Christoph D. Schubart

Rapid evolution to terrestrial life in Jamaican crabs

Crabs of the family Grapsidae are abundant organisms in most intertidal communities. However, relatively few species live in complete independence of the sea. Of those species that do, Jamaicas nine endemic species of land crabs are unique in their exceptional adaptations to terrestrial life, which include the only active brood-care for larvae and juveniles known in crabs. These adaptations, and the morphological similarity to a group of southeast Asian land-dwelling crabs, have raised the question of the number and age of land invasions of the Jamaican species. Here we present molecular evidence that Jamaican land crabs represent a single adaptive radiation from a marine ancestor that invaded terrestrial habitats only 4 million years (Myr) ago. A Late-Tertiary origin has also been found for lizards and frogs of Jamaica and probably reflects the Mid-Tertiary inundation of that island.

GLYPTOGRAPSIDAE, A NEW BRACHYURAN FAMILY FROM CENTRAL AMERICA: LARVAL AND ADULT MORPHOLOGY, AND A MOLECULAR PHYLOGENY OF THE GRAPSOIDEA

Abstract During an ongoing systematic revision of the Decapoda Grapsoidea (here defined as including the families Gecarcinidae, Grapsidae s. str., Plagusiidae, Sesarmidae, and Varunidae; see Schubart et al., 2000a), it became evident that adult and larval morphology of two Central American genera, Glyptograpsus and Platychirograpsus, differs greatly from that of all other genera within this superfamily. Several important morphological characters are shared by these two genera and represent synapomorphies when compared to the other grapsoids. Both of these genera lack a pleurobranch on the sixth thoracic segment. Adult males of Glyptograpsus and Platychirograpsus are all strikingly heterochelous with the major chela being conspicuously flattened anteriorly and showing a subproximal articulation with the carpus. The distal portion of the male gonopod is uncinate, with a narrowed, elongate terminus. The anteriormost portion of the sternum consists of a narrow, fused sternite terminated in a distinctly flanged tip inserted between the mouth appendages. The male abdomen exhibits immobility in the joints between segments 3 to 5 and very limited mobility in the joint between segments 1 and 2. Zoeae of the two genera show a 1,2 setation pattern on the maxillar endopod. A molecular phylogeny of the Grapsoidea, based on 16S mtDNA and including type genera of the five recognized families, confirms that the species of Glyptograpsus and Platychirograpsus together form a well-defined monophyletic unit that is distinct from all other taxa within the Grapsoidea. We therefore describe a new family, the Glyptograpsidae, to accommodate these two genera of crabs.

Production of different phenotypes from the same genotype in the same environment by developmental variation

SUMMARY The phenotype of an organism is determined by the genes, the environment and stochastic developmental events. Although recognized as a basic biological principle influencing life history, susceptibility to diseases, and probably evolution, developmental variation (DV) has been only poorly investigated due to the lack of a suitable model organism. This obstacle could be overcome by using the recently detected, robust and highly fecund parthenogenetic marbled crayfish as an experimental animal. Batch-mates of this clonal crayfish, which were shown to be isogenic by analysis of nuclear microsatellite loci, exhibited surprisingly broad ranges of variation in coloration, growth, life-span, reproduction, behaviour and number of sense organs, even when reared under identical conditions. Maximal variation was observed for the marmorated coloration, the pattern of which was unique in each of the several hundred individuals examined. Variation among identically raised batch-mates was also found with respect to fluctuating asymmetry, a traditional indicator of the epigenetic part of the phenotype, and global DNA methylation, an overall molecular marker of an animals epigenetic state. Developmental variation was produced in all life stages, probably by reaction–diffusion-like patterning mechanisms in early development and non-linear, self-reinforcing circuitries involving behaviour and metabolism in later stages. Our data indicate that, despite being raised in the same environment, individual genotypes can map to numerous phenotypes via DV, thus generating variability among clone-mates and individuality in a parthenogenetic species. Our results further show that DV, an apparently ubiquitous phenomenon in animals and plants, can introduce components of randomness into life histories, modifying individual fitness and population dynamics. Possible perspectives of DV for evolutionary biology are discussed.

