GENE FLOW VERSUS LOCAL ADAPTATION IN THE NORTHERN ACORN BARNACLE, SEMIBALANUS BALANOIDES: INSIGHTS FROM MITOCHONDRIAL DNA VARIATION

Abstract In reciprocal transplant experiments, Bertness and Gaines (1993) found that Semibalanus balanoides juveniles that had settled in an upper Narragansett Bay estuary survived better in that estuary that did juveniles from coastal localities. The observed pattern of survivorship led to the claim that local adaptation may result from a combination of limited gene flow between and strong selection within these habitats. Here we test the hypothesis that limited gene flow has led to habitat‐specific population differentiation using sequence and restriction fragment length polymorphism analyses of the mitochondrial DNA D‐loop region of S. balanoides. Samples were analyzed from replicated coastal and estuary localities in both Narragansett Bay, Rhode Island, and Damariscotta River, Maine. The patterns of F_ST indicate that gene flow between coast and estuary is extensive (Nm > 100) and is not lower in the estuary with lower flushing rates (Narragansett Bay). Given the high estimate of genetic exchange, adaptations for unpredictable environments seem more likely than local adaptation in this species because loci that respond to selection in one generation are essentially homogenized by the next seasons' settlement. Nevertheless, these estimates of neutral gene flow can help identify the strength of selection necessary for local adaptation to accumulate in Semibalanus.     

COLOR-PATTERN VARIATION IN LAKE ERIE WATER SNAKES: THE ROLE OF GENE FLOW

In an effort to clarify the evolutionary processes influencing color-pattern variation in Lake Erie island water snake (Nerodia sipedon) populations, rates of gene flow among island and mainland populations were estimated from patterns of allozymic variation detected using electrophoresis. Rates of gene flow were high with Nm, the number of migrants per generation, averaging 25.5 among island sites, 9.2 between the Ontario mainland and the islands, and 3.6 between the Ohio mainland and the islands. Based on estimates of current population size from mark-recapture work and of past population size extrapolated from the extent of shoreline habitat, values of m between island and mainland populations ranged from 0.0008–0.01. Synthesis of estimates of the rate of gene flow with information on inheritance of color pattern, the strength of natural selection, and population history supports the hypothesis that color-pattern variation in island populations results from a balance between gene flow and natural selection. However, depending on the mode of inheritance of color pattern, stochastic processes such as drift may have been important in the initial stages of differentiation between island and mainland populations.     

MITOCHONDRIAL DNA VARIATION IN THE CRITICALLY ENDANGERED VAQUITA PHOCOENA SINUS NORRIS AND MACFARLAND, 19581

The vaquita, Phocoena sinus, is one of two critically endangered cetacean species, and is listed as an endangered species in both the United States and Mexico. These listings result from a small population size, estimated to be 224 animals, and a rapid decline in abundance (18% per annum) thought to be caused by human activities. To characterize the genetic composition of the vaquita, we have sequenced a portion of the mitochondrial DNA control region from 43 individuals collected between 1985 and 1993. All animals had identical sequences. While low genetic variability has been reported for cetacean species, this complete lack of polymorphism in the control region is unique. This result is concordant with the hypothesis that the evolutionary history of the species includes a bottleneck or founder event, possibly at species inception, followed by a small long-term effective population size. We recommend parallel studies of nuclear variability, which could provide understanding of the basic biology of the vaquita for use in conservation efforts.     

MITOCHONDRIAL DNA VARIATION AND MATERNAL GENE FLOW AMONG HUMPBACK WHALES OF THE SOUTHERN HEMISPHERE

Samples of skin tissue were collected by biopsy darting from humpback whales ( Megaptera novaeangliae ) in six seasonal habitats representing three stocks and four regions: Groups IV (western Australia), V western component (eastern Australia), V eastern component (New Zealand and Tonga) and VI (the Antarctic Peninsula and Gorgona Island, Colombia, South America) of the Southern Hemisphere. A variable section of the mitochondrial DNA control region was amplified and sequenced from 84 of these individuals, distinguishing a total of 48 unique sequences ( i. e. , mtDNA nucleotypes). Phylogenetic reconstructions suggested that these nucleotypes form three clades, corresponding to those previously described in a world-wide survey of humpback whale mtDNA variation, although bootstrap support for two of the clades was relatively low (<50%). An analysis of variance adapted for molecular information showed significant differentiation of nucleotypes among the three Groups (Stocks) and heterogeneity of haplotype diversity among the four regions. A pattern of interchange within and between oceanic basins was demonstrated by the presence of shared identical nucleotypes among humpback whales in regions of the Southern and Northern Hemispheres.     

