Phylogeography of Rhinichthys cataractae (Teleostei: Cyprinidae): pre‐glacial colonization across the Continental Divide and Pleistocene diversification within the Rio Grande drainage

The longnose dace, Rhinichthys cataractae, is a primary freshwater fish inhabiting riffle habitats in small headwater rivers and streams across the North American continent, including drainages east and west of the Continental Divide. The mitochondrially encoded cytochrome b gene (1140 bp) and 2298–2346 bp of the nuclear‐encoded genes S7 and RAG1 were obtained from 87 individuals of R. cataractae (collected from 17 sites throughout its range) and from several close relatives. Phylogenetic analyses recovered a monophyletic R. cataractae species‐group that contained Rhinichthys evermanni, Rhinichthys sp. ‘Millicoma dace’, and a non‐exclusive R. cataractae. Within the R. cataractae species‐group, two well‐supported lineages were identified, including a western lineage (containing R. evermanni, R. sp. ‘Millicoma dace’ and individuals of R. cataractae from Pacific slope drainages) and an eastern lineage (containing individuals of R. cataractae from Arctic, Atlantic, and Gulf slope drainages). Within the eastern lineage of R. cataractae, two well‐supported groups were recovered: a south‐eastern group, containing individuals from the Atlantic slope, southern tributaries to the Mississippi River, and the Rio Grande drainage; and a north‐eastern group, containing individuals from the Arctic slope and northern tributaries to the Mississippi River. Estimates of the timing of divergence within the R. cataractae species‐group, combined with ancestral area‐reconstruction methods, indicate a separation between the eastern and western lineages during the Pliocene to early‐Pleistocene, with a direction of colonization from the west of the Continental Divide eastward. Within the southern portion of its range, R. cataractae likely entered the Rio Grande drainage during the Pleistocene via stream capture events between the Arkansas River (Mississippi River drainage) and headwaters of the Rio Grande. A close relationship between populations of R. cataractae in the Rio Grande drainage and the adjacent Canadian River (Mississippi River drainage) is consistent with hypothesized stream capture events between the Pecos (Rio Grande drainage) and Canadian rivers during the late‐Pleistocene. The population of R. cataractae in the lower Rio Grande may have become separated from other populations in the Rio Grande drainage (upper Rio Grande and Pecos River) and Canadian River during the late‐Pleistocene, well before initiation of recent and significant anthropogenic disturbance within the Rio Grande drainage. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 317–333.     

Species delimitation in a range-restricted group of cascudinhos (Loricariidae: Epactionotus) supports morphological and genetic differentiation across coastal rivers of southern Brazil

Epactionotus species are known for inhabiting the rocky-bottom stretches of fast-flowing rivers in a limited geographic area along the Atlantic coast of southern Brazil. These species are endemic to single coastal river drainages (two neighbouring drainages for Epactionotus bilineatus) isolated from each other by the coastal lacustrine environments or the Atlantic Ocean. E. bilineatus is from the Maquiné and Três Forquilhas River basins, both tributaries of the Tramandaí River system, whereas E. itaimbezinho is endemic to the Mampituba River drainage and Epactionotus gracilis to the Araranguá River drainage. Recent fieldwork in the Atlantic coastal drainages of southern Brazil revealed new populations in the Urussanga, Tubarão, d'Una and Biguaçu River drainages. Iterative species delimitation using molecular data (cytochrome c oxidase subunit I) and morphology (morphometrics and meristics) was applied to evaluate species recognition of isolated populations. With regard to new data, the genus was re-diagnosed, the status of Epactionotus species/populations was re-evaluated, formerly described species were supported and population structure was recognized. As for the newly discovered populations, both morphological and molecular data strongly support the population from the Biguaçu River drainage, in Santa Catarina State, as a new species. Molecular data revealed strong per-basin population structure, which may be related to species habitat specificity and low or no dispersal among drainages.     

