Phylogeographic structure, gene flow and species status in blue grouse (Dendragapus obscurus)

We investigated the genetic population structure and species status of a relatively sedentary bird that is a permanent resident of western North American forests, the blue grouse (Dendragapus obscurus). Phylogenetic analysis of complete mitochondrial control region DNA sequences resulted in the identification of three basal clades of haplotypes that were largely congruent with well-known biogeographical regions. These clades corresponded to the parapatric sooty (D. o. fuliginosus) and dusky (D. o. obscurus) subspecies groups of blue grouse plus a previously unrecognized division between northern and southern dusky grouse populations; the latter does not correspond closely to any currently recognized subspecies boundary. Approximately 66% of the total genetic variation was distributed among these three regions. Maximum likelihood estimates of gene flow between the regions were low or asymmetric; gene flow has been insufficient to prevent genetic divergence between dusky and sooty grouse. Estimates of gene flow among populations within sooty grouse were large except across the Columbia River valley. Among populations of dusky grouse, estimates of gene flow were heterogeneous and asymmetrical, reflecting large-scale fragmentation of the distribution due to landscape features and associated vegetation. Genetic, morphological and behavioural evidence suggest that sooty and dusky grouse are species-level taxa; the specific status of a third clade remains ambiguous.     

Differential influences of local subpopulations on regional diversity and differentiation for greater sage‐grouse (Centrocercus urophasianus)

The distribution of spatial genetic variation across a region can shape evolutionary dynamics and impact population persistence. Local population dynamics and among‐population dispersal rates are strong drivers of this spatial genetic variation, yet for many species we lack a clear understanding of how these population processes interact in space to shape within‐species genetic variation. Here, we used extensive genetic and demographic data from 10 subpopulations of greater sage‐grouse to parameterize a simulated approximate Bayesian computation (ABC) model and (i) test for regional differences in population density and dispersal rates for greater sage‐grouse subpopulations in Wyoming, and (ii) quantify how these differences impact subpopulation regional influence on genetic variation. We found a close match between observed and simulated data under our parameterized model and strong variation in density and dispersal rates across Wyoming. Sensitivity analyses suggested that changes in dispersal (via landscape resistance) had a greater influence on regional differentiation, whereas changes in density had a greater influence on mean diversity across all subpopulations. Local subpopulations, however, varied in their regional influence on genetic variation. Decreases in the size and dispersal rates of central populations with low overall and net immigration (i.e. population sources) had the greatest negative impact on genetic variation. Overall, our results provide insight into the interactions among demography, dispersal and genetic variation and highlight the potential of ABC to disentangle the complexity of regional population dynamics and project the genetic impact of changing conditions.     

Spatially explicit models of seasonal habitat for greater sage‐grouse at broad spatial scales: Informing areas for management in Nevada and northeastern California

Defining boundaries of species' habitat across broad spatial scales is often necessary for management decisions, and yet challenging for species that demonstrate differential variation in seasonal habitat use. Spatially explicit indices that incorporate temporal shifts in selection can help overcome such challenges, especially for species of high conservation concern. Greater sage‐grouse Centrocercus urophasianus (hereafter, sage‐grouse), a sagebrush obligate species inhabiting the American West, represents an important case study because sage‐grouse exhibit seasonal habitat patterns, populations are declining in most portions of their range and are central to contemporary national land use policies. Here, we modeled spatiotemporal selection patterns for telemetered sage‐grouse across multiple study sites (1,084 sage‐grouse; 30,690 locations) in the Great Basin. We developed broad‐scale spatially explicit habitat indices that elucidated space use patterns (spring, summer/fall, and winter) and accounted for regional climatic variation using previously published hydrographic boundaries. We then evaluated differences in selection/avoidance of each habitat characteristic between seasons and hydrographic regions. Most notably, sage‐grouse consistently selected areas dominated by sagebrush with few or no conifers but varied in type of sagebrush selected by season and region. Spatiotemporal variation was most apparent based on availability of water resources and herbaceous cover, where sage‐grouse strongly selected upland natural springs in xeric regions but selected larger wet meadows in mesic regions. Additionally, during the breeding period in spring, herbaceous cover was selected strongly in the mesic regions. Lastly, we expanded upon an existing joint–index framework by combining seasonal habitat indices with a probabilistic index of sage‐grouse abundance and space use to produce habitat maps useful for sage‐grouse management. These products can serve as conservation planning tools that help predict expected benefits of restoration activities, while highlighting areas most critical to sustaining sage‐grouse populations. Our joint–index framework can be applied to other species that exhibit seasonal shifts in habitat requirements to help better guide conservation actions.     

