taxonID	type	description	language	source
03E687F8FFA3FFA73C9C2441FECE594E.taxon	description	Our trees recover a non-monophyletic Pedicellasteridae, which places putative pedicellasterids on three different clades: the Pedicellasteridae; the Ampheraster clade, including new genus 2; and the six-rayed pedicellasterid clade (new genus 3). The Pedicellasteridae clade includes the name-bearing Pedicellaster, Hydrasterias, and a new genus and species of pedicellasterid (new genus 1). The second ‘ pedicellasterid’ clade contains the pedicellasterids Tarsaster alaskanus, Ampheraster marianus, the Hawaiian asteriid Tarsastrocles, and a new pedicellasterid genus and species 2. Finally, the third clade includes the six-rayed new genus / species 3, which was collected from the Gorda Ridge region and adjacent areas in the North Pacific. The genus Tarsaster is not supported as monophyletic, with Tarsaster galapagensis recovered among the Stichasteridae and Tarsaster alaskanus placed with Ampheraster, Tarsastrocles, and new genus / species 1. Although Tarsaster has historical seniority over Ampheraster, the lack of monophyly in Tarsaster has dictated that for the time being, Ampheraster be used to describe the non-Stichasteridae clade, which includes Tarsaster alaskanus, until taxonomic issues in Tarsaster can be fully resolved. Depending on whether the type species for Tarsaster, Tarsaster stoichodes is placed among the Stichasteridae or among the members of the ‘ Ampheraster ’ clade, assignments to new genera will probably be necessary. Although not recovered as monophyletic, pedicellasterids were never recovered as part of the restricted Asteriidae or any of the more recently derived forcipulatacean lineages, supporting the interpretation of at least some pedicellasterids as basal or stemward forcipulataceans. Although the multiple pedicellasterid lineages suggest that the characters associated with pedicellasterids are independently derived, it seems more likely, especially based on phylogenetic hypotheses based on fossils (e. g. Blake & Hagdorn, 2003), that these characters are plesiomorphic for the Forcipulatacea.	en	Mah, Christopher, Foltz, David (2011): Molecular phylogeny of the Forcipulatacea (Asteroidea: Echinodermata): systematics and biogeography. Zoological Journal of the Linnean Society 162 (3): 646-660, DOI: 10.1111/j.1096-3642.2010.00688.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00688.x
03E687F8FFA0FFA73F9A260DFAE95B8B.taxon	description	Our results uphold Stichaster as monophyletic, but not Cosmasterias. Cosmasterias dyscrita was separated from a larger clade of Stichasteridae in both trees. In the three-gene tree, Cosmasterias lurida is a member of the sister group to the large clade containing C. dyscrita. In the two-gene tree, C. lurida is supported as part of the same clade as C. dyscrita. Based on the position of Neomorphaster within the stichasterid clade, the monotypic Neomorphasteridae Fisher, 1923 should be synonymized with the Stichasteridae Perrier, 1885. Our results do not support Neomorphaster as the sister taxon to Zoroaster, as indicated by Mah (2000). However, the shared morphological similarities may be indicative of plesiomorphic character states between zoroasterids and stichasterids. Sampled members of the Stichasteridae lineage are limited to temperate waters in the Southern Hemisphere, with two exceptions: the deep-water Neomorphaster occurring in the Northern Hemisphere in the North Atlantic (none have been reported from the South Atlantic), and Neosmilaster sp. nov. (supported as the sister to Neosmilaster steineni) from the North Pacific. The unsampled North Atlantic Stichasterella was placed within the Stichasteridae by Mortensen (1927), based on several shared morphological characters. If Stichasterella is upheld as a member of this clade, it would also be included among the Northern Hemisphere Stichasteridae. There is evidence that the stichasterid lineage has experienced some extinction and range restriction. Blake & Peterson (1993) described the Neomorphaster - like Pegaster from the Cretaceous of California. A significant level of forcipulatacean diversity that is restricted to the Australia / New Zealand / South Pacific region includes members of the stichasterid clade, including Stichaster, Pseudechinaster, Allostichaster, Smilasterias, and ‘ Cosmasterias ’ dyscrita (a separate lineage from C. lurida). Only the shallow-water South Australian Uniophora was absent from our sampling. However, Uniophora shares several morphological characters with other stichasterids, and therefore we suggest it will probably be included as a member of the Stichasteridae in future molecular phylogenetic studies.	