Intraspecific life history variation in the threatened Louisiana pearlshell mussel, Margaritifera hembeli

1. Margaritifera hembeli is a threatened mussel limited to twenty-two headwater streams in the Red River drainage in central Louisiana, USA. This study evaluated intraspecific variation in density, growth, size and age structure and shell morphology among several isolated populations. This study also identified the host fish and considered the role that host fish distribution played in determining mussel recruitment. 2. Mussels were aggregated in beds and average densities differed among streams. However, maximum mussel densities in beds were similar in all streams; the observed maxima were among the largest for monospecific mussel beds in North America, often exceeding 300 individuals m^–2. 3. The maximum size reached by individuals differed among streams, but all size distributions were skewed towards larger individuals. A repeated measures analysis of tagged mussels in four populations, over a 2-year period, indicated 2-fold differences in growth rates among streams, and significant variation among years. Growth rates were not affected by local population density. Maximum ages reached, determined indirectly by comparing growth rates, varied from 45 to 75 years. A canonical discriminant analysis also revealed significant differences in shell morphology across populations. 4. Half of the populations showed evidence of recent recruitment, and these sites had fish assemblages dominated by the host fish Noturus phaeus (Taylor). Host fish abundance appeared more important than adult mussel density in explaining recruitment patterns. 5. Considerable intraspecific life history variation suggests that management strategies for this species should be stream-specific, with emphasis on ensuring long-term habitat stability.     

The effects of age,sex, and habitat on body size and shape of the blackstripe topminnow,Fundulus notatus (Cyprinodontiformes: Fundulidae) (Rafinesque 1820)

Lake and stream habitats pose a variety of challenges to fishes due to differences in variables such as water velocity, habitat structure, prey community, and predator community. These differences can cause divergent selection on body size and/or shape. Here, we measured sex, age, length, and eight different morphological traits of the blackstripe topminnow, Fundulus notatus, from 19 lake and stream populations across four river drainages in central Illinois. Our goal was to determine whether size and shape differed consistently between lake and stream habitats across drainages. We also considered the effects of age and sex as they may affect size and morphology. We found large differences in body size of age 1 topminnows where stream fish were generally larger than lake fish. Body shape mainly varied as a function of sex. Adult male topminnows had larger morphological traits (with the exception of body width) than females, in particular longer dorsal and anal base lengths. Subtle effects of habitat were present. Stream fish had a longer dorsal fin base than lake fish. These phenotypic patterns may be the result of genetic and/or environmental variation. As these lakes are human‐made, the observed differences, if genetic, would have had to occur relatively rapidly (within about 100 years). © 2013 The Linnean Society of London     

Microsatellite DNA Population Structure and Stock Identification of Steelhead Trout (<Emphasis Type="BoldItalic">Oncorhynchus mykiss</Emphasis>) in the Nass and Skeena Rivers in Northern British Columbia

Population structure and the application to genetic stock identification for steelhead (Oncorhynchus mykiss) in the Nass and Skeena Rivers in northern British Columbia was examined using microsatellite markers. Variation at 8 microsatellite loci (Oki200, Omy77, Ots1, Ots3, Ssa85, Ots100, Ots103, and Ots108) was surveyed for approximately 930 steelhead from 7 populations in the Skeena River drainage and 850 steelhead from 10 populations in the Nass River drainage, as well as 1550 steelhead from test fisheries near the mouth of each river. Differentiation among populations within rivers accounted for about 1.9 times the variation observed among years within populations, with differences between drainages less than variation among populations within drainages. In the Nass River, winter-run populations formed a distinct group from the summer-run populations. Winter-run populations were not assessed in the Skeena River watershed. Simulated mixed-stock samples suggested that variation at the 8 microsatellite loci surveyed should provide relatively accurate and precise estimates of stock composition for fishery management applications within drainages. In the Skeena River drainage in 1998, Babine River (27%) and Bulkley drainage populations (31%) comprised the main components of the returns. For the Nass River in 1998 steelhead returning to Bell-Irving River were estimated to have comprised 39% of the fish sampled in the test fishery, with another 27% of the returns estimated to be derived from Cranberry River. The survey of microsatellite variation did not reveal enough differentiation between Nass River and Skeena River populations to be applied confidently in estimation of stock composition in marine fisheries at this time. Received January 14, 2000; accepted July 13, 2000.     

