Integration of cytochrome c oxidase I barcodes and geometric morphometrics to delimit species in the genus Gnopharmia (Lepidoptera: Geometridae,Ennominae)

We tested the hypothesis of species taxonomy in the genus Gnopharmia (Macariini, Ennominae) that was recently established in a review based on discrete morphological characters. For this objective we integrated both DNA‐based and morphometric approaches in order to infer species boundaries. A 658‐bp fragment of the mitochondrial cytochrome c oxidase subunit 1 (CO1) (DNA barcode) was analysed from populations of five species distributed throughout the Middle East to assess their consistency with traditionally defined morphospecies. Signals in the morphological variation of the aedeagus of all relevant populations were evaluated using geometric landmarks. Consistent groupings compatible with the current taxonomic classification were found with both approaches. The results strongly support the distinction of seven closely related species. © 2013 The Linnean Society of London     

Clinal variation or validation of a subspecies? A case study of the Graptemys nigrinoda complex (Testudines: Emydidae)

Widely distributed species often display intraspecific morphological variation due to the abiotic and biotic gradients experienced across their ranges. Historically, in many vertebrate taxa, such as birds and reptiles, these morphological differences within a species were used to delimit subspecies. Graptemys nigrinoda is an aquatic turtle species endemic to the Mobile Bay Basin. Colour pattern and morphological variability were used to describe a subspecies (G. n. delticola) from the lower reaches of the system, although it and the nominate subspecies also reportedly intergrade over a large portion of the range. Other researchers have suggested that these morphological differences merely reflect clinal variation. Our molecular data (mtDNA) did not support the existence of the subspecies, as the haplotypes were differentiated by only a few base pairs and one haplotype was shared between the putative subspecies. While there were significant morphological and pattern differences among putative specimens of G. n. nigrinoda, G. n. delticola and G. n. nigrinoda × delticola, these differences probably represent clinal variation as they were also related to environmental variables [i.e. cumulative drainage area and drainage (categorical)]. Specimens occupying slow‐current, high‐turbidity river reaches (e.g. the Tensaw River) exhibited greater relative carapace heights and more dark pigmentation, while specimens occupying fast‐current, clearer rivers (e.g. the upper Alabama, Cahaba and Tallapoosa rivers) exhibited lower carapace heights and more yellow pigmentation. Given the absence of clear molecular and morphological differences that are related to drainage characteristics, we suggest that there is not sufficient evidence for the recognition of G. n. delticola as a distinct subspecies. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 810–822.     

Populations composed entirely of hybrid colonies: bidirectional hybridization and polyandry in harvester ants

In eusocial Hymenoptera, haplodiploid life cycles, obligate sterile castes, and polyandry may facilitate selection for hybridization. We analyzed a broad hybrid zone between the ecologically distinct seed‐harvester ants Pogonomyrmex occidentalis (Cresson) and Pogonomyrmex maricopa (Wheeler) using mitochondrial (mt)DNA sequence data, eight morphological markers, and 14 random amplified polymorphic DNA (RAPD) markers. Average mtDNA sequence divergence among parental species was 11.34%, indicating secondary contact. RAPD markers were significantly correlated with morphological variation, confirming the interspecific hybrid origin of all morphologically putative hybrid colonies. A morphological hybrid index indicates an abundance of both F₁ hybrids and parental morphotypes within colonies. Individual character frequencies plotted against distance show coincident and concordant clines, suggesting little to no introgression. The structure of the hybrid zone is two‐fold. Within the western region, stark reversals in character frequencies coincide with overt soil differences, indicating a mosaic hybrid zone structure. The eastern region is a riparian habitat where four adjacent populations were composed entirely of hybrid colonies. These habitat associations suggest that hybrid worker genomes permit dispersal into intermediate environments that select against one or both parental species. The present study suggests that, in addition to retaining reproductive compatibility, ecologically distinct species of ants may generate hybrid colonies maintained by environmental selection. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 320–336.     

