Molecular phylogeny and zoogeography of the freshwater crayfish genus Cherax Erichson (Decapoda: Parastacidae) in Australia

The evolutionary history and biogeography of freshwater-dependent taxa in Australia is of intrinsic interest given the present-day aridity of this continent. Cherax is the most widespread and one of the most species-rich of Australia's nine freshwater crayfish genera. The phylogenetic relationships amongst 19 of the 23 Australian Cherax were established from mitochondrial DNA sequences representing the 12S rRNA and 16S rRNA gene regions. The relationships among species support an initial east–west separation, followed by a north–south divergence in eastern Australia. Molecular clock estimations suggest that these divergences date back to the Miocene. The phylogenetic relationships support endemic speciation within geographical regions and indicate that long-distance dispersal has not led to recent speciation as previously hypothesized. This new evolutionary scenario is consistent with the climatic history of Australia and the evolutionary history of other similarly distributed freshwater-dependent organisms in Australia.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 81 , 553–563.     

Amplification of multiple copies of mitochondrial Cytochrome b gene fragments in the Australian freshwater crayfish,Cherax destructor Clark (Parastacidae: Decapoda)

Non-coding copies of fragments of the mitochondrial genome translocated to the nucleus or pseudogenes are being found with increasing frequency in a diversity of organisms. As part of a study to evaluate the utility of a range of mitochondrial gene regions for population genetic and systematic studies of the Australian freshwater crayfish, Cherax destructor (the yabby), we report the first detection of Cytochrome b (Cyt b) pseudogenes in crustaceans. We amplified and sequenced fragments of the mitochondrial Cyt b gene from 14 individuals of C. destructor using polymerase chain reaction (PCR) with primers designed from conserved regions of Penaeus monodon and Drosophila melanogaster mitochondrial genomes. The phylogenetic tree produced from the amplified fragments using these primers showed a very different topology to the trees obtained from sequences from three other mitochondrial genes, suggesting one or more nuclear pseudogenes have been amplified. Supporting this conclusion, two highly divergent sequences were isolated from each of two single individuals, and a 2 base pair (bp) deletion in one sequence was observed. There was no evidence to support inadvertent amplification of parasite DNA or contamination of samples from other sources. These results add to other recent observations of pseudogenes suggesting the frequent transfer of mitochondrial DNA (mtDNA) genes to the nucleus and reinforces the necessity of great care in interpreting PCR-generated Cyt b sequences used in population or evolutionary studies in freshwater crayfish and crustaceans more generally.     

Phylogeographic structure in the gilgie (Decapoda: Parastacidae: Cherax quinquecarinatus): a south‐western Australian freshwater crayfish

The gilgie (Cherax quinquecarinatus) is among the more widespread of the six endemic south‐western Australian freshwater crayfish species. In the present study, the phylogeographic structure of the gilgie was investigated across its distribution to determine whether patterns reflected those identified earlier in a co‐distributed congeneric, the koonac (Cherax preissii). Gilgies were sampled from 20 localities, a 412‐bp fragment of the cytochrome c oxidase subunit I mitochondrial DNA gene was amplified from 75 individuals, and allozyme variation was assayed at nine loci. As in the koonac, three geographically‐restricted lineages were identified: from the north‐western, southern coastal, and intermediate/south‐western regions. Phylogeographic breaks appeared to be congruent with those in the koonac. The extent of genetic differentiation among lineages was comparable to that in the koonac, suggesting temporal congruence of the historical events responsible for the observed structure. A relaxed Bayesian molecular clock suggested that the major clades and lineages in each species diverged in the Late Miocene–Early Pliocene (4.0–9.6 Myr ago), possibly resulting from increasing pulses of aridity. The retrieval of almost‐identical phylogeographic structure in two co‐distributed species suggests that biogeographic regions can be more accurately defined in south‐western Australia. With the geographic fidelity of these lineages, the present data also provide evidence of the translocation of a single individual from the north‐west to the south coast. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101 , 385–402.     

Development of the female reproductive system in the freshwater crayfish Cherax quadricarinatus (Decapoda, Parastacidae)

Abstract. The differentiation of the female reproductive system from a macroscopic and microscopic point of view was studied in Cherax quadricarinatus. For this characterization, 184 females were dissected and processed for the histological analysis. From the differentiation of the ovary up to its maturity, three ovarian morphotypes could be distinguished macroscopically: parallel strands without any contact between them, an H-shaped ovary, and a Y-shaped ovary. These morphotypes were compared within the Astacida. Four ovarian developmental stages were recognized based on ovary color, and the histological structure and relative proportion of cellular types. The post-spawning ovary was also characterized. The components of the female reproductive system sheath were described and its modifications in the ovary and oviducts were determined and compared. Theoretical aspects of the study of sexual differentiation in C. quadricarinatus were discussed within a phylogenetic framework.     