Museum samples provide novel insights into the taxonomy and genetic diversity of Irish red grouse

The taxonomic status of red grouse in Ireland has been the subject of considerable debate over the past century. Irish red grouse are usually classified as Lagopus lagopus scoticus, which is the same subspecies as that found in Britain, but some ornithologists believe that native Irish red grouse constitute an endemic subspecies, namely L. lagopus hibernicus. The considerable decline of Irish red grouse over the past century, along with possible hybridization with introduced grouse from Britain, have highlighted the need to resolve their taxonomic status as part of a biodiversity management plan. However, genetic analysis of samples from a single point in time will provide limited insight into potentially confounding historical events such as hybridization and introgression. We therefore compared mtDNA sequences from both current and historical samples of the two putative subspecies, scoticus and hibernicus, to see if they are or were genetically distinct. Red grouse from Britain and Ireland shared mitochondrial haplotypes, and our historical data suggest that this is unlikely to be the result of recent hybridization and introgression. These findings, combined with a general lack of documented differences in behaviour and ecology, suggest that Irish red grouse should remain classified as L. lagopus scoticus. At the same time, we found evidence that a significant amount of genetic diversity has been lost from Irish red grouse over the past century, presumably as a result of diminishing population sizes and fragmentation of extant populations. A loss of habitat, combined with the declining numbers and genetic diversity of Irish red grouse, justify their designation as an All-Ireland Priority (Red List) species and a Northern Ireland Priority Species for conservation.     

Genetic variation within and among fragmented populations of lesser prairie-chickens (Tympanuchus pallidicinctus)

As a result of recurrent droughts and anthropogenic factors, the range of the lesser prairie-chicken (Tympanuchus pallidicinctus) has contracted by 92% and the population has been reduced by approximately 97% in the past century, resulting in the smallest population size and most restricted geographical distribution of any North American grouse. We examined genetic variation through DNA sequence analysis of 478 base pairs of the mitochondrial genome and by assaying allelic variation at five microsatellite loci from lesser prairie-chickens collected on 20 leks in western Oklahoma and east-central New Mexico. Traditional population genetic analyses indicate that lesser prairie-chickens maintain high levels of genetic variation at both nuclear and mitochondrial loci. Although some genetic structuring among lesser prairie-chicken leks was detected within Oklahoma and New Mexico for both nuclear and mitochondrial loci, high levels of differentiation were detected between Oklahoma and New Mexico populations. Nested-clade analysis of mitochondrial haplotypes revealed that both historic and contemporary processes have influenced patterns of haplotype distributions and that historic processes have most likely led to the level of differentiation found between the Oklahoma and New Mexico populations.     

Microsatellite and mitochondrial DNA homogeneity among phenotypically diverse crossbill taxa in the UK.