BIOCHEMICAL GENETICS OF FUNDULUS HETEROCLITUS (L.). VI. GEOGRAPHICAL VARIATION IN THE GENE FREQUENCIES OF 15 LOCI

Geographic variation in the gene frequencies corresponding to 15 polymorphic enzymes were studied in the common killifish Fundulus heteroclitus. Aat-A, Est-B, Fum-A, H6pdh-A, Mpi-A and Pgm-B showed clinal variation in allelic frequencies along the Atlantic coast of North America, while Aat-B, Ap-A, and the EST-C phenotypes did not. The clinal allelic variation of six previously examined loci (Ldh-B, Mdh-A, Gpi-B, Idh-A, Pgm-A, and 6-Pgdh-A) was extended to locations farther north. Gene diversity was lowest in the cold waters of northern latitudes and highest in the warmer waters of southern latitudes. The variety of clinal shapes and widths suggests that selection has affected the allelic distributions for at least some of these loci. This hypothesis is discussed with regard to the range contractions and extensions caused by the glacial advances and retreats during the Pleistocene.     

PHYLOGEOGRAPHY OF SPOTTED OWL (STRIX OCCIDENTALIS) POPULATIONS BASED ON MITOCHONDRIAL DNA SEQUENCES: GENE FLOW,GENETIC STRUCTURE,AND A NOVEL BIOGEOGRAPHIC PATTERN

Mitochondrial DNA control region sequences of spotted owls (Strix occidentalis) allowed us to investigate gene flow, genetic structure, and biogeographic relationships among these forest-dwelling birds of western North America Estimates of gene flow based on genetic partitioning and the phylogeography of haplotypes indicate substantial dispersal within three long-recognized subspecies. However, patterns of individual phyletic relationships indicate a historical absence of gene flow among the subspecies, which are essentially monophyletic. The pattern of haplotype coalescence enabled us to identify the approximate timing and direction of a recent episode of gene flow from the Sierra Nevada to the northern coastal ranges. The three subspecies comprise phylogenetic species, and the northern spotted owl (S. o. caurina) is sister to a clade of California (S. o. occidentalis) plus Mexican spotted owls (S o lucida); this represents a novel biogeographic pattern within birds. The California spotted owl had substantially lower nucleotide diversity than the other two subspecies; this result is inconsistent with present patterns of population density A causal explanation requires postulating a severe bottleneck or a selective sweep, either of which was confined to only one geographic region.     

GENE FLOW,REFUGIA, AND EVOLUTION OF GEOGRAPHIC VARIATION IN THE SONG SPARROW (MELOSPIZA MELODIA)

We surveyed mtDNA restriction-site variation in song sparrows taken from across their continental range. Despite marked geographic variation in size and plumage color, mtDNA variation was not geographically structured. Subspecies were not identifiable by mtDNA analysis. We suggest that postglaciation dispersal scattered mtDNA haplotypes across the continent, explaining the lack of mtDNA geographic patterns. Evolution of size and plumage coloration has probably proceeded faster than mtDNA evolution, leading to the well-structured continental pattern of morphological variation. We suggest that the nonordered geographic distribution of haplotypes reflects the recency of population establishment following completion of range expansion. Dispersal distance was estimated from the mtDNA data at 6.1 km per generation, an order of magnitude greater than that (0.3 km) estimated from demographic data. Island samples were not especially different from continental ones. Rooting the haplotype cladogram with a putative primitive haplotype identified Newfoundland and the Queen Charlotte Islands as potential sites of recent refugia. We question whether study of geographic variation in song sparrows leads to insights concerning speciation.     