Morphological diversity among fishes in a Great Plains river drainage

Longitudinal succession of stream fish faunas is commonly related to increasing stream size and stability. However, effects of succession on assemblage morphology are seldom quantified. We used an ecomorphological approach to determine differences in faunal structure among distinct stream types of the Cheyenne River drainage in South Dakota, USA. During May–October 2004 we collected fishes monthly from five streams. We examined 28 morphological traits of the dominant fish species and compared morphological structure among faunas using univariate analysis of variance (ANOVA) tests and multivariate ordination and distance calculation techniques. Species richness and composition varied between smaller creeks and larger rivers. Morphological diversity increased with richness, but richer assemblages were also more tightly packed in morphospace, partly because of increased cyprinid richness. Some morphological differences were predicted by variation in mean discharge and discharge flashiness (flow stability). Fishes of more stable or larger river stations characteristically had smaller heads and mouths and longer intestines. Larger mean standard length was also associated with less flashy flow regimes and higher mean discharge. All assemblages were hyperdispersed in morphological space, consistent with the harsh zoogeographical history of the region and suggesting the presence of open niches. Increasing species and morphological diversity despite increasing discharge flashiness suggests higher niche diversity in Great Plains rivers compared to adjacent creeks. Handling editor: J. A. Cambray     

Population genetics of the freshwater mussel, <Emphasis Type="Italic">Amblema plicata</Emphasis> (Say 1817) (Bivalvia: Unionidae): Evidence of high dispersal and post-glacial colonization

Over 70% of North American freshwater mussel species (families Unionidae and Margaritiferidae) are listed as threatened or endangered. Knowledge of the genetic structure of target species is essential for the development of effective conservation plans. Because Ambelma plicata is a common species, its population genetic structure is likely to be relatively intact, making it a logical model species for investigations of freshwater mussel population genetics. Using mtDNA and allozymes, we determined the genotypes of 170+ individuals in each of three distinct drainages: Lake Erie, Ohio River, and the Lower Mississippi River. Overall, within-population variation increased significantly from north to south, with unique haplotypes and allele frequencies in the Kiamichi River (Lower Mississippi River drainage). Genetic diversity was relatively low in the Strawberry River (Lower Mississippi River drainage), and in the Lake Erie drainage. We calculated significant among-population structure using both molecular markers (A.p. Φ_st = 0.15, θ_st = 0.12). Using a hierarchical approach, we found low genetic structure among rivers and drainages separated by large geographic distances, indicating high effective population size and/or highly vagile fish hosts for this species. Genetic structure in the Lake Erie drainage was similar to that in the Ohio River, and indicates that northern populations were founded from at least two glacial refugia following the Pleistocene. Conservation of genetic diversity in Amblema plicata and other mussel species with similar genetic structure should focus on protection of a number of individual populations, especially those in southern rivers.     

Phylogeography and post-glacial colonization patterns of the rainbow darter, Etheostoma caeruleum (Teleostei: Percidae)

Aim To examine the effects of historical climate change and drainage isolation on the distribution of mitochondrial DNA cytochrome b genetic variation within the rainbow darter, Etheostoma caeruleum (Percidae: Etheostomatinae). Location Eastern North American streams including tributaries to the Mississippi River, Great Lakes, Potomac River and Hudson Bay drainages. Methods Parsimony analyses, Bayesian analyses and haplotype networks of mitochondrial DNA sequences. Results Four major clades were recovered from sampled populations of E. caeruleum. Three of four clades are distributed in the western portion of the species’ range (primarily west of the Mississippi River). Samples from this region do not form a monophyletic group, and sequences often vary greatly between samples from adjacent stream systems (up to 7.2% divergence). A basal clade includes samples from the White River system in the Ozark Highlands. The northern Ozarks–upper Midwest clade includes samples from Missouri River tributaries and the upper Midwest (Hudson Bay, upper Mississippi River, and western Lake Michigan drainage). The eastern clade is composed of individuals from the Ohio River, Great Lakes and Potomac River. The Mississippi River corridor clade includes samples from middle and lower Mississippi River tributaries. Main conclusions The four major clades of E. caeruleum are deep allopatric lineages with well‐defined boundaries and have additional phylogeographical structure within each clade. The Ozark Highlands have the greatest levels of diversity relative to distributional area, with marked cytochrome b subdivisions between adjacent stream systems. Samples from previously glaciated areas do not have a subset of the cytochrome b diversity found in unglaciated areas, but four separate source areas are identified based on phylogenetic analyses. Dispersal into previously glaciated areas followed several known glacial outlets and, based on sequence divergence between populations, may have occurred during different glacial or interglacial stages. The disjunct distribution and cytochrome b pattern of E. caeruleum in the Mississippi River corridor clade is consistent with late Pleistocene and Recent changes in the course and characteristics of the middle and lower Mississippi River. Phylogeographical boundaries between clades of E. caeruleum correspond to independent sources of biogeographical information and provide insight into historical stream drainage relationships, post‐glacial colonization and drainage isolation patterns.     