Genetic differentiation within and between the three morphologically divergent crossbill species extant in the UK was assessed by comparison of allele frequencies at five unlinked microsatellite loci and DNA sequence variation across the mitochondrial control region. No significant differences in microsatellite allele frequency were found either between different populations of the same species or between the crossbill species themselves. A similar lack of genetic divergence was apparent from the mitochondrial sequence data. We resolved 33 different haplotypes, separated by low levels of sequence divergence (0-0.15%). Levels of divergence within and between species were not significantly different. Haplotypes formed a polyphyletic phylogeny, indicating that the crossbill species do not form genetically separate clades. Discordance between neutral DNA polymorphisms and adaptive morphological variation is discussed in relation to defining the systematic relationship between crossbill forms. If adaptive differences have arisen without a concomitant divergence in neutral DNA then attempting to define crossbill types from microsatellite and mitochondrial DNA without recourse to ecology and behaviour may be misleading.     

Behaviourally and genetically distinct populations of an invasive ant provide insight into invasion history and impacts on a tropical ant community

Populations of invasive species often exhibit a high degree of spatial and temporal variability in abundance and hence their effects on resident communities. Here, we examine behavioural, genetic and environmental factors that influence variation in populations of the yellow crazy ant, Anoplolepis gracilipes, on the remote Nukunonu Atoll of Tokelau, Pacific Ocean. Behavioural assays revealed high levels of aggression between two groups of yellow crazy ants from different islands, and genetic analysis confirmed the presence of two distinct populations with unique mitochondrial (mt)DNA haplotypes, designated A and D. The two populations likely resulted from two separate invasion events. The populations exhibited significant differences in abundance of A. gracilipes, with a mean sevenfold difference in relative abundance between the two main haplotypes. The higher density haplotype D population coexisted with 50% fewer other ant species and altered ant community composition. Vegetation composition was also significantly different on islands harbouring the two populations. The results suggest genetic differences could play a role in the spatial and temporal variation in the effect of the yellow crazy ant on a small oceanic atoll. We could not differentiate between genetic effects and effects of vegetation. However, our results indicate that spatial variability in behaviour and impacts within populations of invasive species could be in part due to genetic differences, and play a substantial role in influencing the outcome of biological invasions.     

Restoring Sage‐grouse nesting habitat through removal of early successional conifer

Conifer woodlands have expanded into sagebrush (Artemisia spp.) ecosystems and degrade habitat for sagebrush obligate species such as the Greater Sage‐grouse (Centrocercus urophasianus). Conifer management is increasing despite a lack of empirical evidence assessing outcomes to grouse and their habitat. Although assessments of vegetation recovery after conifer removal are common, comparisons of successional trends with habitat guidelines or actual data on habitat used by sage‐grouse is lacking. We assessed impacts of conifer encroachment on vegetation characteristics known to be important for sage‐grouse nesting. Using a controlled repeated measures design, we then evaluated vegetation changes for 3 years after conifer removal. We compared these results to data from 356 local sage‐grouse nests, rangewide nesting habitat estimates, and published habitat guidelines. We measured negative effects of conifer cover on many characteristics important for sage‐grouse nesting habitat including percent cover of forbs, grasses, and shrubs, and species richness of forbs and shrubs. In untreated habitat, herbaceous vegetation cover was slightly below the cover at local nest sites, while shrub cover and sagebrush cover were well below cover at the nest sites. Following conifer removal, we measured increases in herbaceous vegetation, primarily grasses, and sagebrush height. Our results indicate that conifer abundance can decrease habitat suitability for nesting sage‐grouse. Additionally, conifer removal can improve habitat suitability for nesting sage‐grouse within 3 years, and trajectories indicate that the habitat may continue to improve in the near future.     

Phylogeny and taxonomy of the round-eared sengis or elephant-shrews, genus Macroscelides (Mammalia, Afrotheria, Macroscelidea)