Delimiting species in the genus Otospermophilus (Rodentia: Sciuridae), using genetics,ecology, and morphology

We apply an integrative taxonomy approach to delimit species of ground squirrels in the genus Otospermophilus because the diverse evolutionary histories of organisms shape the existence of taxonomic characters. Previous studies of mitochondrial DNA from this group recovered three divergent lineages within Otospermophilus beecheyi separated into northern, central, and southern geographical populations, with Otospermophilus atricapillus nested within the southern lineage of O. beecheyi. To further evaluate species boundaries within this complex, we collected additional genetic data (one mitochondrial locus, 11 microsatellite markers, and 11 nuclear loci), environmental data (eight bioclimatic variables), and morphological data (23 skull measurements). We used the maximum number of possible taxa (O. atricapillus, Northern O. beecheyi, Central O. beecheyi, and Southern O. beecheyi) as our operational taxonomic units (OTUs) and examined patterns of divergence between these OTUs. Phenotypic measures (both environmental and morphological) showed little differentiation among OTUs. By contrast, all genetic datasets supported the evolutionary independence of Northern O. beecheyi, although they were less consistent in their support for other OTUs as distinct species. Based on these data, we support the conclusions from a previous study that synonymized O. atricapillus with O. beecheyi, and we elevate the northern lineage of O. beecheyi to a separate species. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 1136–1151.     

Genetic and chaetal variation in Nais worms (Annelida,Clitellata, Naididae)

The genus Nais is a group of oligochaetous clitellates, common in eutrophic freshwater habitats. About 30 species are described. Species identification is based primarily on chaetal characters, which are often subtle, inconsistent, and even overlapping between nominal species. We investigated the correlation between genetic variation and chaetal morphology in this genus. Eighty‐one individuals from Europe, North America, and China were included in the study. Seventy‐five of these were preserved as vouchers. They were scrutinized with regard to chaetal morphology, and ten different morphotypes were identified. Three molecular markers, two mitochondrial (the COI gene and 16S rDNA) and one nuclear (the ITS region), were used to establish the genetic lineages in the material. Genetic variation was found to be largely congruent with chaetal character patterns. However, at least nine separately evolving lineages (all supported by mitochondrial as well as nuclear data) correspond to at most six nominal species. Four morphotypes/lineages are recognized as Nais barbata, Nais christinae, Nais elinguis, and Nais stolci, respectively, whereas five, or possibly more, lineages represent a morphological continuum covering the variation of the Nais communis/variabilis complex. Thus, cryptic speciation is revealed. Our results indicate that a taxonomic revision of the genus will be needed in the future.     

Genetic variation in the mitochondrial 16S rRNA gene of the American dog tick,Dermacentor variabilis (Acari: Ixodidae)

Genetic variation in the mitochondrial (mt) 16S ribosomal RNA (rRNA) gene was examined for the American dog tick, Dermacentor variabilis (Say, 1821). Nine different haplotypes were detected among 369 adult D. variabilis collected from four localities in Canada. There were eight variable nucleotide positions in the 404 bp sequence alignment. Individuals of haplotype 1 occurred at frequency of >75% at all localities. Five haplotypes were detected at only one of the four localities. High haplotype diversity and low nucleotide diversity, combined with significantly negative F_s values for ticks at three localities, suggest a recent population expansion. Genetic differences were found between populations at different localities, but a Mantel regression analysis revealed no association between genetic differences and geographical distances. There was also no association between tick haplotype and the prevalence of the bacterium, Rickettsia montanensis Weiss and Moulder, 1984, in D. variabilis among localities or on opposite sides of Blackstrap Lake (Saskatchewan). The 16S rDNA haplotypes from Canadian populations of D. variabilis formed a clade with those from the eastern and central U.S.A., to the exclusion of D. variabilis from geographically isolated populations in the western U.S.A. Although sample sizes for D. variabilis in the eastern U.S.A. are small, there may be genetic divergence between populations in Canada and those in the eastern U.S.A., which may have implications for studies on the pathogenic agents transmitted by D. variabilis to its hosts.     

Phylogenetic relationships in the Sceloporus variabilis (Squamata: Phrynosomatidae) complex based on three molecular markers,continuous characters and geometric morphometric data

The monophyly of the Sceloporus variabilis group is well established with five species and two species complexes, but phylogenetic relationships within species complexes are still uncertain. We studied 278 specimens in 20 terminals to sample all taxa in the “variabilis group,” including three subspecies in the “variabilis complex,” and two outgroups (Sceloporus grammicus and Sceloporus megalepidurus). We assembled an extensive morphological data set with discrete and continuous characters (distances and scale counts), including geometric morphometric data (landmark coordinates of three shapes), and a three‐marker molecular data set as well (ND4, 12S and RAG1). We conducted parsimony and Bayesian phylogenetic inferences on these data, including several partitioning and weighting schemes. We suggest elevating three subspecies to full species status. Therefore, we recommend recognition of nine species in the “variabilis group.” First, S. variabilis is sister to Sceloporus teapensis. In turn, Sceloporus cozumelae is sister to Sceloporus olloporus. These four species are a monophyletic group, which is sister to Sceloporus smithi. Finally, Sceloporus marmoratus is sister of the clade of five species. The other species in the “variabilis group” (Sceloporus chrysostictus, Sceloporus couchii and Sceloporus parvus) are a paraphyletic grade at the base of the tree. Our analyses reject the existence of the “variabilis complex.” We conducted a parsimony‐based ancestral reconstruction on body size (snout–vent length), femoral pores and dorsal scales and related morphological changes to geographic distribution of the species. Our phylogenetic hypothesis will allow best designs of comparative studies with species in the “variabilis group,” one of the earliest divergent lineages in the genus.     