Genetic effects of habitat fragmentation and population isolation on Etheostoma raneyi (Percidae)

The use of genetic methods to quantify the effects of anthropogenic habitat fragmentation on population structure has become increasingly common. However, in today’s highly fragmented habitats, researchers have sometimes concluded that populations are currently genetically isolated due to habitat fragmentation without testing the possibility that populations were genetically isolated before European settlement. Etheostoma raneyi is a benthic headwater fish restricted to river drainages in northern Mississippi, USA, that has a suite of adaptive traits that correlate with poor dispersal ability. Aquatic habitat within this area has been extensively modified, primarily by flood-control projects, and populations in headwater streams have possibly become genetically isolated from one another. We used microsatellite markers to quantify genetic structure as well as contemporary and historical gene flow across the range of the species. Results indicated that genetically distinct populations exist in each headwater stream analyzed, current gene flow rates are lower than historical rates, most genetic variation is partitioned among populations, and populations in the Yocona River drainage show lower levels of genetic diversity than populations in the Tallahatchie River drainage and other Etheostoma species. All populations have negative F_IS scores, of which roughly half are significant relative to Hardy–Weinberg expectations, perhaps due to small population sizes. We conclude that anthropogenic habitat alteration and fragmentation has had a profoundly negative impact on the species by isolating E. raneyi within headwater stream reaches. Further research is needed to inform conservation strategies, but populations in the Yocona River drainage are in dire need of management action. Carefully planned human-mediated dispersal and habitat restoration should be explored as management options across the range of the species.     

Adaptive divergence between lake and stream populations of an East African cichlid fish

Divergent natural selection acting in different habitats may build up barriers to gene flow and initiate speciation. This speciation continuum can range from weak or no divergence to strong genetic differentiation between populations. Here, we focus on the early phases of adaptive divergence in the East African cichlid fish Astatotilapia burtoni, which occurs in both Lake Tanganyika (LT) and inflowing rivers. We first assessed the population structure and morphological differences in A. burtoni from southern LT. We then focused on four lake–stream systems and quantified body shape, ecologically relevant traits (gill raker and lower pharyngeal jaw) as well as stomach contents. Our study revealed the presence of several divergent lake–stream populations that rest at different stages of the speciation continuum, but show the same morphological and ecological trajectories along the lake–stream gradient. Lake fish have higher bodies, a more superior mouth position, longer gill rakers and more slender pharyngeal jaws, and they show a plant/algae and zooplankton‐biased diet, whereas stream fish feed more on snails, insects and plant seeds. A test for reproductive isolation between closely related lake and stream populations did not detect population‐assortative mating. Analyses of F1 offspring reared under common garden conditions indicate that the detected differences in body shape and gill raker length do not constitute pure plastic responses to different environmental conditions, but also have a genetic basis. Taken together, the A. burtoni lake–stream system constitutes a new model to study the factors that enhance and constrain progress towards speciation in cichlid fishes.     

An analysis of spatial and environmental factors influencing hybridization between native westslope cutthroat trout (Oncorhynchus clarki lewisi) and introduced rainbow trout (O. mykiss) in the upper Kootenay River drainage, British Columbia

Westslope cutthroat trout (Oncorhynchus clarki lewisi, WCT) and introduced rainbow trout (O. mykiss, RBT) readily hybridize and introgression has occurred in many drainages across the historic native range of WCT. In British Columbia (Canada), the upper Kootenay River drainage is the heart of the westslope cutthroat trout (WCT, Oncorhynchus clarki lewisi) distribution in Canada this drainage harbours native WCT gene pools that are thought to be under threat from hybridization with introduced rainbow trout (RBT, O. mykiss). In this study, we assess the extent and distribution of WCT × RBT hybridization in the upper Kootenay River drainage. We used four diagnostic nuclear loci to determine the extent of hybridization in 981 fish collected from 23 sample localities across 12 different streams in the upper Kootenay River drainage. About 14% (142/981) of individuals were identified as hybrids (an individual with both RBT and WCT alleles), 3.4% (33/981) were identified as pure RBT, and the remaining individuals were identified as pure WCT. Although pure RBT were absent from the majority of locales (20/23), we found evidence of hybridization at 78% (18/23) of the localities and the percentage of heterospecific alleles (% I) ranged from 0.7% to 97.1%. Only 22% (5/23) of the localities showed no evidence of hybridization. Spatial analysis showed clustering among hybridized locations and decreasing hybridization with increasing distance from Koocanusa Reservoir, suggesting that the reservoir acts as a RBT source. We found no evidence that stream order, stream magnitude, or stream elevation influenced the extent of hybridization among localities. We compared our results to an analysis conducted in 1986, which indicated that hybridization is relatively recent in the upper Kootenay River drainage and that it is increasing in magnitude and distribution. In the absence of timely management intervention, the genetic integrity of WCT populations in the heart of their Canadian range may be lost. Our results indicate the dynamic nature of hybridization in fluvial systems and that for closely related taxa such as WCT and RBT, hybridization appears to be largely influenced by physical barriers to dispersal and contact between species.     

