An annotated catalogue of type specimens of the land snail genus Cyclophorus Monfort, 1810 (Caenogastropoda,Cyclophoridae) in the Natural History Museum,London

The collection of land caenogastropod snails in the genus Cyclophorus Monfort, 1810 housed in the Natural History Museum, London (NHM), includes 52 type lots. Lectotypes have been designated for 43 available species-level names to stabilize existing nomenclature, two previously designated lectotype, two holotypes, one paratype, one syntype, one possible syntype and two paralectotypes are also listed. A complete catalogue of the Cyclophorus types in NHM, London is provided for the first time.     

Biochemical assessment of the taxonomic diversity of the operculate land snail, Cyclophorus fulguratus (Gastropoda: Cyclophoridae), from Thailand

Allozyme variation was examined in 223 samples of the operculate land snail Cyclophorus fulguratus from 13 localities across three regions of Thailand. Using horizontal starch gel electrophoresis, 13 allozyme presumed loci (12 polymorphic) were screened. Heterozygosity was moderate in C. fulguratus (H_exp = 0.008–0.127) with a high genetic heterogeneity among samples (F_st = 0.734). Populations showed a greater genetic differentiation in central Thailand (F_st = 0.380) than in northeastern Thailand (F_st = 0.108), suggesting frequent gene flow among populations in northeastern Thailand. C. fulguratus exhibits a strong pattern of isolation by distance over the entire tested species range in Thailand and may potentially have been involved in an extensive local fragmentation. Results of the distance analysis revealed that large genetic divergence has occurred among the central, northeastern and eastern Thailand groups [D = 0.361–0.701], strongly suggesting populations from these three geographical regions may actually represent or else be evolving into separate species.     

Molecular Evidence for Cryptic Speciation in the Cyclophorus fulguratus (Pfeiffer, 1854) Species Complex (Caenogastropoda: Cyclophoridae) with Description of New Species

A high degree of intraspecific variation, both genetic and in shell morphology, of the operculate land snail Cyclophorus fulguratus (Pfeiffer, 1854) suggests that its classification as a single species warrants reconsideration. We sequenced two nuclear (18S and 28S) and two mitochondrial (16S and COI) genes of 46 C. fulguratus specimens and used them to estimate the phylogeny and to determine the validity of species boundaries. Molecular phylogenetic analyses revealed the presence of three lineages corresponding to three geographically disjunctive populations of C. fulguratus in Thailand. Likelihood tests of topologies significantly supported the non-monophyly of the C. fulguratus–complex and Bayesian species delimitation analysis significantly supported the potential representation as distinct species of these three lineages. Discriminant function analysis based on geometric-morphometrics of shell shape allowed for significant distinction of these three candidate species, although they revealed a considerable degree of overlap of shell shape reflecting their crypsis morphologically. The diagnostic characters are provided by color pattern, pattern of protoconch and pattern of jaw. In conclusion, the results support that the C. fulguratus s.l., as currently recognized, consists of three distinct species in Thailand: C. fulguratus s.s., C. rangunensis and C. abditus sp.nov., which are described herein.     

Genetic divergence in the Chorthippus parallelus species group (Orthoptera: Acrididae)

Specimens of Chorthippus parallelus parallelus, C. p. erythropus, C. montanus, C. curtipennis and C. dorsatus were examined electrophoretically. This analysis indicates that C. curtipennis is very closely related to C. p. parallelus and C.p. erythropus (Nei's I = 0.97) while C. montanus is more distantly related ( I = 0.88). The values are consistent with divergence of C. p. erythropus from C. p. parallelus since its separation by the Pyrenean ice sheet during the Pleistocene and of C. curtipennis since invasion of North America by C. parallelus at the same time. A much earlier divergence of C. montanus is implied.     

Genetic divergence among species of the section Strombocarpa,genusProsopis (Leguminosae)

We studied degree of variability and relationships among populations belonging to five species of the section Strombocarpa of the family Leguminosae, namelyProsopis ferox, P. torquata, P. pubescens, P. strombulifera andP. reptans, by isozyme electrophoresis. The genetic similarity betweenP. reptans andP. strombulifera is high enough for the two to be considered subspecies or geographic races ofP. strombulifera instead of good taxonomic species. As expected these species cluster withP. pubescens, all belonging to series Strombocarpae. This group is different isoenzymatically from the remaining species,P. torquata andP. ferox. Comparison of our results with those obtained earlier for species of the section Algarobia showed that genetic variability in the section Strombocarpa is lower than in the section Algarobia, while the species divergence is much higher. These results suggest different adaptive strategies between these sections.     