ARE CHLOROPLAST AND MITOCHONDRIAL DNA VARIATION SPECIES INDEPENDENT IN OAKS?

Extensive introgression of cytoplasmic genomes across oak species is now a well-established fact. To distinguish between ancient hybridization events and ongoing introgression, a direct test for the existence of local exchanges is proposed. Such local exchanges must be comparatively recent, that is, contemporaneous with or later than the last postglacial recolonization. The test is applied to an extensive set of data comprising 377 pure or mixed populations (1744 individuals) of four white oak species in southern France. After demonstrating that local exchanges have occurred frequently between all species pairs, another test is performed to check if species status does nevertheless play some role in restricting cytoplasmic gene flow. The results vary according to the species pairs considered, and the observed pattern may be related to the ecology and/or compatibility of interspecific crosses. It is also shown that, for some of these oak species, the presence of related species in a population significantly influences the intraspecific diversity. Altogether, the results demonstrate that (1) intraspecific cytoplasmic gene flow varies according to the species, (2) interspecific cytoplasmic gene flow varies according to the species pair, and (3) both components of gene flow are at least partly related.     

SPATIAL POPULATION STRUCTURE IN THE WHIRLIGIG BEETLE DINEUTUS ASSIMILIS: EVOLUTIONARY INFERENCES BASED ON MITOCHONDRIAL DNA AND FIELD DATA

The spatial population structure of the pond-living water beetle Dineutus assimilis (Coleoptera: Gyrinidae) was investigated through a field study of population dynamics and dispersal, with a concurrent assessment of the spatial distribution of mitochondrial DNA (mtDNA) restriction-fragment-length polymorphism (RFLP). A comprehensive 2-yr survey within a 60-km² study area revealed pronounced fluctuations in local abundances, including extinctions and colonizations. The recapture of marked individuals showed that dispersal among ponds is frequent in both males and females and connects populations on a large geographic scale (maximum observed flight distance: 20 km). The population structure of D. assimilis is thus characterized by both pronounced genetic drift and frequent gene flow. Together, these two forces generate a pattern of very local and transient genetic differentiation. Mitochondrial DNA samples collected within a few kilometers indicate highly significant spatial structure, if newly founded demes or those that experienced recent bottlenecks are included. These results based on four demes within the study area were placed into a regional context by further samples collected at distances of 100 km and 200 km. F_st estimates computed on increasing spatial scales were variable but showed no increasing trend. Thus, gene flow exerts a strong homogenizing force over a wide geographic range but is counteracted locally by genetic drift. These findings highlight the need to supplement estimates of F_st with additional data to arrive at valid interpretations of the genetic information. More generally, this study raises questions about how to capture the relevant features of dynamic, subdivided populations to understand their evolutionary dynamics.     

GENETIC SUBDIVISIONS AMONG SMALL CANIDS: MITOCHONDRIAL DNA DIFFERENTIATION OF SWIFT,KIT, AND ARCTIC FOXES

Gene flow can effectively suppress genetic divergence among widely separated populations in highly mobile species. However, the same may not be true of species that typically disperse over shorter distances. Using mtDNA restriction-site and sequence analyses, we evaluate the extent of divergence among populations of two small relatively sedentary North American canids, the kit and swift foxes (genus Vulpes). We determine the significance of genetic differentiation among populations separated by distance and those separated by discrete topographic barriers. Our results show the among-population component of genetic variation in kit and swift foxes is large and similar to that of small rodents with limited dispersal ability. In addition, we found two distinct groupings of genotypes, separated by the Rocky Mountains, corresponding to the traditional division between kit and swift fox populations. Previous workers have characterized these morphologically similar populations either as separate species or subspecies. Our mtDNA data also suggest that kit and swift fox populations hybridize over a limited geographic area. However, the sequence divergence between kit and swift foxes is similar to that between these taxa and the arctic fox (Alopex lagopus), a morphologically distinct species commonly placed in a separate genus. This result presents a dilemma for species concepts, and we conclude that kit and swift foxes should be recognized as separate species.     