en	Mah, Christopher, Foltz, David (2011): Molecular phylogeny of the Forcipulatacea (Asteroidea: Echinodermata): systematics and biogeography. Zoological Journal of the Linnean Society 162 (3): 646-660, DOI: 10.1111/j.1096-3642.2010.00688.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00688.x
03E687F8FFA0FFA63C3E2683FDDA58DD.taxon	description	In our two-gene tree (Fig. 2) the monotypic Heliasteridae, represented by the Eastern Pacific species Heliaster kubiniji, is supported (100 % bootstrap) as the sister taxon to the Southern Ocean / sub-Antarctic Labidiaster. The three-gene tree, which lacks Heliaster, shows Labidiaster as sister taxa to a new genus and species of deep-sea six-rayed pedicellasterid from the Central Pacific (86 % bootstrap support), but with much longer branch lengths, suggesting greater divergence between the two taxa. If Labidiaster and Heliaster are sister taxa, this would further suggest a close biogeographic relationship between the South American and Antarctic / sub-Antarctic asteroid faunas. Janosik et al. (2008) have shown pelagic larvae for Labidiaster annulatus present in the Drake Passage, suggesting gene flow between South American and Antarctic populations. Support for Labidiaster as the sister taxon to Heliaster is consistent with the assertion that the Labidiasteridae (sensu Spencer & Wright, 1966) is a purely artificial grouping (e. g. Mah, 2000; Foltz et al., 2007). Labidiaster is a phylogenetically separate taxon from the other labidiasterids (sensu Clark & Mah, 2001), including Plazaster, Coronaster, and Rathbunaster. Because Labidiaster is the type genus for the Labidiasteridae, this places synonymy of the Labidiasteridae into the Heliasteridae rather than the Asteriidae, as has been implied by earlier studies (e. g. Mah, 2000). The other genera within the Labidiasteridae have emerged with phylogenetically distinct clades: Plazaster and Rathbunaster on separate clades within the boreal Asteriidae, and Coronaster with the pantropical Asteriidae. All members of the polyphyletic Labidiasteridae were characterized by large numbers of elongate arms (up to 50 in Labidiaster), biserial tube foot rows, and prominent pedicellariae. Based on our phylogenetic trees, these characters may be independently derived adaptations for benthopelagic predation, which has been observed in Labidiaster (Dearborn, Edwards & Fratt, 1991) and Rathbunaster (Lauerman, 1998).	en	Mah, Christopher, Foltz, David (2011): Molecular phylogeny of the Forcipulatacea (Asteroidea: Echinodermata): systematics and biogeography. Zoological Journal of the Linnean Society 162 (3): 646-660, DOI: 10.1111/j.1096-3642.2010.00688.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00688.x
03E687F8FFA1FFA63E332668FAA45E1D.taxon	description	Mah (1998, 1999) considered Odinella, Novodinia, and Brisingaster as having an intermediate brisingid morphology relative to an asteriid or labidiasterid sister taxon. This was especially the case for Odinella, which has the distinct, wing-like ambulacral ossicles present in asteriids rather than the vertebrae-like ambulacrals present in other brisingidan genera. This perspective is consistent with Mah’s (1998) hypothesis of a bathymetric shift between the shallower-water members of this clade and the deeper-water Freyellidae and Brisingidae. It should be noted that, although a new classification has not been finalized, our placement of the brisingidans as a derived sister branch to the Asteriidae disagrees with the ordinal-level ranking that places it on a parallel with the Forcipulatida.	en	Mah, Christopher, Foltz, David (2011): Molecular phylogeny of the Forcipulatacea (Asteroidea: Echinodermata): systematics and biogeography. Zoological Journal of the Linnean Society 162 (3): 646-660, DOI: 10.1111/j.1096-3642.2010.00688.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00688.x