The round-eared sengis or elephant-shrews (genus Macroscelides) exhibit striking pelage variation throughout their ranges. Over ten taxonomic names have been proposed to describe this variation, but currently only two taxa are recognized (M. proboscideus proboscideus and M. p. flavicaudatus). Here, we review the taxonomic history of Macroscelides, and we use data on the geographic distribution, morphology, and mitochondrial DNA sequence to evaluate the current taxonomy. Our data support only two taxa that correspond to the currently recognized subspecies M. p. proboscideus and M. p. flavicaudatus. Mitochondrial haplotypes of these two taxa are reciprocally monophyletic with over 13% uncorrected sequence divergence between them. PCA analysis of 14 morphological characters (mostly cranial) grouped the two taxa into non-overlapping clusters, and body mass alone is a relatively reliable distinguishing character throughout much of Macroscelides range. Although fieldworkers were unable to find sympatric populations, the two taxa were found within 50 km of each other, and genetic analysis showed no evidence of gene flow. Based upon corroborating genetic data, morphological data, near sympatry with no evidence of gene flow, and differences in habitat use, we elevate these two forms to full species.     

The incomplete history of mitochondrial lineages between two rockfishes, Sebastes longispinis and Sebastes hubbsi (Scorpaeniformes: Scorpaenidae)

The genetic divergence between two closely related rockfishes, Sebastes longispinis and Sebastes hubbsi, was inferred from both mitochondrial DNA (mtDNA) sequence variations and amplified fragment length polymorphism (AFLP) markers. The two species were placed into two distinct clades in a neighbour-joining tree based on the AFLP data, clearly indicating that they represented separate species. Although this evidence, together with a previous morphological study, revealed clear differences between the two species, no obvious clustering of haplotypes by species was detected in the minimum spanning network inferred from sequence variations in the mtDNA control region (c. 500 base pairs). In fact, the significant Φ(ST) estimates indicated only a restriction of gene flow between the two species. Uncorrected pairwise sequence differences in mtDNA between two species were small (1·8% at maximum, on the lower end of the range of control region divergence between previously studied sister species pairs), suggesting their speciation event as having been fairly recent. The incongruent results of AFLP and mtDNA phylogenies suggested incomplete lineage sorting and introgression of mtDNA in the course of the evolution of the two species. Differences in their main distributional ranges and the small level of sequence divergence in mtDNA suggests that speciation and dispersal may have been associated with glacio-eustatic sea level fluctuations between the Japanese Archipelago and the Korean Peninsula during the past 0·4 million years.     

Matrilineal genetic structure and female-mediated gene flow in red grouse (Lagopus lagopus scoticus): an analysis using mitochondrial DNA

Abstract.— DNA sequence variation at the hypervariable 5'end of the mitochondrial control region was examined in 247 individuals to detect genetic divergence among 14 populations of red grouse ( Lagopus lagopus scoticus ) in northeastern Scotland. Ten haplotypes were resolved, several of which were shared among populations. Analysis of molecular variance, Nei's yST, and a cladistic estimate of the amount of gene flow indicated a lack of overall population differentiation. Patterns of overall panmixia are in stark contrast to previous reports of localized subdivision among the same set of populations detected using hypervariable microsatellite markers. Because grouse cocks are territorial and show extreme natal philopatry and females are the dispersing sex, such discordance could be explained by sexbiased dispersal, with extensive female-mediated gene flow preventing mitochondrial DNA divergence. However, it is difficult to reconcile how effective dispersal of females would not homogenize both mitochondrial and nuclear structure simultaneously. We use a model that examines the spatial and temporal dynamics of diparentally and uniparentally inherited genes to show that, under realistic ecological scenarios and with specific differences in the dispersal of males and females, the local effective size of the nuclear genome can be less than that of the mitochondrial and the patterns of structuring we observe are meaningful.     

