Savel R. Daniels

Global diversity of crabs (Crustacea: Decapoda: Brachyura) in freshwater

An assessment of the global freshwater crab diversity is presented. A total of 1,476 species in 14 families are currently known from all zoogeographical regions (except Antarctica), including 1,306 species in eight exclusively freshwater families (Pseudothelphusidae, Trichodactylidae, Potamonautidae, Deckeniidae, Platythelphusidae, Potamidae, Gecarcinucidae and Parathelphusidae). Estimates of true freshwater crab diversity including likely numbers of undescribed taxa suggest that the field remains largely in a “discovery” phase. Main ideas on the origins, diversification, and phylogeny of true freshwater crabs are briefly discussed. The economic importance of freshwater crabs is also highlighted.

A living fossil tale of Pangaean biogeography

The current distributions of widespread groups of terrestrial animals and plants are supposedly the result of a mixture of either vicariance owing to continental split or more recent trans-oceanic dispersal. For organisms exhibiting a vicariant biogeographic pattern—achieving their current distribution by riding on the plates of former supercontinents—this view is largely inspired by the belief that Pangaea lacked geographical or ecological barriers, or that extinctions and dispersal would have erased any biogeographic signal since the early Mesozoic. We here present a time-calibrated molecular phylogeny of Onychophora (velvet worms), an ancient and exclusively terrestrial panarthropod group distributed throughout former Pangaean landmasses. Our data not only demonstrate that trans-oceanic dispersal does not need be invoked to explain contemporary distributions, but also reveal that the early diversification of the group pre-dates the break-up of Pangaea, maintaining regionalization even in landmasses that have remained contiguous throughout the history of the group. These results corroborate a growing body of evidence from palaeontology, palaeogeography and palaeoclimatic modelling depicting ancient biogeographic regionalization over the continuous landmass of Pangaea.

Molecular data suggest that melanistic ectotherms at the south-western tip of Africa are the products of Miocene climatic events: Evidence from cordylid lizards

In the present study the evolutionary relationships among Cordylus niger–oelofseni–cordylus are investigated using molecular sequence data to: (1) estimate the divergence within this group; (2) date the origin of melanism in the clade and to evaluate possible palaeoecological events responsible for its evolution; (3) examine the systematic relationships among the three isolated populations of C. oelofseni that were previously shown to be polyphyletic using allozyme data. The recovered topology for the combined sequence data (16S and NADH dehydrogenase component 2 (ND2)) among C . niger–oelofseni–cordylus suggests that melanism evolved only once in the clade. The application of a molecular clock to the data demonstrates that melanism evolved during the Miocene epoch, 17–15 million years ago. We believe that the development of a cold-water current and upwelling system along the south-west coast of Africa during this epoch was instrumental in the evolution of melanism in this lizard clade and possibly also in other ectotherm clades with melanistic forms occurring in the area. The results show that C. oelofseni , as presently construed, is composed of three distinct evolutionary lineages, with no shared haplotypes present and marked sequence divergence. Two of the three lineages represent undescribed species. Cordylus cordylus is composed of two distinct clades, a montane clade and a coastal lowland clade. Cladogenic activity in mountain areas of the Western Cape in South Africa has been particularly pronounced and the systematic diversity of many taxa may have been underestimated.

Geographic patterns of genetic and morphological divergence amongst populations of a river crab (Decapoda, Potamonautidae) with the description of a new species from mountain streams in the Western Cape, South Africa

Recent systematic research has revealed that Potamonautes brincki comprises two genetically and morphologically distinct population groups. The systematic affinities between these population groups have remained uncertain. In the present study, the relationship between the population groups was examined. Eleven populations were collected from high mountain streams in the Western Cape, South Africa and used in the genetic and morphological analyses. Allozyme electrophoresis of 13 protein coding loci separated two main population groups: group A (Cape Peninsular groups) and group B (Hottentot’s Holland) at I = 0.73. Two additional genetic groups were evident, with group B being conspecific to group A, and group D being conspecific to group C. Morphological examination of pleopod 1 and the terminal segment of the mandibular palp showed considerable differences between the two main population groups, with groups A and B being similar and groups C and D being similar. The morphometric data for the four main groups were examined using discriminant functions analysis and the two main groups were compared using analyses of covariance. Discriminant functions analysis showed a moderate degree of overlap between the groups. Additional morphometric data showed a clear discrimination between the two main population groups. The genetic and morphometric data sets exhibited congruent patterns of variation and the data showed the presence of a species boundary. A new freshwater crab species, P. parvicorpus sp. n., is described. The results of the present study are discussed in the light of historical and contemporary factors that are likely to have contributed to speciation.