Parallel adaptive radiations in arid and temperate Australia: molecular phylogeography and systematics of the Egernia whitii (Lacertilia: Scincidae) species group

It has been an enduring belief that increasing aridity combined with Pliocene‐Pleistocene glacial cycles resulted in the formation of distinct arid zone and temperate zone faunas within Australia. We assembled a molecular phylogeny for the Egernia whitii species group, an endemic group of skinks that comprises representatives from arid and temperate Australia, in order to test several biogeographical hypotheses regarding the origin of the Australian arid zone fauna. Sequence data were obtained from ten of the 11 species within the species group, plus three other Egernia species and an outgroup (Eulamprus heatwolei). We targeted portions of the ND4 (696 base pairs) and 16S rRNA (500 bp) mitochondrial genes and the β‐Fibrinogen 7th Intron nuclear gene (648 bp). The edited alignment comprised 1844 characters, of which 551 (30%) were variable and 382 (69%) were parsimony informative. We analysed the data using maximum likelihood and Bayesian techniques and produced a single optimal tree. Our phylogeny strongly supports two major clades within the species group, corresponding to temperate‐adapted rock‐dwelling species and arid‐adapted obligate burrowing species. However, the phylogenetic affinities of E. pulchra were not resolved. Our topology indicates that the New South Wales population of E. margaretae is actually E. whitii and reveals that E. margaretae margaretae and E. m. personata are distinct species. There also appears to be a major phylogeographical break within E. whitii occurring in eastern Victoria. Although our data supported several previously proposed phylogenetic relationships, Shimodaira–Hasegawa tests soundly rejected several suggested affinities between certain species. The arid zone members of the E. whitii species group had been suggested to have originated as a result of multiple periods of colonization during the Pleistocene glaciation cycles. However, our genetic data suggest a single origin (presumably from a semiarid E. multiscutata‐like ancestor) for the arid zone members of the group prior to the Plio‐Pleistocene, probably during the late Miocene to early Pliocene. Our topology displays substantial sequence divergence between species with short internodes and long terminal branches, indicating rapid adaptive radiations within the arid and temperate zones. The presence of temperate‐adapted species within more mesic refugia of the arid zone suggests that the necessary adaptations to aridity for colonizing the dry interior of the continent have not evolved since the initial period of adaptive radiation, despite the long evolutionary history of the species group. © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 83 , 157–173.     

Phylogeography of the widespread Australian freshwater prawn, Macrobrachium australiense (Decapoda, Palaemonidae)

Aim To investigate the phylogeographic structure of the widespread freshwater prawn, Macrobrachium australiense, within and between major Australian drainage basins using mitochondrial sequence data. This will enable the investigation of historical connections between major drainages and examination of hypotheses of biogeographic associations among Australian freshwater basins. Location Inland, eastern and northern Australia. Methods Sequencing 16S rRNA and ATPase 6 protein coding mitochondrial DNA genes from M. australiense from 19 locations from inland, eastern and northern Australia. Results Within drainage basins, haplotype trees are monophyletic, with the exception of the Finke River from the Lake Eyre Basin. Macrobrachium australiense from the two main inland drainages, the Murray–Darling and Lake Eyre Basin are divergent from each other and do not form a monophyletic group, instead the Murray–Darling Basin haplotypes clade with eastern coastal haplotypes. Haplotypes from neighbouring eastern coastal drainages were found to be quite divergent from each other. Main conclusions The phylogeographic relationships among M. australiense suggest that the two major inland drainages, the Murray–Darling Basin and the Lake Eyre Basin, are not biogeographically closely associated to each other. Instead the Murray–Darling Basin is more closely allied with the eastern coastal drainages across the Great Dividing Range. Despite their proximity the neighbouring southeast Queensland coastal Mary and Brisbane Rivers are also biogeographically divergent from each other. The results also indicate that the Finke River appears to have been isolated from the remainder of the Lake Eyre Basin catchment for a significant period of time.     