Genetic variation in Holocene bowhead whales from Svalbard

Bowhead whales (Balaena mysticetus) are distributed in the Arctic in five putative stocks. All stocks have been heavily depleted due to centuries of exploitation. In the present study, nucleotide sequence variation of the mitochondrial control region was determined from bone remains of 99 bowhead whales. The bones, 14C dated from recent to more than 50,000 bp, were collected on Svalbard (Spitsbergen) and are expected to relate to ancestors of the today nearly extinct Spitsbergen stock. Fifty-eight haplotypes were found, a few being frequent but many only found in one individual. The most abundant haplotypes of the Spitsbergen stock are the same as those most abundant in the extant Bering-Chukchi-Beaufort (BCB) Seas stock of bowhead whales. Although F(ST) indicates a slight but statistically significant genetic differentiation between the Spitsbergen and the BCB stocks this was not considered informative due to the very high levels of genetic diversity of mitochondrial DNA haplotypes in both bowhead whale stocks. Other measures such as K(ST) also indicated very low genetic differentiation between the two populations. Nucleotide diversity and haplotype diversity showed only minor differences between the Spitsbergen and BCB stocks. The data suggest that the historic Spitsbergen stock--before the severe bottleneck caused by whaling--did not have substantially more genetic variation than the extant BCB stock. The similar haplotypes of the Holocene Svalbard samples and the current BCB stock indicate significant migration between these two stocks and question the current designation of five distinct stocks of bowhead whales in the Arctic.     

Population genetic structure of two ecologically distinct Amazonian spiny rats: separating history and current ecology

Abstract. Population history and current demographic and ecological factors determine the amount of genetic variation within and the degree of differentiation among populations. Differences in the life history and ecology of codistributed species may lead to differences in hierarchical population genetic structure. Here, we compare patterns of genetic diversity and structure of two species of spiny rats in the genus Proechimys from the Rio Jurua of western Amazonian Brazil. Based on the ecological and life-history differences between the two species, we make predictions as to how they might differ in patterns of genetic diversity and structure. We use mitochondrial sequence data from the cytochrome b gene to test these predictions. Although both species maintain nearly the same number of mitochondrial haplotypes across the sampled range, they differ in levels of genetic diversity and geographic structure. Patterns of gene flow are also different between the two species with average M-values of nearly three in P. steerei and less than one in P. simonsi . Our initial predictions are largely upheld by the genetic data and where conflicting hypotheses arise, we suggest further studies that may allow us to distinguish among evolutionary scenarios. Separating the effects of history and ongoing demography on patterns of genetic diversity is challenging. Combining genetic analyses with field studies remains essential to disentangling these complex processes.     

Mitochondrial DNA sequence variation among geographical isolates of Opisthorchis viverrini in Thailand and Lao PDR, and phylogenetic relationships with other trematodes

The present study compared the genetic variation among 14 different geographical isolates of Opisthorchis viverrini sensu lato from Thailand and Lao PDR using sequence data for 2 mitochondrial DNA genes, the subunit 1 of NADH dehydrogenase gene (nad1) and cytochrome c oxidase gene (cox1). Four different nad1 haplotypes were detected among isolates, all of which were identical at the amino acid sequence level. Nucleotide sequence variation among 14 isolates ranged from 0 to 0.3% for nad1. Two different cox1 haplotypes were detected among isolates. These two haplotypes differed at 2 nucleotide positions, one of which resulted in a change in the amino acid sequence. Nucleotide sequence variation among isolates for cox1 ranged from 0 to 0.5%. Comparison of cox1 sequences of O. viverrini to those of other trematodes revealed nucleotide differences of 13-31%. A phylogenetic analysis of the cox1 sequence data revealed strong statistical support for a clade containing O. viverrini and 2 other species of opisthorchid trematodes; O. felineus and Clonorchis sinsensis.     

Relationships of eelpouts of genus Zoarces (Zoarcidae, Pisces) inferred from molecular genetic and morphological data

Molecular genetic and morphological analysis of eelpouts of the genus Zoarces was carried out. Based on the mitochondrial DNA sequence variation, haplotypes of notched-fin eelpout, Z. elogatus, more closely related Fedorov eelpout, Z. fedorovi, and common eelpout, Z. viviparus, as well as of Andriyashev eelpout, Z. andriashevi, were grouped in one macrocluster. Haplotypes of American eelpout, Z. americanus, and blotched eelpout, Z. gillii, clustered separately from other species. The genetic differences between Z. gillii and the other eelpout species were very high for within-genus comparisons, constituting 7.62%. Species divergence in terms of morphological characters was generally consistent with molecular genetic data and confirmed distinct isolation of American eelpout, and especially of blotched eelpout.     