Genetic divergence and reproductive isolation in the Ochthebius (Calobius) complex (Coleoptera: Hydraenidae)

The reproductive isolation in hydrenid beetles of the Ochthebius complex was studied by analysing gene exchange in natural populations of O. quadricollis, Ochthebius sp. A and O. brevicollis steinbuehleri collected along the Mediterranean coasts. The ranges of these three species are largely allopatric, but sympatric areas were detected between contiguous taxa, ie, O. quadricollis and Ochthebius sp. A; Ochthebius sp. A and O. b. steinbuehleri. Three levels of reproductive isolation and genetic divergence were observed. One level involves extensive intraspecific genetic divergence within the biological species O. quadricollis, Ochthebius sp. A and O. brevicollis, associated with both physical barriers (eg, sea and sand stretches) and the low dispersal capacity of larvae and adults. The finding of transitional samples between the most differentiated population groups should indicate, however, that there is still some gene flow between the populations of the three taxa. Another level is found between Ochthebius sp. A and O. b. steinbuehleri, whose gene pools appear to be fairly distinct in spite of the fact that reproductive isolation is still incomplete: in their few syntopic sites, some F1 hybrids appeared indeed to have lower fitness, since backcrosses or recombinant genotypes were never observed. The final level in the evolution of reproductive isolation (full reproductive isolation) has been achieved by the species O. quadricollis and Ochthebius sp. A. No F1 or F(n) hybrids, nor backcrosses were found in their sympatric areas. The relative importance of ecological factors and evolutionary forces in the prevention of gene exchange between taxa are discussed.     

Genetic and morphological divergence in theAndrocymbium gramineum complex (Colchicaceae)

Narrow endemism is a poor predictor of genetic potential in the six taxa that conform theAndrocymbium gramineum complex, as shown by the unexpectedly high values of variability associated with ten allozymic loci and 13 biometrical variables. Although both levels are shown not to be correlated, variability is always concentrated within populations. This result, together with environmental, reproductive and historical data strongly suggests differentiation in local isolates, where changes can be quickly assimilated by stochastic processes. A key implication for conservation is that sampling within the largest population will save effort while neglecting only very low frequency alleles.     

Genetic subdivision and morphological variation in a freshwater snail species complex formerly referred to as Viviparus georgianus (Lea)

Genetic and morphological variation was studied in a brooding (ovoviviparous) and morphologically variable freshwater snail [ Viviparus georgianus (Lea)] species complex in southern Georgia and Florida. Eleven populations were clustered into three genetically isolated, allopatric groups characterized by 7 to 14 diagnostic loci out of the 38 loci examined. These allopatric groups were identified as V. georgianus (in eastern and southern Florida), V. limi (in the Ocblockonee River) and V. goodrichi (in the Florida panhandle). Nei's standard genetic distances between the species were large (0.23-0.52) compared to intraspecific distances (0.00-0.06). Hierarchical F-statistics for differentiation among sites within drainage systems (F_SD) for V. goodrichi and V. georgianus were large (0.519 and 0.387, respectively). The F_DT values (differentiation among drainage systems within the total area sampled) were negative or close to zero, so most of the intraspecific genetic differentiation was due to differences among populations within drianage systems, rather than to differences among systems. Canonical discriminant analysis of nine shell morphological measurements separated all three species with little overlap. Principal components analysis of the shell morphology revealed intraspecific variation among sites possibly due to environmental effects. This study demonstrates the marked genetic and morphological variation among and within drainage systems in freshwater prosobranchs.     