Aggressive interactions between three species of freshwater crayfish of the genus Cherax (Decapoda: Parastacidae)

The freshwater crayfish, Cherax destructor Clark, native to southeastern Australia, was first introduced to farm dams in southwestern Western Australia in 1932. The geographic range of the crayfish in Western Australia has increased substantially since then, and in recent years it has become established in natural waterways where it co-occurs with species of freshwater crayfish endemic to southwestern Australia, Cherax cainii and Cherax quinquecarinatus. The potential for competitive exclusion of these endemic species by C. destructor was investigated through laboratory experiments measuring aggressive behaviour. Body mass and species were found to be important factors governing aggressive dominance between C. cainii and C. destructor, with C. cainii winning significantly more interactions only when they were larger in body mass than their opponent. In trials between C. quinquecarinatus and C. destructor of similar body mass, there was no difference between the number of interactions 'won' by the two species. The implications for natural populations are discussed.     

Phylogeography and conservation genetics of endangered European Margaritiferidae (Bivalvia: Unionoidea)

Margaritifera margaritifera and M. auricularia are among the most endangered freshwater mussels in the world, and the only species of the genus found in Europe. Our genetic study explores allozymic variability (27 loci) and differentiation at the mitochondrial sequence level (partial COI and 16S rRNA gene sequences). The Spanish M. auricularia population showed genetic parameters of variation that were of the same order as those of other freshwater molluscs (though at the lower end of the range), probably permitting its potential recovery. The difference between this species and M. margaritifera was clearly established (ten diagnostic allozymic loci, Nei = 0.462, and mean nucleotide divergence around 9.4%). The M. margaritifera populations analysed showed a certain degree of population genetic structure (according to allozyme data) that was not, however, related to a geographical cline. Nevertheless, two mitochondrial lineages (albeit very closely related) were identified: a northern lineage extending from Ireland to the Kola Peninsula including the western Atlantic coast, and a second cluster distributed from Ireland to the Iberian Peninsula. The phylogenetic relationships between these two species and other related taxa were established. The putative M. m. durrovensis could be considered an 'ecophenotype'. Palaeobiogeographical scenarios are presented and indicate unexpected 'recent' gene flow between M. margaritifera populations that were theoretically isolated in the early Tertiary.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society 2003, 78, 235–252.     

Sea‐level changes and palaeo‐ranges: reconstruction of ancient shorelines and river drainages and the phylogeography of the Australian land crayfish Engaeus sericatus Clark (Decapoda: Parastacidae)

Historical sea levels have been influential in shaping the phylogeography of freshwater‐limited taxa via palaeodrainage and palaeoshoreline connections. In this study, we demonstrate an approach to phylogeographic analysis incorporating historical sea‐level information in a nested clade phylogeographic analysis (NCPA) framework, using burrowing freshwater crayfish as the model organism. Our study area focuses on the Bass Strait region of southeastern Australia, which is marine region encompassing a shallow seabed that has emerged as a land bridge during glacial cycles connecting mainland Australia and Tasmania. Bathymetric data were analysed using Geographical Information Systems (GIS) to delineate a palaeodrainage model when the palaeocoastline was 150 m below present‐day sea level. Such sea levels occurred at least twice in the past 500 000 years, perhaps more often or of larger magnitude within the last 10 million years, linking Victoria and Tasmania. Inter‐locality distance measures confined to the palaeodrainage network were incorporated into an NCPA of crayfish (Engaeus sericatus Clark 1936) mitochondrial 16S rDNA haplotypes. The results were then compared to NCPAs using present‐day river drainages and traditional great‐circle distance measures. NCPA inferences were cross‐examined using frequentist and Bayesian procedures in the context of geomorphological and historical sea‐level data. We found distribution of present‐day genetic variation in E. sericatus to be partly explained not only by connectivity through palaeodrainages but also via present‐day drainages or overland (great circle) routes. We recommend that future studies consider all three of these distance measures, especially for studies of coastally distributed species.     

The molecular systematics of Leiopotherapon unicolor (Günther, 1859): testing for cryptic speciation in Australia's most widespread freshwater fish

Leiopotherapon unicolor is the most widespread freshwater fish species in Australia. A comprehensive allozyme and mitochondrial DNA 16S rRNA data set was assembled from 141 specimens of L. unicolor collected Australia-wide in order to test for cryptic speciation in this far-ranging species. Surprisingly, little genetic diversity was observed within L. unicolor and provided no evidence for the existence of cryptic species within this lineage. In contrast, a small sample set of L. aheneus used as the outgroup showed two highly divergent haplotypes strongly suggestive of cryptic speciation. L. unicolor has a number of ecological and life history attributes that may explain the lack of significant genetic divergence over substantial geographical distances. The occurrence of other widespread fish and crustacean species that also display only limited genetic diversity indicate that climate conditions more favourable to dispersal across central and northern Australia than is suggested by the extent of present-day aridity have occurred in the relatively recent geological past.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 537–552.     