03E687F8FFA1FFA93CCB2668FAE45CAF.taxon	description	The largest clade, containing the greatest number of species within the holarctic Asteriidae, is the one upholding the genus Leptasterias, a highly diverse lineage that has shown extensive diversification across the Arctic and temperate to high-latitude North Atlantic and North Pacific. All included species of Leptasterias were recovered on a single clade, which also included the North Pacific Evasterias and the Southern Hemisphere Perissasterias as sister taxa within the Leptasterias lineage. Previous phylogenetic studies of the genus Leptasterias employing more taxa and longer sequence reads than those analyzed here (Hrincevich, Rocha-Olivares & Foltz, 2000; Foltz et al., 2008) have reported somewhat different results from those in Figures 2 and 3: (1) Evasterias was strongly supported as the sister group to Leptasterias; (2) Leptasterias fisheri + Leptasterias stolacantha + Leptasterias muelleri was strongly supported as the sister group to the six-armed subgenus Hexasterias; (3) Leptasterias polaris (Polaris section of Fisher, 1930) was strongly supported as the sister group to the remaining Hexasterias (i. e. Fisher’s Camtschatica section); (4) Leptasterias leptodoma was supported as the sister group to the remaining members of the Camtschatica section. Leptasterias hylodes, Leptasterias ochotensis, Leptasterias squamulata were not sampled in the earlier studies. These differing results probably reflect the difficulty of inferring phylogenetic relationships within a recent, rapid radiation (Foltz et al., 2008). Two Southern Hemisphere genera, Perissasterias and Taranuiaster, were present among the predominantly Northern Hemisphere boreal Asteriidae. Further data are needed to make any definitive conclusions, but if the results are taken at face value, then two interpretations are possible for one or both of these genera. Perissasterias and Taranuiaster may belong to mostly extinct lineages of Asteriidae, which suggests a greater number of taxa with a more widespread distribution in the past, especially into the Southern Hemisphere. Alternatively, larval or adult dispersal may have transported ancestors of these taxa to the Southern Hemisphere, providing suitable environmental conditions to settle, thrive, and establish an adult population. Fisher (1930) noted that Perissasterias was one of the few Southern Hemisphere genera to share characters with Northern Hemisphere Asteriidae. Also, Perissasterias and Taranuiaster are deep-sea taxa, and live in environments that may parallel cold-water settings similar to those of Leptasterias and other boreal Asteriidae. Pycnopodia was supported as the sister taxon to Rathbunaster. Pycnopodia has been classified as one of two genera (the other being Lysastroma) within the Pycnopodiidae (see Clark & Mah, 2001). However, Lysastroma was not included in our treatment, which for the moment, prevents us from completely testing the monophyly and validity of the Pycnopodiidae.	en	Mah, Christopher, Foltz, David (2011): Molecular phylogeny of the Forcipulatacea (Asteroidea: Echinodermata): systematics and biogeography. Zoological Journal of the Linnean Society 162 (3): 646-660, DOI: 10.1111/j.1096-3642.2010.00688.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00688.x
03E687F8FFAEFFA83CFE21A7FE8B5B8B.taxon	description	The pantropical asteriid clade was supported as the sister taxon to the Antarctic asteriid clade in Figures 2 and 3, with 94 % bootstrap support on the two-gene tree and 97 % bootstrap support on the three-gene tree. There is a similar pattern observed in between the Antarctic urchin Sterechinus and its tropical South American sister taxa, Loxechinus and Pseudechinus (Lee et al., 2004). This is consistent with a relationship between asteriid taxa from the Antarctic / sub-Antarctic and adjacent tropical regions. Several non-asteriid Forcipulatacea show relationships between tropical and Antarctic / sub-Antarctic taxa. For example, the tropical shallow-water Heliaster is supported as the sister taxon to the Antarctic / sub-Antarctic Labidiaster. The Antarctic brisingidan Odinella is supported as