Phylogeography of the bigeye chub Hybopsis amblops (Teleostei: Cypriniformes): early Pleistocene diversification and post‐glacial range expansion

The bigeye chub, Hybopsis amblops, is a member of the Central Highlands ichthyofauna of eastern North America. Phylogenetic analyses of the H. amblops species group based on a 1059 bp fragment of the mitochondrial DNA cytochrome b gene did not recover a monophyletic group. The inclusion of Hybopsis hypsinotus in the species complex is questionable. Within H. amblops, five strongly supported clades were identified; two clades containing haplotypes from the Ozark Highlands and three clades containing haplotypes from the Eastern Highlands and previously glaciated regions of the Ohio and Wabash River drainages. Estimates of the timing of divergence indicated that prior to the onset of glaciation, vicariant events separated populations east and west of the Mississippi River. East of the Mississippi River glacial cycles associated with the blocking and rerouting of the Teays River system caused populations to be pushed southward into refugia of the upper Ohio River. Following the most recent Wisconsinan glaciation, populations expanded northward into previously glaciated regions and southward into the Cumberland River drainage. In the Ozarks, west of the Mississippi River, isolation of clades appears to be maintained by the lack of stream capture events between the upper Arkansas and the White River systems and a barrier formed by the Arkansas River.     

Creation and maintenance of habitat downstream from a weir for the greenside darter, Etheostoma blennioides – a rare fish in Canada

The biology, microhabitat use and migratory behaviour of the greenside darter, Etheostoma blennioides, was studied at the Mannheim Weir on the Grand River, Ontario during the summer of 1995 and 1996. Officially listed as vulnerable in Canada, greenside darters reached maturity at age 1 and lived up to 4 years. They were found in riffle habitats that consisted of cobble and loose boulders, with large mats of Cladophora. This type of unembedded substrate is uncommon in the Grand River watershed. However, local abundance of greenside darters immediately downstream from the Mannheim Weir was likely due to high water velocities from weir discharge, freshets and ice scour which help maintain unembedded riffle areas. Trap data indicated that greenside darters temporally partition this habitat with the stonecat, Noturus flavus. Other darter species were not commonly found in areas with greenside darters, whose depth selection and habitat choices were influenced by predators and morphology. Denil fishways at the Mannheim Weir rarely passed greenside darters due to prohibitively high water velocities and exclusion by larger fish. This revised version was published online in August 2006 with corrections to the Cover Date.     

Patterns of population differentiation in annual killifishes from the Paraná–Uruguay–La Plata Basin: the role of vicariance and dispersal

Aim To elucidate the role of vicariance versus dispersal at the microevolutionary scale in annual killifish populations belonging to the Austrolebias bellottii species complex (Rivulidae). Within this complex, A. bellottii and A. apaii have low vagility and occur widely within the study area, making them excellent models for testing biogeographic hypotheses of differentiation. Location South America, in the Paraná–Uruguay–La Plata river basin. Methods Molecular data and morphometric analyses were used to reconstruct the phylogeographic history and morphological variation of 24 populations of two taxa of the A. bellottii species complex. Phylogenetic analyses using maximum likelihood (ML) and Bayesian inference (BI) model‐based methods, estimates of clade divergence times implemented in beast , non‐metric multidimensional scaling, analysis of molecular variance results, and morphological analyses elucidated the role of vicariance versus dispersal hypotheses in population differentiation in the aforementioned river basin. Results In the A. bellottii species complex from the Paraná–Uruguay–La Plata river basin, past allopatric fragmentation from vicariance events seems to be the most plausible scenario for diversification since the Late Miocene and more recently since the Plio‐Pleistocene. The Plio‐Pleistocene vicariance produced the differentiation of three major clades in A. bellottii populations. One clade from the eastern Uruguay River drainage was separated from another in western Uruguay and the Paraná–La Plata River drainages. A later vicariance event split populations to the south (lower Paraná–La Plata Basin) and north (middle Paraná) of the western Paraná River drainage. However, our results do not exclude the possibility of dispersal events among A. bellottii populations from both the Uruguay and Paraná river drainages, which could occur in these river basins during hypothesized connectivity cycles of the Late Pliocene and Pleistocene. Main conclusions Past allopatric fragmentation caused by different vicariance events seems to be the main driver of diversification in the A. bellottii species complex since the Plio‐Pleistocene. However, the current molecular data suggest that populations from both drainages of the Paraná–Uruguay rivers may have experienced cycles of connectivity during the Pleistocene, perhaps including multiple vicariance or dispersal events from populations located in the western lower Uruguay River drainage, which encompassed climatic and geological changes in the Paraná–Uruguay–La Plata Basin.     