Phylogeographic patterning in a freshwater crab species (Decapoda: Potamonautidae: Potamonautes) reveals the signature of historical climatic oscillations

Aim The phylogeographic relationships among populations of the common Cape River crab, Potamonautes perlatus, are examined to investigate whether the contemporary population genetic structure is congruent with the hypothesized hydrographic evolution of drainage systems established during the Pliocene, or whether it reflects an older Miocene climatic amelioration. Location 139 samples of P. perlatus were collected from 31 populations distributed among the five major perennial drainage systems and a number of smaller catchments in the Western and Eastern Cape, South Africa. Methods Phylogeographic analysis using parsimony, maximum likelihood, minimum evolution and Bayesian inferences was employed for the 16S rRNA mtDNA gene region, while bootstrapping and posterior probabilities were used to assess the robustness of clades. In addition, nested clade analysis was performed in an attempt to disentangle the contemporary and historical factors that have sculpted genealogical relationships among conspecific populations of P. perlatus. Results Phylogenetic topologies were congruent irrespective of the evolutionary method employed. Two highly distinct reciprocally monophyletic clades characterized by marked levels of corrected sequence divergence were present, with no shared haplotypes between the two major phylogroups. Phylogroup one comprises the populations of the westward‐flowing drainages (mainly the Berg and Olifants drainages), and phylogroup two comprises all of the southward‐flowing drainages and can further be divided into two subclades – one containing the Breede River populations, and the other containing the Gamtoos and Gourits drainage systems. The nested clade analysis demonstrated restricted gene flow and long‐distance dispersal for a number of higher clade levels. The higher‐level groups and results for the total cladogram suggest either fragmentation or isolation by distance. Main conclusions Freshwater crabs are generally highly philopatric, and dispersal, although not common, has occurred historically. The westward‐flowing drainages (Berg, Olifants, Eerste, Liesbeeck and Tokai) are isolated from the southward‐flowing drainages by the Cape Fold Mountains, while the southward‐flowing drainages have a number of tributaries that extend into the low‐lying regions, allowing for gene flow between these three major drainages systems (Breede, Gamtoos and Gourits). Among the westward‐flowing drainages, a more intensive sampling regime is required to understand evolutionary relationships. Our molecular results suggest that the observed patterns pre‐date the formation of contemporary hydrographic patterns in the Cape. This suggests that an older Late Miocene event has severely impacted the contemporary population structure in this species, as recent Pliocene hydrographic boundaries do not correspond to the phylogeographic pattern observed. Conservation efforts for aquatic taxa should clearly be directed at the catchments, in an attempt to conserve biological diversity.     

南海野生斑节对虾的线粒体16S rRNA基因序列单倍型分析

采用聚合酶链式反应(PCR)技术对我国南海海域5个地点(三亚、深圳、阳江、湛江、北海)的野生斑节对虾100个个体的mtDNA 16S rRNA序列进行扩增,扩增产物经纯化后进行测序.用Clustal_X排序软件对所得的100个mtDNA 16S rRNA序列进行比对.通过ARLEQUIN软件对所得100个mtDNA 16S rRNA序列进行比较分析,共检测出28个变异位点,19种单倍型.19单倍型序列的碱基组成显示出较高的A T比例(69.0%).19种单倍型序列的遗传距离在0.002～0.033.根据构建的单倍型进化关系网,5个地点群体中的单倍型呈现一种混杂的分布格局.此外,根据单倍型在5个地点群体出现的频率和单倍型进化关系网,单倍型7是最为原始的单倍型,其他18种单倍型均起源于单倍型7.     