MITOCHONDRIAL DNA VARIATION WITHIN AND BETWEEN SPECIES OF THE PAPILIO MACHAON GROUP OF SWALLOWTAIL BUTTERFLIES

Species limits and phylogenetic relationships in the Papilio machaon species group are potentially confounded by a complex pattern of Pleistocene range fragmentation, hybridization, and ecological race formation. Mitochondrial DNA (mtDNA) restriction-site analysis has been used to define genetic affinities and genetic population structure within this species group. The distribution of mtDNA haplotypes generally confirms prior phylogenetic hypotheses and species delineations, but there is poor correspondence between ecological races and mtDNA haplotypes. The amount and distribution of mtDNA sequence variation within species vary among species, reflecting differences in current patterns of gene flow and/or historical population structure. In spite of wing pattern characters that ally them with P. polyxenes, both P. joanae and P. brevicauda have mtDNA that is closely related to that of P. machaon. We suggest that P. joanae and P. brevicauda are of hybrid origin.     

THE EFFECTS OF THE RELATIVE GEOGRAPHIC SCALES OF GENE FLOW AND SELECTION ON MORPH FREQUENCIES IN THE WALKING-STICK TIMEMA CRISTINAE

Gene frequencies in large populations are determined by a balance between selection and gene flow between neighborhoods of different selection regimes. This balance is affected by the area of the patches of a given selection regime relative to the gene-flow distance. If patches are small relative to gene-flow distance, similarity in the total area occupied by different patch types is a crucial condition for the stability of polymorphisms. However, if patches are larger than the gene-flow distance, then the relative area of different patch types is less important because of reduced gene flow resulting from isolation by distance. Two morphs (striped and unstriped) of the walking-stick Timema cristinae were each strongly associated with patches of distinct species of food plants on which they are most cryptic. The frequency of a morph was high on the plant on which it is most cryptic when either: (1) the area occupied by the food plant (patch) was very large; (2) the patch was completely isolated from other patches; or (3) the patch was larger than adjacent patches. Results (1) and (2) are consistent with isolation-by-distance models, and result (3) is consistent with Levene's multiple-niche polymorphism model.     

线粒体DNA和人类进化

线粒体ＤＮＡ（ｍｔＤＮＡ）由于自身比较独特的遗传特性（母系遗传、缺乏重组和进化速率高）而被广泛地应用于人类群体的起源和演化研究。通过对其全序列的限制性酶切和Ｄ－环高变区序列数据的分析,ｍｔＤＮＡ较好地阐明了人类学中诸如现代人类起源、人群过去动态的估计以及单个人群的区域性微分化和人口历史学等问题。综述了近年来世界各人群ｍｔＤＮＡ的研究进展、研究方法的改进、ｍｔＤＮＡ与核基因标记结果的异同、ｍｔＤＮＡ     

两种(鱼尝)科鱼类在长江和珠江流域Cytb基因序列变异性分析

采用PCR技术获得了长江和珠江流域的两种鲿科鱼类1137bp Cytb基因的片段,进行了序列测定,并用Mega软件进行了序列分析。研究发现两种鲿科鱼类的全序列从第十位碱基起有一个三联体密码子的缺失,推测为鲇形目鱼类Cytb序列所特有特征。黄颡鱼和大鳍鳠种群间的序列差异分别为0．4％和1．6％,介于一般淡水鱼类mtDNA种内序列的变异范围,而高于洄游鱼类序列变异。黄颡鱼属和鲮属Cytb基因序列间的差异为17．6％-18．0％,大大高于种间的序列差异率,也从分子水平上证实了二者属级分类单元的有效性。依据分子进化速率推测,长江和珠江两水系间大鳍鳠的分歧时间为距今27万年左右的更新世中晚期,黄颡鱼的分歧时间在距今7万年左右的更新世末期。推测造成二者有着不同分歧时间的可能原因是：大鳍鳠属暖水性鱼类,在第四纪冰川期间,由于全球性气候变冷,大鳍鳠退至长江以南,并在各水系间产生隔离,各自独立分化至今。而黄颡鱼具有很强的适应性,在第四纪数次冰川期间可能进行过广泛的交流。     