Genetic, ecological, and behavioural divergence between two sibling snapping shrimp species (Crustacea: Decapoda: Alpheus)

Examination of genetic and ecological relationships within sibling species complexes can provide insights into species diversity and speciation processes. Alpheus angulatus and A. armillatus, two snapping shrimp species with overlapping ranges in the north‐western Atlantic, are similar in morphology, exploit similar ecological niches and appear to represent recently diverged sibling species. We examined phylogenetic and ecological relationships between these two species with: (i) sequence data from two mitochondrial genes (16S rRNA and COI); (ii) data on potential differences in microhabitat distribution for A. armillatus and A. angulatus; and (iii) data from laboratory experiments on the level of reproductive isolation between the two species. DNA sequence data suggest A. armillatus and A. angulatus are sister species that diverged subsequent to the close of the Isthmus of Panama, and that haplotype diversity is lower in A. armillatus than in A. angulatus. Both species are distantly related to A. heterochaelis and A. estuariensis, two species with which A. angulatus shares some similarities in coloration. Ecological data on the distribution of A. angulatus and A. armillatus from two locations revealed differences in distribution of the two species between habitat patches, with each patch dominated by one or the other species. However, there was no apparent difference in distribution of the two species within habitat patches with respect to microhabitat location. Ecological data also revealed that heterospecific individuals often occur in close proximity (i.e. within metres or centimetres) where sympatric. Behavioural data indicated that these species are reproductively isolated, which is consistent with speciation in transient allopatry followed by post‐divergence secondary contact. Our data further resolve taxonomic confusion between the sibling species, A. armillatus and A. angulatus, and suggest that sympatry in areas of range overlap and exploitation of similar ecological niches by these two recently diverged species have selected for high levels of behavioural incompatibility.

When Indian crabs were not yet Asian--biogeographic evidence for Eocene proximity of India and Southeast Asia.

BackgroundThe faunal and floral relationship of northward-drifting India with its neighboring continents is of general biogeographic interest as an important driver of regional biodiversity. However, direct biogeographic connectivity of India and Southeast Asia during the Cenozoic remains largely unexplored. We investigate timing, direction and mechanisms of faunal exchange between India and Southeast Asia, based on a molecular phylogeny, molecular clock-derived time estimates and biogeographic reconstructions of the Asian freshwater crab family Gecarcinucidae.ResultsAlthough the Gecarcinucidae are not an element of an ancient Gondwana fauna, their subfamily Gecarcinucinae, and probably also the Liotelphusinae, evolved on the Indian Subcontinent and subsequently dispersed to Southeast Asia. Estimated by a model testing approach, this dispersal event took place during the Middle Eocene, and thus before the final collision of India and the Tibet-part of Eurasia.ConclusionsWe postulate that the India and Southeast Asia were close enough for exchange of freshwater organisms during the Middle Eocene, before the final Indian-Eurasian collision. Our data support geological models that assume the Indian plate having tracked along Southeast Asia during its move northwards.

Marine biogeographic boundaries and human introduction along the European coast revealed by phylogeography of the prawn Palaemon elegans

A phylogeographic analysis is carried out for the widely distributed European littoral prawn Palaemon elegans in order to test for potential genetic differentiation and geographic structure. Mitochondrial sequences were obtained from 283 specimens from the northeastern Atlantic, the Baltic, Mediterranean, Black and Caspian Seas. Our study revealed a surprisingly complex population structure. Three main haplogroups can be separated: one from the Atlantic (Type I) and two from the Mediterranean (Types II and III). While the Mediterranean types occur in sympatry, a clear phylogeographic break was observed along the Almería-Oran Front separating Type I and giving evidence for a genetic isolation of Atlantic and Mediterranean populations. Type III represents the most distinct haplogroup with high levels of nucleotide divergence, indicating the occurrence of a cryptic species with a Messinian origin. The colonization of the southeastern Baltic Sea is most likely due to human introduction.

Phylogenetic and morphometric differentiation reveal geographic radiation and pseudo-cryptic speciation in a mangrove crab from the Indo-West Pacific

The presence of boundaries to dispersal has been recently documented for many Indo-West Pacific (IWP) species with planktonic propagules and a widespread distribution. We studied the phylogeography of the mangrove crab Neosarmatium meinerti (Brachyura: Sesarmidae) and the phylogenetic relationship to its presumed sister species N. fourmanoiri in the IWP in order to compare intraspecific with interspecific diversity. Portions of the mitochondrial genes 16S and CoxI were sequenced for 23 specimens of N. meinerti and 5 N. fourmanoiri, while a fragment of the 28S was obtained for a subset of specimens. Genetic data are supplemented by morphometric and based on 37 adult males of N. meinerti and 9 males of N. fourmanoiri. The conserved nuclear 28S reveals the existence of a genetic break between the Indian and Pacific oceans. Otherwise, mitochondrial genes as well as morphometry clearly support the presence of a species complex within N. meinerti composed by four well structured and geographically defined lineages: East African coast; western Indian Ocean islands; South East Asia; and Australia.