Genetic and morphological variation in two littorinid gastropods: evidence for recent population expansions along the East African coast

In marine species, population diversity and differentiation is affected by the population history and by the complex interaction between oceanographic dynamics and ecological traits. In the present study, we examined two species of marine gastropods (the mangrove periwinkle Littoraria scabra and the rocky shore Littoraria glabrata) along the East African coast, using both genetic and geometric morphometric methods. We report a greater variation of shell shape in L. scabra compared to the slightly smaller variation in L. glabrata. This variation was probably associated with variation of environmental factors along the coast, such as temperature and hydrodynamics. Despite morphological variation, we found low mitochondrial genetic differentiation among samples from different localities for both species, which is probably a consequence of the ongoing gene flow during the free‐swimming larval stage of these gastropods. Additionally, high levels of haplotype diversity, low nucleotide diversity, and ‘star‐like’ genealogies were found in both species. These observations and the results from mismatch distributions, indicate a possible signature of recent population expansions in both species, which probably started during interglacial periods of the Pleistocene and led to the colonization of the Indian Ocean coast. © 2013 The Linnean Society of London     

Species delimitation and phylogeography of the Abies chensiensis complex inferred from morphological and molecular data

Species delimitation in the Abies chensiensis complex, consisting of A. chensiensis, A. recurvata, A. ernestii and A. ernestii var. salouenensis, has been the subject of a long‐term dispute, as various taxonomic combinations have been proposed. We combined different lines of evidence, including morphological characters and mitochondrial and plastid DNA sequence data, to assess species delimitation. Plastid DNA sequence variation was highly consistent with the morphological result and separated A. recurvata from the other three taxa, but the mitochondrial DNA genealogy was more strongly related to geographical distribution rather than species delimitation, as proposed in previous studies. On the basis of morphological characters and plastid DNA sequence variation, the current species status of A. recurvata was well supported, and we also accepted A. ernestii and A. ernestii var. salouenensis as varieties of A. chensiensis. Furthermore, the phylogeographical history of A. chensiensis was surveyed by molecular data and ecological niche modelling. On the basis of molecular data, the current phylogeographical pattern of the A. chensiensis complex was probably shaped by recent rapid expansions from multiple refugia in the Hengduan Mountains and habitat contraction after the last glaciation. This hypothesis was supported by the results of ecological niche modelling and previous fossil records. Our findings suggest that these fir taxa might have continued to expand their range after the largest glaciation until the end of the last glaciation, but have contracted since then with the increasing temperature and habitat changes. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 177 , 175–188.     

Limited gene exchange between two sister species of leaf beetles within a hybrid zone in the Alps

Analysing genomic variation within and between sister species is a first step towards understanding species boundaries. We focused on two sister species of cold‐resistant leaf beetles, Gonioctena quinquepunctata and G. intermedia, whose ranges overlap in the Alps. A previous study of DNA sequence variation had revealed multiple instances of mitochondrial genome introgression in this region, suggesting recent hybridization between the two species. To evaluate the extent of gene exchange resulting from these hybridization events, we sampled individuals of both species inside and outside the hybrid zone and analysed genomic variation among them using RAD‐seq markers. Individual levels of introgression in the nuclear genome were estimated first by defining species‐specific SNPs (displaying a fixed difference between species) a priori and second by using model‐based methods. Both types of analyses indicated little gene exchange, if any, between species at the level of the nuclear genome. Whereas the first method suggested slightly more gene flow, we argue that it has likely overestimated introgression in the phylogeographic context of this study. We conclude that strong intrinsic barriers prevent genetic exchange at the level of the nuclear genome between the two species. The apparent discrepancy observed between introgression occurring in the nuclear and mitochondrial genomes could be explained by selection acting in favour of the latter. Also, these results have consequences for the phylogeographic study of each species, since we can assume that genetic diversity in the overlapping portion of their ranges is not the product of introgression.     