海南岛宽额鳢(Channa gachua)群体遗传变异与生物地理过程

为了解海南岛宽额鳢(Channa gachua)的群体遗传分化和亲缘生物地理过程,采集了云南元江和海南岛5个水系(昌化江、陵水河、藤桥河、万泉河及南渡江)共6个种群168个宽额鳢个体,基于线粒体细胞色素b(Cyt b)基因全序列(1142 bp)对其遗传多样性和遗传分化程度进行了评估,并探讨了地质和气候等因素如何塑造了这一物种的亲缘地理结构及演化历史。基于Cyt b序列构建的系统树结果将所有个体分成两个主要谱系(A和B),谱系A包括海南岛所有种群,其中,部分昌化江个体形成独立的亚支(A2),其余个体聚为另一亚支(A1),谱系B为云南元江的全部个体,各谱系间的遗传分化指数均较高。种群历史动态分析表明,各谱系均没有发生种群扩张,但A1亚支与谱系B曾在约1万年前发生过有效种群数量减小的事件。根据研究结果推测,更新世冰期期间,北部湾因海平面下降而暴露,大陆和海南岛的水系发生接触,越南北部水系(包括元江/红河)通过一条联系雷州半岛和海南岛的古河道流入南海,因而冰期期间宽额鳢有机会从元江(红河)扩散至海南岛西南部,随后在海南岛内部,宽额鳢进一步扩散,并以五指山为种群间基因交流的重要地理障碍,各水系间种群发生基因交流和遗传分化。     

中间黄颡鱼群体遗传变异与亲缘生物地理

通过分析81尾采自华南西部12条水系的中间黄颡鱼(Pelteobagrus intermedius)mtDNA控制区435 bp的序列,研究其群体遗传变异及亲缘生物地理格局。结果显示,中间黄颡鱼群体间的遗传分化较小,核苷酸变异只有0.54%。12条水系的群体共有7个单倍型,其中一些现已相互隔离的水系中的群体共享同一个单倍型,提示这些水系曾经有非常密切的联系。根据嵌套进化支序分析,中间黄颡鱼可能起源于峒中河、北仑河、防城河所在的广西与越南交界地区,并通过两条途径向华南沿海西部诸独立水系和海南岛扩散,在演化过程中,曾发生片断化事件,长距离建群和持续的分布区扩张。     

Phylogeography of the greenside darter complex, Etheostoma blennioides (Teleostomi: Percidae): a wide-ranging polytypic taxon

The greenside darter, Etheostoma blennioides (Teleostomi: Percidae), is a wide-ranging polytypic taxon that occurs throughout eastern North America. A previous morphological study recognized four subspecies (blennioides, newmanii, gutselli, and pholidotum), several morphological races, and three zones of morphological intergradation. We generated complete cytochrome b (1140bp) sequence data for 51 individuals from across the range of the greenside darter inclusive of all of the currently recognized taxa to assess genetic variation and taxonomic boundaries. Both maximum parsimony and mixed model Bayesian analyses resulted in two strongly supported deeply divergent clades including (1) a Tennessee River drainage clade, and (2) an Ohio River and Great Lakes basins, Interior Highlands, and Atlantic slope clade. Etheostoma blennius, a closely related congener, nested within the Tennessee River clade of E. blennioides, rendering the complex paraphyletic. Test of alternative topologies failed to support the current taxonomic designations. The inclusion of nuclear sequence data from intron 1 of the S7 ribosomal protein (523bp) from a subset of the populations was included to independently test whether the currently recognized taxa conform to distinct evolutionary lineages and also to clarify potential issues associated with ancestral hybridization. Although the nuclear data was less variable than the mitochondrial data, the monophyly of several of the subspecies could not be rejected.     