Phylogeography of the lacertid lizard, Psammodromus algirus, in Iberia and across the Strait of Gibraltar

Aim  To determine genetic substructuring within the lacertid lizard Psammodromus algirus . To compare levels of variation across a geological barrier, the Strait of Gibraltar, and to compare this against the known age of the barrier using a molecular clock hypothesis. To compare the effect of the barrier within this species with previously published data from other organisms.
Location  The Iberian Peninsula and North Africa.
Methods  Partial sequences from the mitochondrial cytochrome b , 12S rRNA and 16S rRNA genes were obtained from 101 specimens belonging to the subfamily Gallotiinae and used in this study. The data set was aligned using C lustal X and phylogenetic trees produced using both maximum-parsimony and maximum-likelihood methods. Maximum likelihood estimates of divergence times for the combined data set (12S + 16S + cytochrome b ) were obtained after discovery of lineage rate constancy across the tree using a likelihood ratio test.
Results  Psammodromus algirus contains divergent eastern and western mtDNA clades within the Iberian Peninsula. The western clade has northern and southern lineages in Iberia and one in North Africa. This phylogeographical pattern indicates that the lizard invaded North Africa after the opening of the Strait, presumably by natural rafting.
Main conclusions  As in several other species, current patterns of genetic diversity within P. algirus are not directly related to the opening of the Strait of Gibraltar. Widespread sampling on both sides of the barrier is necessary to determine its effect on species in this area accurately.     

Reconstruction of phylogenetic relationships within Grapsidae (Crustacea: Brachyura) and comparison of trans-isthmian versus amphi-atlantic gene flow based on mtDNA

Following taxonomic revisions in recent years, the originally large family Grapsidae MacLeay, 1838 has become a relatively small and morphologically homogeneous family in terms of adult and larval morphology. Most available molecular studies including more than one genus of the family have also suggested monophyly of the corresponding taxa. However, no single phylogenetic study has ever included all constituent genera of the Grapsidae. In the current study, a molecular phylogeny based on sequences of the mitochondrial 16S rRNA gene from all eight grapsid genera and 34 species is presented and suggests that up to four genera are not monophyletic. This is mainly due to the polyphyletic nature of the genus Pachygrapsus which can be found in six different lineages of the phylogeny, suggesting that the genus currently does not represent a single evolutionary lineage and is in need of taxonomic revision. Amphi-atlantic and trans-isthmian species pairs or populations in four genera are compared and reveal relatively constant and pronounced divergences across the Panama Isthmus as opposed to moderate divergences across the Atlantic Ocean, thereby suggesting occurrence of gene flow across the Atlantic Ocean during the past three million years.     

Endemic evolutionary radiation of Rhagada land snails (Pulmonata: Camaenidae) in a continental archipelago in northern Western Australia

In the Dampier Archipelago of Western Australia's Pilbara Region, several locally endemic, morphologically distinctive species of Rhagada land snails occur, contrasting with the morphologically conservative species with wider distributions on the adjacent mainland. To test alternative origins of this unusual local diversity in a continental archipelago, we examined sequences of the cytochrome oxidase subunit 1 and 16S mitochondrial genes in 22 described species and eight undescribed forms, including all known morphospecies from the Pilbara Region's Dampier Archipelago and adjacent mainland. Phylogenetic analyses consistently resolved four, deep clades within the Pilbara Region, with a mean sequence divergence of 15–18%. All but one of the species from the Dampier Archipelago formed one of the major clades, indicating that the morphological radiation in the archipelago evolved locally, rather than through multiple, relictual mainland lineages. Morphological divergence spanning almost that of the entire genus was within a subclade with sequence divergence < 4%, highlighting the disconnection between morphological diversification and levels of molecular genetic divergence. This in situ morphological radiation in the Dampier Archipelago, which transcends variation seen over much larger distances on the mainland, is unusual for a continental archipelago, and may relate to local heterogeneity of land forms. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 316–327.     

Phylogeography of postglacial range expansion in Nigronia serricornis Say (Megaloptera: Corydalidae)

We have examined the effects of post-Wisconsinan glacial range expansion on the phylogeography of the saw-combed fishfly, Nigronia serricornis Say (Megaloptera: Corydalidae) because aquatic insects are under-represented in postglacial studies (and in phylogeography in general), and because the effects of ecological degradation on the population genetics of environmental indicator species like N. serricornis cannot be measured unless the underlying phylogeography is understood. Sequence data from a 630-base fragment of the mitochondrial cytochrome oxidase I (COI) gene were subjected to amova and nested clade analysis for 30 populations (n = 344) of N. serricornis. Both the amova and nested clade analysis revealed substantial population structure; 44.4% of the variance occured among populations. Three northward migrations are apparent: one from Tennessee into Illinois, Indiana, Wisconsin, Michigan and Ontario, a second that radiated eastward from Pennsylvania, and a third that moved along the coast from North Carolina into Connecticut, Massachusetts, Maine and then into New York. The latter two of these migrations were the result of contiguous range expansions, while the former expansion, out of Tennessee, appears to have been rapid with little gene flow from the source population. Additional clades included a group of haplotypes in central Kentucky that appear to have expanded along preglacial drainages, and clades in North Carolina and Georgia that have remained centrally located. Haplotype diversity decreased from south to north, a pattern that has been widely reported for animal and plant populations that expanded with the retreat of the Wisconsinan glaciation.     