Phylogeography of an Italian endemic salamander (genus Salamandrina): glacial refugia,postglacial expansions,and secondary contact

The Italian endemic genus Salamandrina has been historically regarded as monotypic but, recently, studies based on both mitochondrial and nuclear markers have indicated the existence of two distinct species of spectacled salamanders: Salamandrina perspicillata, in central and northern Italy, and Salamandrina terdigitata, in southern Italy. We analyzed nucleotide variation at mitochondrial and nuclear genes [cytochrome b, 12S and 16S rRNA, recombination activating gene (RAG 1)] in 223 individuals from 56 locations, aiming to investigate their genetic structure and recent evolutionary histories. Phylogenetic and phylogeographical analyses revealed the existence of three and two genetically distinct groups of populations in northern and southern salamander, respectively. Historical demographic analyses led to the inference of range expansion for both species in the late Pleistocene. During the last glacial stage, each salamander survived in a single refugium, namely the southern in Calabria and the northern in central Italy. At the end of this period, both lineages expanded northward and established secondary contact. Spatial distribution of RAG 1 haplotype variation revealed two differentiated population groups corresponding to the major mitochondrial (mt)DNA clades. Nuclear pattern of introgressive hybridization was more extensive than the highly limited introgression of mtDNA markers. From a conservation standpoint, southern Latium and Calabria proved to be the major genetic diversity reservoirs, thus deserving particular conservation efforts. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 903–922.     

Morphometric analysis and taxonomic revision of the Vriesea paraibica complex (Bromeliaceae)

The species related to Vriesea paraibica (Bromeliaceae, Tillandsioideae) have controversial taxonomic limits. For several decades, this group has been identified in herbarium collections as V. × morreniana, an artificial hybrid that does not grow in natural habitats. The aim of this study was to assess the morphological variation in the V. paraibica complex through morphometric analyses of natural populations. Two sets of analyses were performed: the first involved six natural populations (G1) and the second was carried out on taxa that emerged from the first analysis, but using material from herbarium collections (G2). Univariate ANOVA was used, as well as discriminant analysis of 16 morphometric variables in G1 and 18 in G2. The results of the analyses of the two groups were similar and led to the selection of diagnostic traits of four species. Lengths of the lower and median floral bracts were significant for the separation of red and yellow floral bracts. Vriesea paraibica and V. interrogatoria have red bracts; these two species are differentiated by the widths of the lower and median portions of the inflorescence and by scape length. These structures are larger in the former and smaller in the latter. Of the species with yellow floral bracts, V. eltoniana is distinguished by longer leaf blades and scapes and V. flava is characterized by its shorter sepal lengths. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 159 , 163–181.     

Morphological variation versus genetic divergence: a taxonomic implication for Mogannia species (Cicadidae: Cicadinae)

Species showing intraspecific morphological variation tend to be very difficult to identify using morphological characters. One such example is the cicada genus Mogannia where some species show considerable intraspecific variation mainly exhibited by wing pattern and body colouration. Thirty-one variants covering different putative species of Mogannia were recognized and illustrated in the present paper. Molecular data of mitochondrial COI and Cytb sequences were employed to test the level of variation and phylogeny of them. The existence of a ‘barcoding gap’ between intraspecific and interspecific genetic divergences and the reciprocally monophyletic clades indicate that all the closely related variants represent a single species, and that all these variants correspond to six species, respectively. However, the evolutionary relationships of intraspecific variants are not resolved possibly due to insufficient genetic variation among them. Our results indicated that some morphological characters, especially the wing pattern and body colouration, and even the number of apical processes of the aedeagus in a couple of related species, must be used with great caution in delimiting Mogannia species and their relatives. The factors responsible for intraspecific morphological variation and phylogeny of Mogannia spp. are preliminarily discussed.     