Population structure, propagule pressure, and conservation biogeography in the sub‐Antarctic: lessons from indigenous and invasive springtails

The patterns in and the processes underlying the distribution of invertebrates among Southern Ocean islands and across vegetation types on these islands are reasonably well understood. However, few studies have examined the extent to which populations are genetically structured. Given that many sub-Antarctic islands experienced major glaciation and volcanic activity, it might be predicted that substantial population substructure and little genetic isolation-by-distance should characterize indigenous species. By contrast, substantially less population structure might be expected for introduced species. Here, we examine these predictions and their consequences for the conservation of diversity in the region. We do so by examining haplotype diversity based on mitochondrial cytochrome c oxidase subunit I sequence data, from two indigenous (Cryptopygus antarcticus travei, Tullbergia bisetosa) and two introduced (Isotomurus cf. palustris, Ceratophysella denticulata) springtail species from Marion Island. We find considerable genetic substructure in the indigenous species that is compatible with the geological and glacialogical history of the island. Moreover, by employing ecological techniques, we show that haplotype diversity is likely much higher than our sequenced samples suggest. No structure is found in the introduced species, with each being represented by a single haplotype only. This indicates that propagule pressure is not significant for these small animals unlike the situation for other, larger invasive species: a few individuals introduced once are likely to have initiated the invasion. These outcomes demonstrate that sampling must be more comprehensive if the population history of indigenous arthropods on these islands is to be comprehended, and that the risks of within- and among-island introductions are substantial. The latter means that, if biogeographical signal is to be retained in the region, great care must be taken to avoid inadvertent movement of indigenous species among and within islands. Thus, quarantine procedures should also focus on among-island movements.

Deep genealogical lineages in the widely distributed African helmeted terrapin: Evidence from mitochondrial and nuclear DNA (Testudines: Pelomedusidae: Pelomedusa subrufa)

We investigated the phylogeographic differentiation of the widely distributed African helmeted terrapin Pelomedusa subrufa based on 1503 base pairs of mitochondrial DNA (partial cyt b and ND4 genes with adjacent tRNAs) and 1937 bp of nuclear DNA (partial Rag1, Rag2, R35 genes). Congruent among different analyses, nine strongly divergent mitochondrial clades were found, representing three major geographical groupings: (1) A northern group which includes clades I from Cameroon, II from Ghana and Ivory Coast, III from Benin, Burkina Faso and Niger, IV from the Central African Republic, and V from Kenya, (2) a northeastern group consisting of clades VI from Somalia, and VII from Saudi Arabia and Yemen, and (3) a southern group comprising clade VIII from Botswana, the Democratic Republic of Congo, Madagascar and Malawi, and clade IX from South Africa. Malagasy and continental African populations were not clearly differentiated, indicating very recent arrival or introduction of Pelomedusa in Madagascar. The southern group was in some phylogenetic analyses sister to Pelusios, rendering Pelomedusa paraphyletic with respect to that genus. However, using partitioned Bayesian analyses and sequence data of the three nuclear genes, Pelomedusa was monophyletic, suggesting that its mitochondrial paraphyly is due to either ancient introgressive hybridization or phylogenetic noise. Otherwise, nuclear sequence data recovered a lower level of divergence, but corroborated the general differentiation pattern of Pelomedusa as revealed by mtDNA. This, and the depth of the divergences between clades, indicates ancient differentiation. The divergences observed fall within, and in part exceed considerably, the differentiation typically occurring among chelonian species. To test whether Pelomedusa is best considered a single species composed of deep genealogical lineages, or a complex of up to nine distinct species, we suggest a future taxonomic revision that should (1) extend the geographical sampling of molecular data, specifically focusing on contact zones and the possible sympatric occurrence of lineages without admixture, and (2) evaluate the morphology of the various genealogical lineages using the type specimens or topotypical material of the numerous junior synonyms of P. subrufa.

The mitochondrial genome of the onychophoran Opisthopatus cinctipes (Peripatopsidae) reflects the ancestral mitochondrial gene arrangement of Panarthropoda and Ecdysozoa

The ancestral genome composition in Onychophora (velvet worms) is unknown since only a single species of Peripatidae has been studied thus far, which shows a highly derived gene order with numerous translocated genes. Due to this lack of information from Onychophora, it is difficult to infer the ancestral mitochondrial gene arrangement patterns for Panarthropoda and Ecdysozoa. Hence, we analyzed the complete mitochondrial genome of the onychophoran Opisthopatus cinctipes, a representative of Peripatopsidae. Our data show that O. cinctipes possesses a highly conserved gene order, similar to that found in various arthropods. By comparing our results to those from different outgroups, we reconstruct the ancestral gene arrangement in Panarthropoda and Ecdysozoa. Our phylogenetic analysis of protein-coding gene sequences from 60 protostome species (including outgroups) provides some support for the sister group relationship of Onychophora and Arthropoda, which was not recovered by using a single species of Peripatidae, Epiperipatus biolleyi, in a previous study. A comparison of the strand-specific bias between onychophorans, arthropods, and a priapulid suggests that the peripatid E. biolleyi is less suitable for phylogenetic analyses of Ecdysozoa using mitochondrial genomic data than the peripatopsid O. cinctipes.