Genetic and morphological divergence within the Sebastes pachycephalus complex (Scorpaeniformes: Scorpaenidae)

Genetic and morphological divergence among the four subspecies in the Sebastes pachycephalus complex (S. pachycephalus pachycephalus, S. p. nigricans, S. p. nudus and S. p. chalcogrammus) was clarified. Principal coordinate analysis (PCoA) based on AFLP clearly divided 55 specimens of the complex into two groups, the S. p. pachycephalus?CS. p. nigricans group (P-Ni group) and the S. p. nudus?CS. p. chalcogrammus group (Nu-C group), although three specimens occupied intermediate positions. The minimum spanning network (MSN) based on partial sequences of the mitochondrial control region (mtCR) failed to separate either the P-Ni and Nu-C groups or the four subspecies into distinct clades, although restricted gene flow and genetic differentiation between the former were indicated by the F _ST estimation. Differences in morphological characters, including counts of pectoral fin rays and counts of dorsal fin spines lacking basal scales, were also evident between the two groups. However, little or no genetic or morphological difference was found between the two subspecies within each group. It was concluded that the P-Ni and Nu-C groups of the S. pachycephalus complex actually represent two different species, which is further supported by their sympatric distribution. Differences in dorsal body coloration and the presence or absence of brown spots on the ventral surface, which were formerly used to discriminate between four ??subspecies,?? may simply represent intraspecific variation. The three specimens occupying intermediate positions in the AFLP PCoA also occupied equivocal positions between the two species in the principal component analysis (PCA) based on morphometric characters, suggesting that they were hybrids between the two species. The star-shaped MSN of mtCR, which lacks distinct clades representing the two species, may be due to not only interspecific hybridization but also the sharing of ancestral haplotypes.     

Systematics of the Anopheles barbirostris species complex (Diptera: Culicidae: Anophelinae) in Thailand

This study deals with five species of the Barbirostris Complex of Anopheles subgenus Anopheles that are known to occur in Thailand. Three new species of the complex, A nopheles dissidens sp. nov. , A nopheles saeungae sp. nov. , and A nopheles wejchoochotei sp. nov. , are characterized and compared with Anopheles barbirostris van der Wulp and Anopheles campestris Reid based on specimens of molecularly identified progeny broods. For practical purposes, the five species are essentially isomorphic and can only be unequivocally identified from diagnostic mitochondrial and ribosomal DNA sequences. Based on overall morphological similarity, An. campestris is considered to be a member of the Barbirostris Complex rather than a separate member of the Barbirostris Subgroup. The molecular data, mitotic karyotypes, bionomics, and distributions of the species are reviewed and discussed. It is concluded that integrated molecular epidemiological studies of the complex throughout the Oriental Region are needed to unambiguously elucidate the individual species and their relation to disease. © 2015 The Linnean Society of London     

Genetic divergence within frog species is greater in topographically more complex regions

Most global hotspots of biodiversity and endemism are in montane regions. One explanation is that montane regions have intrinsically higher speciation rates than lowland regions because complex mountain topography and climate variation facilitate genetic isolation among populations. Here, we ask from an intraspecific perspective whether frog species whose haplotypes are connected by topographically/climatically complex regions display strong genetic isolation (greater scaled genetic distances), compared with species whose haplotypes are connected by less complex regions. We analysed published DNA sequences of several frog species from tropical Central and South America for the mitochondrial cob, cox1 and 16S rRNA genes. Pairwise genetic distances among haplotypes within each species were scaled to the geographic distances between each pair of haplotypes. Topographic complexity was positively correlated with scaled genetic distances, and isolation‐by‐resistance was supported only in species from more topographically complex regions. This suggests that heterogeneous topographies increase landscape resistance, which in turn favours the appearance of isolation‐by‐resistance. Moreover, we found that the potential barriers that restrict gene flow within species are more closely related to factors associated with temperature and topography than to precipitation.     

Genetic divergence and geographic variation in the deep-water Conus orbignyi complex (Mollusca: Conoidea)

Puillandre, N. et al. (2010) Genetic divergence and geographic variation in a deep-water cone lineage: molecular and morphological analyses of the Conus orbignyi complex (Mollusca: Conoidea).The cone snails (family Conidae) are a hyperdiverse lineage of venomous gastropods. Two standard markers, COI and ITS2, were used to define six genetically-divergent groups within a subclade of Conidae that includes Conus orbignyi; each of these was then evaluated based on their shell morphology. We conclude that three forms, previously regarded as subspecies of Conus orbignyi are distinct species, now recognized as Conus orbignyi, Conus elokismenos and Conus coriolisi. In addition, three additional species (Conus pseudorbignyi, Conus joliveti and Conus comatosa) belong to this clade. Some of the proposed species (e.g., Conus elokismenos) are possibly in turn complexes comprising multiple species. Groups such as Conidae illustrate the challenges generally faced in species delimitation in biodiverse lineages. In the case of the Conus orbignyi complex, not only are there definable, genetically divergent lineages, but also considerable geographic variation within each group. Our study suggests that an intensive analysis of multiple specimens within a single locality helps to minimize the confounding effects of geographic variation and can be a useful starting point for circumscribing different species within such a confusing complex.     