Molecular phylogeny of the family Tephritidae (Insecta: Diptera): New insight from combined analysis of the mitochondrial 12S, 16S,and COII genes

The phylogeny of the family Tephritidae (Diptera: Tephritidae) was reconstructed from mitochondrial 12S, 16S, and COII gene fragments using 87 species, including 79 tephritid and 8 outgroup species. Minimum evolution and Bayesian trees suggested the following phylogenetic relationships: (1) A sister group relationship between Ortalotrypeta and Tachinisca, and their basal phylogenetic position within Tephritidae; (2) a sister group relationship between the tribe Acanthonevrini and Phytalmiini; (3) monophyly of Plioreocepta, Taomyia and an undescribed new genus, and their sister group relationship with the subfamily Tephritinae; (4) a possible sister group relationship of Cephalophysa and Adramini; and (5) reconfirmation of monophyly for Trypetini, Carpomyini, Tephritinae, and Dacinae. The combination of 12S, 16S, and COII data enabled resolution of phylogenetic relationships among the higher taxa of Tephritidae.     

Vitellogenin levels in hemolymph, ovary and hepatopancreas of the freshwater crayfish Cherax quadricarinatus (Decapoda: Parastacidae) during the reproductive cycle

The freshwater crayfish Cherax quadricarinatus is a tropical species of great interest for aquaculture. Vitellogenin (Vg), a lipoprotein precursor of the vitellum accumulated in spawned eggs, can be synthesized in the ovary and/or hepatopancreas of most crustaceans, being the hemolymph the way for transporting Vg throughout the reproductive cycle. Concentration of Vg in hemolymph, ovary and hepatopancreas of Cherax quadricarinatus adult females was measured by means of ELISA, specifically developed after purifying the native Vg. Measurements were made at four periods of the reproductive cycle: pre-reproductive, mid-reproductive, late reproductive and post-reproductive. Besides, both hepatosomatic (HSI) and gonadosomatic (GSI) indexes were determined in each period. Significant variations in Vg levels were detected in both hemolymph and hepatopancreas, being the highest values observed during the mid-reproductive period. Besides, such variations were positively correlated to the HSI. A positive correlation between Vg levels in hepatopancreas and ovary was also seen. These results support previous evidences about the central role of the hepatopancreas as a site of Vg synthesis in the studied species, together with the relevancy of hemolymph for transporting Vg from the hepatopancreas to the ovary. For aquaculture purposes, Vg monitoring in hemolymph could be used as a non-injurious method, to check the reproductive activity of C. quadricarinatus females.     

Convergent evolution of shell shape in freshwater limpets: the African genus Burnupia

Convergent evolution of shell shape is a well known phenomenon in gastropods that has caused much confusion in taxonomic and systematic studies. A paradigm is the patelliform shell shape in several taxa of the order Basommatophora. Historically, most freshwater limpets were assigned to the family Ancylidae. Based on anatomical data, some taxa were subsequently moved to different families. However, there are still doubts about the monophyly of the remaining ancylids. This is also true for the African limpets and particularly for species of the genus Burnupia Walker, 1912. In the present paper, two independent molecular markers (COI, 18S rRNA) are used to (a) infer the position of Burnupia within the Basommatophora and (b) to test whether the shell shape of Burnupia evolved independently. Bayesian and maximum likelihood analyses of 12 genera of limnic Basommatophora indicate that Burnupia not only appears to be distinct from the Ancylidae, but also from all other representatives of the superfamily Planorboidea studied here. Based on a generally well-supported phylogeny and preliminary anatomical data, it is concluded that patelliform shell shapes evolved at least three times in the taxa studied here and that the shell shape of Burnupia represents another case of convergent evolution. Previously proposed functional and evolutionary scenarios for the evolution of patelliform shell and their relevance for freshwater limpets are discussed.  © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society , 2004, 140 , 577–586.