Exploring Pandora's Box: Potential and Pitfalls of Low Coverage Genome Surveys for Evolutionary Biology

High throughput sequencing technologies are revolutionizing genetic research. With this “rise of the machines”, genomic sequences can be obtained even for unknown genomes within a short time and for reasonable costs. This has enabled evolutionary biologists studying genetically unexplored species to identify molecular markers or genomic regions of interest (e.g. micro- and minisatellites, mitochondrial and nuclear genes) by sequencing only a fraction of the genome. However, when using such datasets from non-model species, it is possible that DNA from non-target contaminant species such as bacteria, viruses, fungi, or other eukaryotic organisms may complicate the interpretation of the results. In this study we analysed 14 genomic pyrosequencing libraries of aquatic non-model taxa from four major evolutionary lineages. We quantified the amount of suitable micro- and minisatellites, mitochondrial genomes, known nuclear genes and transposable elements and searched for contamination from various sources using bioinformatic approaches. Our results show that in all sequence libraries with estimated coverage of about 0.02–25%, many appropriate micro- and minisatellites, mitochondrial gene sequences and nuclear genes from different KEGG (Kyoto Encyclopedia of Genes and Genomes) pathways could be identified and characterized. These can serve as markers for phylogenetic and population genetic analyses. A central finding of our study is that several genomic libraries suffered from different biases owing to non-target DNA or mobile elements. In particular, viruses, bacteria or eukaryote endosymbionts contributed significantly (up to 10%) to some of the libraries analysed. If not identified as such, genetic markers developed from high-throughput sequencing data for non-model organisms may bias evolutionary studies or fail completely in experimental tests. In conclusion, our study demonstrates the enormous potential of low-coverage genome survey sequences and suggests bioinformatic analysis workflows. The results also advise a more sophisticated filtering for problematic sequences and non-target genome sequences prior to developing markers.

Is the Species Flock Concept Operational? The Antarctic Shelf Case

There has been a significant body of literature on species flock definition but not so much about practical means to appraise them. We here apply the five criteria of Eastman and McCune for detecting species flocks in four taxonomic components of the benthic fauna of the Antarctic shelf: teleost fishes, crinoids (feather stars), echinoids (sea urchins) and crustacean arthropods. Practical limitations led us to prioritize the three historical criteria (endemicity, monophyly, species richness) over the two ecological ones (ecological diversity and habitat dominance). We propose a new protocol which includes an iterative fine-tuning of the monophyly and endemicity criteria in order to discover unsuspected flocks. As a result nine « full » species flocks (fulfilling the five criteria) are briefly described. Eight other flocks fit the three historical criteria but need to be further investigated from the ecological point of view (here called « core flocks »). The approach also shows that some candidate taxonomic components are no species flocks at all. The present study contradicts the paradigm that marine species flocks are rare. The hypothesis according to which the Antarctic shelf acts as a species flocks generator is supported, and the approach indicates paths for further ecological studies and may serve as a starting point to investigate the processes leading to flock-like patterning of biodiversity.

High regional differentiation in a North American crab species throughout its native range and invaded European waters: a phylogeographic analysis

Rhithropanopeus harrisii (Gould 1841) has a native distribution from New Brunswick (Canada) to Veracruz (Mexico) and is considered an invasive species in northwestern North American (Oregon and California), South American (Brazil) and European estuaries and rivers. In Europe, it was observed for the first time in 1874, in The Netherlands. We sequenced and analyzed part of the cytochrome oxidase subunit I gene (mitochondrial DNA) of eight populations, three from the east coast of the United States of America (USA) and five from Europe, in order to assess their genetic diversity and to determine a potential founder population. European populations are characterized by a lower number of haplotypes than the whole native region of the eastern USA, suggesting that genetic bottlenecks occurred during the European colonisation. Along the North American East Coast, there is evidence of clearcut genetic heterogeneity, New Jersey being the most similar population in its genetic structure to the postulated Europe-founding population. Also the different European populations are heterogeneous and there is a tendency of higher genetic diversity in the populations founded earlier. R. harrisii is still in the process of expansion in Europe and may have been introduced once or repeatedly by different invasion mechanisms. The pronounced lack of gene flow among populations is of great ecological significance, since it may facilitate rapid adaptation and specialization to local conditions within single estuarine systems.