sister taxon to a clade containing basal brisingidans Brisingaster and Novodinia. Although Novodinia is widely distributed, Brisingaster is known only from the Indian Ocean and New Caledonia (Mah, 1999). A more complete species-level data set, for both the Antarctic and the pantropical clade, will be needed to discover the closest sister taxon to the Antarctic fauna. The pantropical Asteriidae is well-supported as the sister group to the diverse Antarctic Asteriidae. The pantropical Asteriidae is composed of Coronaster, members of which occur mostly in the deep sea, as the sister clade to four shallow-water tropical genera: Astrostole, Astrometis, Coscinasterias, and Meyenaster. Astrostole, Astrometis, and Meyenaster are known only from the shallow-water settings from the Eastern Tropical Pacific, whereas Coronaster and Coscinasterias occur widely in the Atlantic and the Indo-Pacific. Coronaster occurs primarily in the deep sea (> 200 m), although some species (e. g. Coronaster marchenus) do occur in shallow waters (<10 m). Nearly every known genus of nominal Antarctic Asteriidae was supported as part of a single lineage, occurring only at high latitudes in the Southern Hemisphere. Relative to more stemward Antarctic / sub-Antarctic forcipulataceans, such as Granaster, Neosmilaster, or Labidiaster, the Antarctic Asteriidae clade evolved more recently and as a single event. Our trees did not support Diplasterias as monophyletic, but Diplasterias forms a species complex in the Southern Ocean region similar to the one formed by Leptasterias in the boreal north. This suggests that further taxonomic sampling and additional genes with longer reads are likely to contribute to a better understanding of Diplasterias, similar to the prior studies of Leptasterias discussed above. The diversification of the Antarctic Asteriidae may be associated with the isolation of the Southern Ocean fauna and the formation of the Antarctic Counter Current at the end of the Eocene / early Oligocene (25 Ma), as outlined by Clarke & Crame (1992) and Aronson et al. (2009). Distribution data for several of the genera included in the Antarctic asteriid clade (Fisher, 1940; Bernasconi, 1973, C. Mah, unpubl. data) show that Anasterias, Diplasterias, Lysasterias, and Psalidaster occur well into the South Atlantic as well as the Southern Ocean, with Adelasterias, Notasterias, and Saliasterias apparently limited to the Southern Ocean. We suggest that the ancestors to the Antarctic asteriid lineage occupied a much broader distribution from the Southern Ocean and the adjacent sub- Antarctic region, including the South Atlantic and South Pacific, prior to the formation of the Antarctic Circumpolar Current (ACC). The ACC would have isolated the Southern Ocean fauna from the adjoining population, leaving the adjoining population ‘ outside’ the ACC boundary in the South Atlantic / sub- Antarctic region. Several accounts have documented other invertebrate species that have a similar distribution, but which demonstrate either limited or no gene flow across the ACC (e. g. Hunter & Halanych, 2008; Thornhill et al., 2008). This suggests that nominal Antarctic asteriid species, with identical morphology on either side of the ACC, could be genetically distinct, cryptic species complexes. The presence of Southern Ocean and sub-Antarctic populations separated by the ACC would also be consistent with widespread circumpolar distribution of so many of the Antarctic asteriids, which lack planktonic larvae. The sub-Antarctic populations would conceivably be ancestral relative to those in the Southern Ocean. This notion would also be consistent with the seeming plesiomorphic morphology shared by Psalidaster, Saliasterias, and several multi-armed and widespread members of the pantropical Asteriidae, such as Coronaster and Astrostole. However, further comprehensive biogeographic analyses of the clade members would be necessary to test this hypothesis.	en	Mah, Christopher, Foltz, David (2011): Molecular phylogeny of the Forcipulatacea (Asteroidea: Echinodermata): systematics and biogeography. Zoological Journal of the Linnean Society 162 (3): 646-660, DOI: 10.1111/j.1096-3642.2010.00688.x, URL: http://dx.doi.org/10.1111/j.1096-3642.2010.00688.x
