S. Gale. Research approaches used limited sampling, and much literature treatment is not accurate and therefore misleading. We review these concerns, seeking to clarify argumentation on differing interpretations.

Difference in larval type explains patterns of nonsynonymous substitutions in two ancient paralogs of the histone H3 gene in sea stars

SUMMARY Paralogous genes frequently show differences in patterns and rates of substitution that are typically attributed to different selection regimes, mutation rates, or local recombination rates. Here, two anciently diverged paralogous copies of the histone H3 gene in sea stars, the tandem‐repetitive early‐stage gene and a newly isolated gene with lower copy number that was termed the “putative late‐stage histone H3 gene” were analyzed in 69 species with varying mode of larval development. The two genes showed differences in relative copy number, overall substitution rates, nucleotide composition, and codon usage, but similar patterns of relative nonsynonymous substitution rates, when analyzed by the dN/dS ratio. Sea stars with a nonpelagic and nonfeeding larval type (i.e., brooding lineages) were observed to have dN/dS ratios that were larger than for nonbrooders but equal between the two paralogs. This finding suggested that demographic differences between brooding and nonbrooding lineages were responsible for the elevated dN/dS ratios observed for brooders and refuted a suggestion from a previous analysis of the early‐stage gene that the excess nonsynonymous substitutions were due to either (1) gene expression differences at the larval stage between brooders and nonbrooders or (2) the highly repetitive structure of the early‐stage histone H3 gene.

Recent relaxation of purifying selection on the tandem-repetitive early-stage histone H3 gene in brooding sea stars.

Patterns of nucleotide substitution differ between marine species that have a pelagic feeding (planktotrophic) larval stage and related species that lack such a stage, for both adaptive and non-adaptive reasons. Here, patterns of nucleotide and inferred amino acid substitution are analyzed for the tandem-repetitive early-stage histone H3 gene in 36 sea star species of the order Forcipulatida with documented larval habitat. The relative rate of nonsynonymous substitution (expressed as ω=d(N)/d(S)) was significantly higher in lineages with a brooded non-feeding (lecithotrophic) larval form than in lineages with a planktotrophic larval form. There was also a significant excess of conservative over radical substitutions. The increase in ω for brooders as compared to non-brooders was much greater than for previously analyzed mitochondrial sequences in echinoderms. These data are consistent with the hypothesis that purifying selection on this gene has been relaxed in brooding lineages compared to non-brooding lineages. The hypotheses of adaptive or neutral evolution are less plausible, although recent pseudogenization following a period of relaxed purifying selection could also explain the results.