Fixed, Free, and Fixed: The Fickle Phylogeny of Extant Crinoidea (Echinodermata) and Their Permian-Triassic Origin

Although the status of Crinoidea (sea lilies and featherstars) as sister group to all other living echinoderms is well-established, relationships among crinoids, particularly extant forms, are debated. All living species are currently placed in Articulata, which is generally accepted as the only crinoid group to survive the Permian–Triassic extinction event. Recent classifications have recognized five major extant taxa: Isocrinida, Hyocrinida, Bourgueticrinina, Comatulidina and Cyrtocrinida, plus several smaller groups with uncertain taxonomic status, e.g., Guillecrinus, Proisocrinus and Caledonicrinus. Here we infer the phylogeny of extant Crinoidea using three mitochondrial genes and two nuclear genes from 59 crinoid terminals that span the majority of extant crinoid diversity. Although there is poor support for some of the more basal nodes, and some tree topologies varied with the data used and mode of analysis, we obtain several robust results. Cyrtocrinida, Hyocrinida, Isocrinida are all recovered as clades, but two stalked crinoid groups, Bourgueticrinina and Guillecrinina, nest among the featherstars, lending support to an argument that they are paedomorphic forms. Hence, they are reduced to families within Comatulida. Proisocrinus is clearly shown to be part of Isocrinida, and Caledonicrinus may not be a bourgueticrinid. Among comatulids, tree topologies show little congruence with current taxonomy, indicating that much systematic revision is required. Relaxed molecular clock analyses with eight fossil calibration points recover Articulata with a median date to the most recent common ancestor at 231–252 mya in the Middle to Upper Triassic. These analyses tend to support the hypothesis that the group is a radiation from a small clade that passed through the Permian–Triassic extinction event rather than several lineages that survived. Our tree topologies show various scenarios for the evolution of stalks and cirri in Articulata, so it is clear that further data and taxon sampling are needed to recover a more robust phylogeny of the group

Molecular cloning and characterization of the polymorphic MHC class II DBB from the tammar wallaby (Macropus eugenii)

Genes of the major histocompatibility complex (MHC) have been characterized in all extant lineages of mammals. The tammar wallaby (Macropus eugenii) is well established as a model marsupial species; however, no classical MHC sequences have been described from this species. We have isolated two MHC class II β-chain sequences from a tammar wallaby spleen cDNA library using a tammar MHC class II β probe. These sequences belong to the marsupial MHC class II DBB gene family. Two additional DBB sequences were amplified from tammar wallaby genomic DNA. All four sequences were obtained from the same individual, indicating that there are at least two DBB loci in the tammar wallaby.5 page(s

Reduced MHC class II diversity in island compared to mainland populations of the black-footed rock-wallaby (Petrogale lateralis lateralis)

Many animal populations that are endangered in mainland areas exist in stable island populations, which have the potential to act as an “ark” in case of mainland population declines. Previous studies have found neutral genetic variation in such species to be up to an order of magnitude lower in island compared to mainland populations. If low genetic variation is prevalent across fitness-related loci, this would reduce the effectiveness of island populations as a source of individuals to supplement declining mainland populations or re-establish extinct mainland populations. One such species, the black-footed rock-wallaby (Petrogale lateralis lateralis), exists within fragmented mainland populations and small island populations off Western Australia. We examined sequence variation in this species within a fitness-related locus under positive selection, the MHC class II DAB β1 locus. The mainland populations displayed greater levels of allelic diversity (4–7 alleles) than the island population, despite being small and isolated, and contained at least two DAB gene copies. The island population displayed low allelic diversity (2 alleles) and fewer alleles per individual in comparison to mainland populations, and probably possesses only one DAB gene copy. The patterns of DAB diversity suggested that the island population has a markedly lower level of genetic variation than the mainland populations, in concordance with results from microsatellites (genotyped in a previous study), but preserved unique alleles which were not found in mainland populations. Where possible, conservation actions should pool individuals from multiple populations, not only island populations, for translocation programs, and focus on preventing further declines in mainland populations.13 page(s

Multifaceted genetic analysis of the "critically endangered" brush-tailed rock-wallaby Petrogale penicillata in Victoria, Australia : implications for management

The use of molecular genetic techniques can aid wildlife managers in setting priorities and devising management strategies for scattered populations of threatened taxa. In this study, six remnant populations of the "critically endangered" brush-tailed rock-wallaby (Petrogale penicillata) in Victoria, Australia, were examined using karyotypic, microsatellite (11 loci) and mitochondrial DNA (mtDNA) control region sequence analysis. Each remnant population was found to be genetically distinct (unique microsatellite alleles and control region haplotypes), but had low genetic diversity. This distribution of genetic diversity between, rather than within populations, is most likely a consequence of recent severe reductions in population size and dispersal that have occurred since European settlement. The six mtDNA control region haplotypes identified in the Victorian populations were all closely related (average 1.3% sequence divergence), and only 2% divergence separated haplotypes from East Gippsland and the Grampians (550 km to the west). In contrast there was considerable sequence divergence (7.7%) between the Victorian haplotypes and those found in P. penicillata from elsewhere in the species range. In comparison, 8.8% divergence separates P. penicillata from the closely related P. herberti. The Victorian haplotypes also formed a distinct and well supported monophyletic group that excluded haplotypes from other P. penicillata and P. herberti. In light of these data, we recommend that the remnant Victorian populations of P. penicillata be managed separately from remaining populations in New South Wales and Queensland; and that individuals be regularly exchanged amongst the Victorian populations to increase their diversity and reduce the likelihood of inbreeding depression

Genetic analysis of a population crash in brush-tailed rock-wallabies (Petrogale penicillata) from Jenolan Caves, south-eastern Australia

Although the theoretical effects of a severe reduction in effective population size (i.e. a bottleneck) are well known, relatively few empirical studies of bottlenecks have been based on extensive temporally spaced samples of a population both before and after a bottleneck. Here we describe the results of one such study, utilising the Jenolan Caves (JC) population of the brush-tailed rock-wallaby (Petrogale penicillata). When first sampled in 1985 (n = 20) the JC population comprised ~90 individuals. Subsequently the population crashed, and by 1992 only seven individuals remained. In 1996 the entire population (n = 10) was again sampled. Genetic diversity in the pre- and post-crash JC population was compared using 11 polymorphic microsatellite loci and PCR–SSCP analysis of the mitochondrial DNA control region. Only a single unique control region haplotype was detected in the pre- and post-crash JC population, although variant haplotypes were present in other P. penicillata populations. Of the 35 microsatellite alleles present in the pre-crash population, nine (26%) were lost during the bottleneck. The average number of rare alleles declined by 72%, allelic diversity was reduced by 30% and average heterozygosity declined by 10%. These observations are consistent with theoretical predictions. Additional analyses revealed that a P. penicillata female at Wombeyan Caves was the only survivor of a 1990/91 reintroduction attempt using animals from JC. Of the microsatellite alleles detected in this female, 21% (4/19) were no longer present in the post-crash JC population. Furthermore, the genetic profiles of animals from the recently discovered Taralga population indicate that they are not derived from JC stock, but represent a threatened remnant of a hitherto undetected natural P. penicillata population