Genets (Carnivora, Viverridae) in Africa: an evolutionary synthesis based on cytochrome b sequences and morphological characters

The taxonomy of the genets (genus Genetta) has long been discussed, thus hampering endeavours towards evolutionary reconstruction. Sequence data from the complete cytochrome b gene (cyt b) were generated for 50 specimens representing 15 morphological species in order to allow the production of the first exhaustive molecular phylogeny of the genets. Second, a revised morphological matrix comprising 50 characters was combined with the cyt b data to estimate the level of morphological homoplasy. Phylogenetic analyses were conducted using parsimony, maximum likelihood and Bayesian procedures. Our results based on cyt b contradict a part of the traditional taxonomy of genus Genetta, the servaline and small‐spotted genets being paraphyletic, but confirmed the species status recently re‐investigated for three genets belonging to the large‐spotted complex, including the newly described G. bourloni. The combined analysis yielded similar results although morphological characters were clearly homoplasic. Partitioned Bremer supports indicated conflicting signals between the two data sets throughout the tree, and species‐diagnostic characters, useful for delimiting species boundaries, were significantly correlated to habitat. However, morphological data supported the monophyly of clades (G. victoriae, other genets) (G. servalina, G. cristata), large‐spotted genet complex and forest forms. Our results suggest a complex evolutionary history of the genets in Africa, with a Poiana‐like ancestor inhabiting rain forest, and then a diversification involving two independent invasions of open habitats and one reversion to rain forest. Divergence estimates based on cyt b revealed that splitting events within genets partly follow a climatic speciation model during the cyclical periods of the Quaternary, although ‘primitive’ rain forest lineages diverged earlier, during the Late Miocene and Early Pliocene. © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 81 , 589–610.     

Taxonomic significance of leaf surface morphology in Aloe section Pictae (Xanthorrhoeaceae)

Leaf surface morphology was analysed in 32 species representing the maculate species complex (the poorly resolved section Pictae) in the genus Aloe (Xanthorrhoeaceae). Few comparative morphological data are available for the complex. Leaf surface and stomatal characters observed by scanning electron microscopy show taxonomically significant interspecific variation. Most species are characterized by irregularly outlined, four‐ to six‐sided epidermal cells, the periclinal walls of which are flat and embellished with micropapillae and the anticlinal walls of which are indicated by channels on the leaf surface. The outer stomatal pore is typically sunken or plane and surrounded by four lobes on the leaf surface that may overarch the epistomatal chamber. The guard cells have distinct outer and inner stomatal ledges. Two geographical groups, comprising southern and east African species, are distinguishable by their leaf surface morphology. These characters are diagnostic in A. ellenbeckii, A. prinslooi and A. suffulta and support changes in the delimitation of A. greatheadii, A. macrocarpa and A. swynnertonii. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 160 , 418–428.     

Hybridization between genetically and morphologically divergent forms of Rhagada (Gastropoda: Camaenidae) snails at a zone of secondary contact

Congeneric sympatry is rare in Australian camaenid land snails, and the reasons are poorly understood. As the first example of contact between species of the Western Australian genus Rhagada, we examined the parapatric zone between the coastally distributed Rhagada convicta and an undescribed inland species, which differ in size, shape, and banding of their shells and in their reproductive systems. Phylogenetic analysis of sequences of the mitochondrial 16S RNA gene indicated secondary contact between two distinct clades, with an average sequence divergence of 6.5%. Despite these substantial differences, the two forms hybridize in the narrow contact zone, resulting in intermediate phenotypes for shells and reproductive anatomy, and incongruence of the mitochondrial DNA (mtDNA) lineage and morphology. Hybridization appears to be asymmetric, however, as all morphological intermediates possessed the R. convicta mtDNA. Interbreeding between these deeply divergent forms challenges the use of reproductive structures as indicators of reproductive isolation in snail taxonomy. This is the first contact zone detected between mainland species of Rhagada and indicates that the widespread pattern of geographical replacement is not simply a result of allopatric divergence without subsequent contact. Instead, the perspective for understanding the radiation and geographical distributions must include interactions between the species. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 348–362.     

Molecular phylogeny of the Eremias persica complex of the Iranian plateau (Reptilia: Lacertidae), based on mtDNA sequences

The Persian racerunner Eremias persica Blanford, 1875 is confined to the Iranian plateau, and forms one of the most widespread but rarely studied species of the family Lacertidae. With many local populations inhabiting a variety of habitats, and exhibiting considerable morphological, genetic, and ecological variations, it represents a species complex. We analysed sequences of mitochondrial cytochrome b and 12S ribosomal RNA (rRNA) genes derived from 13 geographically distant populations belonging to the E. persica complex. Using our knowledge of palaeogeographical events, a molecular clock was calibrated to assess the major events in fragmentation, radiation, and intraspecific variation. The sequence data strongly support a basal separation of the highland populations of western Iran from those of the open steppes and deserts, occurring in the east. The subsequent radiation, fragmentation, and evolution of these major assemblages have led to several discernable geographical lineages across the wide area of the Iranian plateau. The results indicate a middle‐Miocene origin for the clade as a whole. The first split, isolating the western and eastern clades, appears to have occurred 11–10 Mya. Further fragmentations and divergence within the major clades began about 8 Mya, with an evolutionary rate of 1.6% sequence divergence per million years among the lineages in the genes studied (combined data set). Molecular and morphological data strongly support a taxonomic revision of this species complex. At least four of the discovered clades should be raised to species, and two to subspecies, rank. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 158 , 641–660.     