Population structure, propagule pressure, and conservation biogeography in the sub-Antarctic: Lessons from indigenous and invasive springtails: Biodiversity research

The patterns in and the processes underlying the distribution of invertebrates among Southern Ocean islands and across vegetation types on these islands are reasonably well understood. However, few studies have examined the extent to which populations are genetically structured. Given that many sub-Antarctic islands experienced major glaciation and volcanic activity, it might be predicted that substantial population substructure and little genetic isolation-by-distance should characterize indigenous species. By contrast, substantially less population structure might be expected for introduced species. Here, we examine these predictions and their consequences for the conservation of diversity in the region. We do so by examining haplotype diversity based on mitochondrial cytochrome c oxidase subunit I sequence data, from two indigenous (Cryptopygus antarcticus travei, Tullbergia bisetosa) and two introduced (Isotomurus cf. palustris, Ceratophysella denticulata) springtail species from Marion Island. We find considerable genetic substructure in the indigenous species that is compatible with the geological and glacialogical history of the island. Moreover, by employing ecological techniques, we show that haplotype diversity is likely much higher than our sequenced samples suggest. No structure is found in the introduced species, with each being represented by a single haplotype only. This indicates that propagule pressure is not significant for these small animals unlike the situation for other, larger invasive species: a few individuals introduced once are likely to have initiated the invasion. These outcomes demonstrate that sampling must be more comprehensive if the population history of indigenous arthropods on these islands is to be comprehended, and that the risks of within- and among-island introductions are substantial. The latter means that, if biogeographical signal is to be retained in the region, great care must be taken to avoid inadvertent movement of indigenous species among and within islands. Thus, quarantine procedures should also focus on among-island movements.

Congruent patterns of genetic variation in a burrowing freshwater crab revealed by allozymes and mt DNA sequence analysis

Five populations of the burrowing freshwater crab, Potamonautes calcaratus representing a total of 100 specimens, were collected from the Kruger National Park, South Africa. The population genetic structure of this species was investigated using both nuclear genetic markers (allozymes), and direct sequencing of a 610 base pair fragment the cytochrome oxidase 1(CO 1) subunit of the mitochondrial DNA. Electrophoresis of 21 allozyme loci revealed that moderate levels of genetic differentiation (F(ST) = 0.12) was present among populations. Sequence data for 20 individuals revealed the presence of ten haplotypes, the distribution of which showed no geographic structuring. The ΦCT of 0.43 corroborates a moderate degree of genetic structuring. The nucleotide diversity (π) was low and ranged from 0.00 to 0.007. Sequence divergence amongst populations ranged from 0.49% to 1.47%. Both genetic markers revealed moderate population structuring, supporting the conclusion that populations share a common recent ancestry, with moderate levels of recent gene flow. These results provide evidence that allozyme and sequencing data may be congruent and that these independent markers can detect similar patterns of genetic differentiation. Results are discussed in light of contemporary factors that have been likely in sculpting the genetic structure.

A molecular phylogeny for the South African limbless lizard taxa of the subfamily Acontinae (Sauria: Scincidae) with special emphasis on relationships within Acontias

In the present study, relationships among three genera Acontias, Acontophiops, and Typhlosaurus, that comprise the South African limbless lizard subfamily Acontinae, were assessed with partial sequences of the 16S rRNA mitochondrial DNA gene. In addition, relationships within Acontias were further investigated using sequence data from the cytochrome oxidase I gene (COI). Maximum likelihood and maximum parsimony analyses of the 16S rRNA mtDNA data revealed that within this subfamily, Typhlosaurus is basal while Acontophiops and Acontias are sister taxa. Based on the 16S rRNA mtDNA data, the relationships within Acontias placed A. meleagris orientalis as the sister taxon of A. percivali tasmani, with A. m. orientalis lineacauda morph and A. m. meleagrus being the sister taxa to this group. The small-bodied skinks A. lineatus lineatus and A. l. tristis formed a monophyletic group, with the medium-bodied species A. gracilicauda gracilicauda being their sister taxon. Analyses of the COI gene for Acontias place A. m. orientalis as the sister taxon of A. p. tasmani with both A. meleagris meleagris and A. m. orientalis lineacauda being distinct. In contrast to the 16S rRNA mtDNA data, the COI data placed A. g. gracilicauda as the sister taxon to these medium-bodied species; while the subspecies status of the small-bodied taxa A. l. lineatus and A. l. tristis is reaffirmed. Combined analysis of both gene fragments for Acontias taxa recovered the same clades as found using only COI data. Systematic affinities in Acontias are discussed. These results indicate that Acontias is more species rich than previously thought.