Population structure and comparative phylogeography of jack species (Caranx ignobilis and C. melampygus) in the high Hawaiian Islands

Members of the family Carangidae are top-level predators and highly prized food and sport fishes. Although ecologically and economically important, little is known about the biology of numerous species in the family. This is particularly true of the jacks Caranx ignobilis and C. melampygus, which have experienced recent population reductions around the high Hawaiian Islands due to overfishing. Previous studies have documented territorial tendencies as well as cases of long-distance excursions in both species, suggesting populations may exhibit a range of structure at the genetic level. To explore this possibility, mitochondrial DNA ATPase6 and ATPase8 gene sequence variation was assessed from 91 individuals (33 C. ignobilis and 58 C. melampygus) spanning the islands of Kaua'i, O'ahu, Moloka'i, Maui, and Hawai'i. Although a total of 20 distinct haplotypes (8 for C. ignobilis; 12 for C. melampygus) were recovered, no evidence of population structure was found for either species across the examined geographic range. However, distinct demographic patterns were identified, implying differing evolutionary histories and/or population dynamics. Additionally, ～ 6% of the examined C. ignobilis were C. ignobilis × C. melampygus hybrids because they harbored mitochondrial haplotypes typical of C. melampygus. These hybrids contribute to measurable gene flow between the species and may play a significant role in the evolution of the genus.     

Ecological and biogeographical inferences on two sympatric and enigmatic Andean cat species using genetic identification of faecal samples

The carnivore community of the altiplano ecosystem of the high Andes, including the Andean mountain cat (Leopardus jacobita) and pampas cat (Leopardus colocolo), is one of the least studied in the world. We determined the origin of 186 carnivore samples (184 faeces and two skulls) collected above 3000 m above sea level in northern Chile, including 33 from the Andean mountain cat and 75 from the pampas cat using diagnostic molecular genetic sequence variation. We determined for the first time food habits, habitat and physiographic associations, and general patterns of molecular genetic variation of the Andean mountain cat and the pampas cat in Chile. Both species had narrow dietary niches dominated by small rodents and there was a wide overlap in diet composition (0.82), suggesting low levels of prey partitioning between species. The mountain viscacha (Lagidium viscacia) made up a large proportion of the biomass of the diet of both species, especially for the Andean mountain cat (93.9% vs. 74.8% for the pampas cat), underscoring the importance of further research and conservation focus on this vanishing prey species. Although the probability of finding Andean mountain cat scats increased with altitude and slope, there was substantial geographical overlap in distribution between species, revealing that the pampas cat distribution includes high-altitude grassland habitats. The Andean mountain cat had relatively low levels of mitochondrial DNA (mtDNA) genetic variation (two mtDNA haplotypes) compared with the pampas cat (17 mtDNA haplotypes), suggestive of a distinct evolutionary history and relatively smaller historic populations. These insights will facilitate and provide tools and hypotheses for much-needed research and conservation efforts on these species and this ecosystem.     

Molecular phylogenetic status of the Korean goral and Japanese serow based on partial sequences of the mitochondrial cytochrome b gene