Morphological and molecular variation in the least madtom Noturus hildebrandi (Siluriformes: Ictaluridae), a Mississippi Embayment endemic: evidence for a cryptic lineage in the Hatchie River

Sheared principal component analysis of 40 morphometric characteristics measured for 146 individuals and relative frequencies of pigmentation patterns scored for 980 individuals of the least madtom Noturus hildebrandi, a diminutive catfish endemic to eastern lowland drainages of the Mississippi Embayment region of North America, suggested a clinal pattern of morphological variation extending across the range from north to south. DNA sequence data representing 90 individuals from the mitochondrial gene cytochrome b (cytb) analysed using Bayesian phylogenetic methods recovered four major haplotype clades, suggestive of a high degree of isolation by drainage. Individual gene trees of cytb and four additional nuclear loci as well as trees based on concatenated datasets of these genes consistently recovered a cryptic lineage of individuals from the Hatchie River drainage that is morphologically indistinguishable from surrounding populations. Gene‐tree analyses failed to recover a monophyletic N. hildebrandi with respect to Noturus baileyi. A coalescence‐based species tree analysis, however, did recover N. hildebrandi monophyly with high support, suggesting that relationships reflected in individual gene trees and concatenated datasets are in part artefacts of incomplete lineage sorting or an ancient introgressive event. Results are consistent with the hypothesis of an ancient connection between the Hatchie and Tennessee River systems. Current subspecific designations are of limited utility as they reflect morphological variation and are not entirely consistent with phylogeny. Discrepancies between the pattern of variation observed in the morphological and molecular data may be explained by recent local adaptation to individual stream conditions that masks deeper evolutionary divergences.     

Developmental responses to predation risk in morphologically defended mayflies

Densities and species composition of predators could affect morphological defences, larval development and the timing of emergence of their prey. To address this issue we studied the morphology and life history of an ephemerellid mayfly, Ephemerella invaria, from two streams in a deciduous forested drainage basin in central New York. Both streams contained predatory fish, but densities and species composition of fish differed. A field survey provided evidence that Ephemerella inhabiting a stream with 10 fish species and high relative densities of fish emerged several weeks earlier and at smaller sizes than Ephemerella inhabiting a nearby tributary with ~2 fish species and low relative densities of fish. However, the two populations of mayflies showed no differences in defensive morphology or growth rates. In laboratory rearing experiments, we exposed Ephemerella larvae from these two locations to fish chemical cues or control water (no fish) over 2 months to test whether differences in life histories could be attributed to fish. Fish cues induced faster larval development, but also smaller size of mature Ephemerella individuals from both high and low predator locations. Although shorter development times in more dangerous environments could increase larval survival, smaller size of females results in a fecundity cost associated with this life history shift. Consistent with the field studies, laboratory rearing experiments revealed no effects of fish cues on Ephemerella's morphological defences. These data suggest that variation in the density or species composition of predators may favour the evolution of developmental plasticity to reduce mortality in the larval environment.     

华南沿海西部美丽小条鳅基于线粒体控制区的种群遗传变异及亲缘地理格局

为研究其种群遗传变异和亲缘地理格局,分析了107尾采自华南西部和海南岛的12条水系的美丽小条鳅(Micronoema-cheilus pulcher Nichols)控制区934-938 bp的序列,其中有79个核苷酸变异位点。分子变异分析(AMOVA)表明,种群间的遗传变异占46.88%,种群内的遗传变异占55.06%。基于36个单倍型的系统树显示,12条水系的种群聚成两支。其中,广西沿海诸独立水系(防城河、峒中河、北仑河、南流江)和西江水系与广东漠阳江和潭江水系关系密切,而海南岛万泉河和南渡江与广东鉴江水系关系密切。根据嵌套进化枝系地理分析(NCPA)推测,防城河周边地区可能是美丽小条鳅的扩散中心,该物种可由此区域通过两条途径扩散:(1)沿西江水系向广西沿海独立水系至广东漠阳江和潭江水系扩散;(2)向海南岛诸水系再至雷州半岛的鉴江水系扩散。在演化过程中,曾发生片断化事件、长距离建群和持续的分布区扩张。