Genetic structure of the common terrestrial pulmonate snail,Cryptozona siamensis (Pfeiffer, 1856), in Thailand

Allozyme variation was examined in 432 specimens of the pulmonate snail, Cryptozona siamensis, from 24 localities across three regions of Thailand plus an additional population from Malaysia. Although this snail is a cosmopolitan species it was predicted that C. siamensis should exhibit a large degree of genetic subdivision across these regions due to a low level of inter-population gene flow. To test this hypothesis, 10 enzyme systems were screened using horizontal starch gel electrophoresis, from which 14 presumed allozyme loci were scored and 13 found to be polymorphic. For these 13 polymorphic loci the expected heterozygosity levels were moderate (H_exp = 0.036–0.183, mean = 0.101). Genetic differentiation among the samples was low, with a low F_ST value (0.254) and genetic distance (D = 0.000–0.124, mean = 0.034) and no significant isolation by distance (IBD) across the whole sampled range of Thailand or Thailand plus Malaysia. However, an IBD pattern in the population substructure in the central Thailand region was detected. The low F_ST and moderate H_exp values in various populations of C. siamensis suggests a high degree of gene flow/migration between populations across the three geographic regions of Thailand (northern, central and southern) as well as the sampled population in Malaysia, perhaps associated with the anthropochoric effect of snail dispersal. The apparent IBD in the central Thailand region could be the consequence of local dynamics though this region.     

Hidden Mediterranean biodiversity: molecular evidence for a cryptic species complex within the reef building vermetid gastropod Dendropoma petraeum (Mollusca: Caenogastropoda)

The reef building vermetid gastropod Dendropoma petraeum inhabits the warmest waters of the Mediterranean Sea and is considered a threatened marine species. The aim of this study was to characterize its genetic structure throughout its whole distribution range using mitochondrial and nuclear sequence data. Because of its sessile adult lifestyle and lack of a pelagic larval stage, we expected a markedly subdivided population structure with limited levels of gene flow. Fragments of the mitochondrial genes cytochrome c oxidase subunit I (COI) and 16S rRNA (16S), were sequenced, along with the nuclear ribosomal cluster (internal transcribed spacer; ITS) in specimens from 18 localities. Our analyses identified four highly distinct phylogroups separated by a mean divergence of > 14% according to the COI sequence data or > 9% according to 16S, but differing only slightly in morphology. The nuclear data (ITS) indicated a lower substitution rate (divergence among groups of around 1%). These large genetic distances among the four lineages clearly point to the existence of a cryptic species complex within D. petraeum comprising at least four species. Differences in the characteristics of intracapsular larval development and protoconch were also detected among these lineages. The allopatric distribution of these cryptic species supports a predominantly vicariant-based cladogenetic pattern for the genus Dendropoma in the Mediterranean.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 898–912.     

Cytological evidence for a complex of species within the taxon Bactrocera tau (Diptera: Tephritidae) in Thailand

Analysis of mitotic karyotypes of wild specimens of larvae of the Bactrocera tea-like fruit flies (Diptera: Tephritidae) in Thailand has revealed seven distinct chromosome forms, based on the amount and distribution of heterochromatin in sex chromosomes and autosomes. Such cytological differences are perfecdy correlated with morphological observations and molecular genetics data. These findings clearly suggest that B. tau is a cluster of at least seven closely related species temporarily designated as species A (= B. tau) , B, C, D, E, F and G. On die basis of the gross quantity of heterochromatin accumulation in the genome, three groups of mitotic karyotypes can be recognized. Group 1 comprises species A and E. Species E specifically occurs only in fruits of Strychnos thorelii while species A attacks many kinds of host plants. Group 2 contains species B, F and G. Species B has been found only in Siphonodon celastrineus fruit, whereas species F and G attack die same host species, the medically important plant, Hydnocarpus anthelminthicus , albeit in different localities. Group 3 includes species C and D, each of which comprises larger amounts of pericentric heterochromatin in all chromosomes than die other two groups. Hence, diese two species are cytologically remote from those of groups 1 and 2. Species C and D occur allopatrically but they attack the same host plant species, Momordica cochinchinensis. Thus, genetic differentiation at the chromosomal level to specific host plant species and geographic isolation seem to play an important role in speciation of members of the B. tau complex.     