Taxonomy and molecular phylogeny of the Asian Paraleucophenga Hendel (Diptera,Drosophilidae)

This paper reports on nine Asian species of the genus Paraleucophenga, of which four are new to science: Paraleucophenga brevipenis sp. nov. , Paraleucophenga hirtipenis sp. nov. , Paraleucophenga longiseta sp. nov. , and Paraleucophenga tanydactylia sp. nov. We also report on a new synonym, Paraleucophenga shanyinensis Chen & Toda, 1994 syn. nov. A key to all of the species examined, based on morphological data, is provided, together with a ‘molecular’ key to seven Paraleudophenga species based on DNA sequence data of the mitochondrial NADH dehydrogenase subunit 2 (ND2) gene. The phylogenetic relationships among seven Paraleucophenga species are reconstructed based on DNA sequences of the mitochondrial ND2 gene, using two Leucophenga species as outgroups. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 615–629.     

Phylogeography and phylogenetic relationships of Malagasy tree and ground boas

Three species of boid snakes are recognized in Madagascar, namely the genus Sanzinia (one species and two subspecies) and the genus Acrantophis (two species). In the present study, we studied the patterns of genetic variation of these species across Madagascar using a fragment of the mitochondrial 16S rRNA gene in 77 specimens. To support the phylogenetic relationships of the lineages identified, three further gene fragments (cytochrome b, 12S rRNA and c‐mos) were analyzed in a reduced but representative set of samples. The results obtained corroborate that the genus Sanzinia includes two highly divergent mitochondrial lineages that evolved independently from each other on the east versus the west side of Madagascar. Each of these lineages presents a further subdivision that separates northern from southern groups. The nuclear marker showed no variation among the Malagasy boas, indicating either very low substitution rates in this gene or relatively recent speciation events coupled with high mitochondrial substitution rates. Because the broad geographic sampling detected no admixture among haplotypic lineages within Sanzinia, it is hypothesized that these may represent distinct species. Deviant haplotypes of snakes morphologically similar to Acrantophis dumerili indicate that this taxon may be a complex of two species as well. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 640–652.     

Differences in population structure estimated within maternally‐ and paternally‐inherited forms of mitochondria in Lampsilis siliquoidea (Bivalvia: Unionidae)

Mussels in several orders possess two separate mitochondrial lineages: a standard female‐inherited form and one inherited only through males. This system of doubly uniparental inheritance (DUI) for mitochondrial genes provides an opportunity to compare the population structure of gene‐lineages passed either mother‐to‐daughter or father‐to‐son. In the present study, we contrast variation in the male and female haplotype lineages of the American freshwater mussel species, Lampsilis siliquoidea (sometimes called Lampsilis radiata luteola), throughout the Lake Erie, Ohio River, and upper Mississippi River watersheds, and contrast variation with the sequences obtained for the related species/subspecies Lampsilis radiata radiata from Maine. The genetic markers were fragments of the cytochrome c oxidase subunit I gene (COI), which occurs in both mitochondrial types, F (female) and M (male). High haplotype diversity was found in the two independent lineages, although purifying selection against amino acid change appeared to be stronger in the female than the male lineage. Phylogeographical patterns also varied between mitochondria passing through females and males. The female lineage exhibited more population structure, with the occurrence of private or nearly‐private haplotypes within two streams, and three others showed restricted haplotype distributions. By contrast to the F‐haplotypes, complex phylogenetic structure occurred for M‐haplotypes, yet this phylogenetic variation coincided with almost no geographical pattern within haplotypes. Basically, F‐haplotypes showed isolation, especially above physical barriers, whereas M‐haplotypes did not. A few individuals in the eastern Lake Erie watershed even possessed M‐haplotypes of an Atlantic Slope (L. radiata radiata) origin, although their F‐haplotypes were typical of Midwestern L. siliquoidea. The finding that mussels package sperm as spermatozuegmata, which float downstream, may underlie greater gene mobility in male‐inherited mitochondria. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 229–240.