To investigate the molecular phylogenetic status of the Korean goral, Nemorhaedus caudatus raddeanus, and Japanese serow, Capricornis crispus, we determined partial sequences of the mitochondrial cytochrome b gene of twelve Korean gorals and sixteen Japanese serows, and compared them with those of the major lineages of Rupicaprini species including two other Nemorhaedus species and two other Capricornis species. The Korean gorals examined possessed two haplotypes with only one nucleotide difference between them, while the Japanese serows showed slightly higher sequence diversity with five haplotypes. Genetic distances and molecular phylogenetic trees indicated that there is considerable genetic divergence between the Korean goral and N. caudatus (the Chinese goral) [Groves and Shields (1996)], but virtually none between Korean and Russian gorals. The Korean and Russian gorals may therefore be distinct from the Chinese goral. The data highlight the importance of conservation of the goral populations of these regions, and the need to reconsider the taxonomic status of Korean and Russian gorals. Our study also clearly demonstrated sufficient genetic distance between serows and gorals to justify their assignment to separate genera. Of the three species of Capricornis, the Formosan serow, C. swinhoei is more closely related to C. sumatraensis than to the Japanese serow, suggesting that the Formosan serow is a distinct species. Preliminary data on intraspecific genetic variation in the Japanese serow are also presented.     

Phylogeographic analysis of mitochondrial gene flow and introgression in the salamander, Plethodon shermani

Plethodon shermani comprises a series of geographically disjunct populations occupying high-elevation mountain isolates. These populations hybridize at their borders with salamanders of the Plethodon glutinosus species complex, and past range expansions inferred from Pleistocene climatic cycles may have increased the possible genetic interactions between P. shermani and species of the P. glutinosus complex. Because mitochondrial DNA haplotypes often show introgression across species borders, we survey mtDNA variation for evidence of past and ongoing genetic interactions between P. shermani, its close relative Plethodon cheoah, and species of the P. glutinosus complex. Ongoing hybridization with the P. glutinosus-complex species Plethodon teyahalee is accompanied by extensive mitochondrial introgression in some Unicoi populations of P. shermani, but it has little genetic impact on P. shermani populations outside hybrid zones at three other isolates (Tusquitee, Wayah Bald, Standing Indian). Some Unicoi populations of P. shermani exhibit mtDNA evidence of past hybridization with diverse lineages from P. aureolus and P. glutinosus. The Tusquitee isolate of P. shermani is also characterized by mtDNA haplotypes most closely related to Plethodon aureolus and P. glutinosus, presumably introduced by past genetic contact with these species or with introgressed populations of Unicoi P. shermani. The mtDNA variation in sampled populations of the Wayah Bald and Standing Indian isolates of P. shermani appears largely unaffected by ongoing hybridization. Principal components analyses of allozymic data indicate that P. shermani isolates collectively form a genetically homogeneous unit clearly demarcated from species with which they have had current or past genetic interactions. Rapid mtDNA introgression associated with transient contacts between P. shermani and other species permits a fine-level resolution of evolutionary lineages not evident from allozymic data.     

Macrophyte development in unimpacted lowland rivers in Poland

Three ecologically and morphologically distinct forms of Arctic charr (Salvelinus alpinus L.) have been identified in Loch Rannoch, Scotland, whose evolutionary status and origins are incompletely understood. A study was made of restriction fragment length polymorphism (RFLPs) detected variation in the D-loop, ND1 and cytochrome b regions of the mitochondrial genome, encompassing >3500 bp. Eight RFLP haplotypes were identified that clustered into three distinct clans based on restriction differences and into four clans based on sequence differences. Significant differences in RFLP frequencies were found among all morph groups. The pelagic morph was highly divergent from the two benthic forms, with the benthic forms having variants from only one genetic clan while the pelagic was dominated by a single variant from another clan. The relative divergence observed among benthic and pelagic forms is ~10 fold greater when nucleotide divergence among the haplotypes, as well as haplotype frequency differences, is taken into account. Sequence divergence between haplotypes in the two main clans is of a similar order to that between haplotypes in these clans and a charr from North America. In contrast, divergence among the two benthic morphs relates entirely to differences in haplotype frequencies. The study confirms the genetic distinctiveness of the pelagic and benthic forms as well as of the two benthic forms. It strongly supports previous evidence that the genetic divergence between the pelagic and benthic populations is allopatric in origin. Additionally, the results strongly suggest that the two benthic populations have undergone peripatric divergence through the sequential colonisation of the two basins by one lineage, followed by their spatial separation and reproductive isolation.