Cryptic lineage diversity,body size divergence,and sympatry in a species complex of Australian lizards (Gehyra)

Understanding the joint evolutionary and ecological underpinnings of sympatry among close relatives remains a key challenge in biology. This problem can be addressed through joint phylogenomic and phenotypic analysis of complexes of closely related lineages within, and across, species and hence representing the speciation continuum. For a complex of tropical geckos from northern Australia—Gehyra nana and close relatives—we combine mtDNA phylogeography, exon‐capture sequencing, and morphological data to resolve independently evolving lineages and infer their divergence history and patterns of morphological evolution. Gehyra nana is found to include nine divergent lineages and is paraphyletic with four other species from the Kimberley region of north‐west Australia. Across these 13 taxa, 12 of which are restricted to rocky habitats, several lineages overlap geographically, including on the diverse Kimberley islands. Morphological evolution is dominated by body size shifts, and both body size and shape have evolved gradually across the group. However, larger body size shifts are observed among overlapping taxa than among closely related parapatric lineages of G. nana, and sympatric lineages are more divergent than expected at random. Whether elevated body size differences among sympatric lineages are due to ecological sorting or character displacement remains to be determined.     

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.     

Genetic differentiation of the endemic Baikalian mollusk Baicalia carinata (Mollusca: Caenogastropoda)

In gastropod mollusk Baicalia carinata Dybowski, 1875, sampled in different sites of the Lake Baikal, comparison of 81 sequences of internal transcribed spacer 1 (ITS1) located between the genes for 18S rRNA and 5.8S rRNA, and of 100 sequences of the fragment of mitochondrial cytochrome-c-oxidase subunit 1 gene (CO1) was performed. Molecular phylogenetic analysis showed that the endemic mollusk species studied formed at least two distinct populations, Southwestern and Eastern. Statistical significance of the species subdivision into two populations was evaluated using the Mann-Whitney rank test.     

A new species of the Fluviopupa group (Caenogastropoda: Tateidae) from north-east Queensland,Australia

Fluviopupa kessneri n. sp. is described from the Mulgrave River and one of its major tributaries, the Little Mulgrave River, in north-east Queensland. It is the first record of the Fluviopupa group from a continental area, other members of the genus being known only from Fiji, New Guinea and a few other Pacific Islands, including New Caledonia.     

石荠苎属的遗传分化与种间关系分析

杂交实验结果表明,石荠苎属(唇形科)中国产的7个种,即小花荠苎Mosla cavaleriei、石香薷M． chinensis、小鱼仙草M．dianthera、杭州石荠苎M．hangchouensis、疏花荠苎M．pauciflora、石荠苎M． scabra和苏州石荠苎M．soochouensis,彼此之间生殖隔离。这7个种中,疏花荠苎是四倍体,由杂交伴随多倍化产生。根据21个代表居群在15个酶系统的28个位点的等位酶资料分析,其余6个二倍体种组成3个亲缘关系密切的种对。杭州石荠苎和石香薷的亲缘关系最近,石香薷可能由于涉及繁育系统类型的大突变而与杭州石荠苎分化开来。小花荠苎与小鱼仙草比较接近,但它们的遗传分化很显著,除了突变之外,染色体结构也有大的变异,最明显的表现是随体的位置不同,小花荠苎的随体位于长臂上,而小鱼仙草与其它种一样,随体位于短臂上。石荠苎和苏州石荠苎在酶基因位点上的分化不大,但生境隔离、花期隔离和繁育系统的差别等能有效地保持种的独立性。不同的生境要求、不同的花期、花的形态结构的分化、繁育系统类型的不同、杂种F_l代高度不育、染色体数目和核型的差异等均为石荠苎属下种间有效的隔离机制,而且近缘种间常是多种机制共同起作用。