GENE FLOW AND SPECIES DELIMITATION: A CASE STUDY OF TWO PINE SPECIES WITH OVERLAPPING DISTRIBUTIONS IN SOUTHEAST CHINA

Species delimitation detected by molecular markers is complicated by introgression and incomplete lineage sorting between species. Recent modeling suggests that fixed genetic differences between species are highly related to rates of intraspecific gene flow. However, it remains unclear whether such differences are due to high levels of intraspecific gene flow overriding the spread of introgressed alleles or favoring rapid lineage sorting between species. In pines, chloroplast (cp) and mitochondrial (mt) DNAs are normally paternally and maternally inherited, respectively, and thus their relative rates of intraspecific gene flow are expected to be high and low, respectively. In this study, we used two pine species with overlapping geographical distributions in southeast China, P. massoniana and P. hwangshanensis, as a model system to examine the association between organelle gene flow and variation within and between species. We found that cpDNA variation across these two pine species is more species specific than mtDNA variation and almost delimits taxonomic boundaries. The shared mt/cp DNA genetic variation between species shows no bias in regard to parapatric versus allopatric species’ distributions. Our results therefore support the hypothesis that high intraspecific gene flow has accelerated cpDNA lineage sorting between these two pine species.     

ALLOZYME VARIATION IN A SNAIL (LITTORINA SAXATILIS)—DECONFOUNDING THE EFFECTS OF MICROHABITAT AND GENE FLOW

It is commonly observed that a restricted gene flow among populations of a species generates genetic differentiation in, for example, allozyme markers. However, recent studies suggest that microhabitat-specific variation may contribute to the total differentiation. To appreciate the relative contributions of geographic variation and habitat-specific variation, we sampled 42 subpopulations of the intertidal snail Littorina saxatilis from three different microhabitats (boulders, low and high rocky intertidal) on five small islands within a distance of 15 km. We used a modified orthogonal version of Nei's gene diversity analysis with a modified analysis of variance (ANOVA) that estimated the significance of habitat and geographic separation and the interaction between them. Between subpopulation differentiation (G_ST) was usually in the range of 5% to 10% but was exceptionally high in one locus (Aat; 53%). Genetic differentiation attributable to different habitats accounted for 10% to 81% (mean 35%) of the between subpopulation variation and was significant (P < 0.05) in six loci. Differentiation due to geographic separation accounted for 11% to 61% (mean 36%) and was significant (P < 0.05) in seven loci. Furthermore, three loci showed interactions between habitat and island, suggesting varying effects of habitat in different islands. Microhabitat-specific variation, probably through spatially varying fitness, seems particularly likely in Aat and Pgm-2. Moderate levels of habitat associated variation added to the observed differentiation due to gene flow in Pgi, Pnp, and Pgm-1, whereas in the remaining three loci either the habitat effect was confused by strong habitat-island interaction (Ark) or was virtually absent (Pep and Mpi).     

MAINTENANCE OF ECOLOGICALLY SIGNIFICANT GENETIC VARIATION IN THE TIGER SWALLOWTAIL BUTTERFLY THROUGH DIFFERENTIAL SELECTION AND GENE FLOW

Differential selection in a heterogeneous environment is thought to promote the maintenance of ecologically significant genetic variation. Variation is maintained when selection is counterbalanced by the homogenizing effects of gene flow and random mating. In this study, we examine the relative importance of differential selection and gene flow in maintaining genetic variation in Papilio glaucus. Differential selection on traits contributing to successful use of host plants (oviposition preference and larval performance) was assessed by comparing the responses of southern Ohio, north central Georgia, and southern Florida populations of P. glaucus to three hosts: Liriodendron tulipifera, Magnolia virginiana, and Prunus serotina. Gene flow among populations was estimated using allozyme frequencies from nine polymorphic loci. Significant genetic differentiation was observed among populations for both oviposition preference and larval performance. This differentiation was interpreted to be the result of selection acting on Florida P. glaucus for enhanced use of Magnolia, the prevalent host in Florida. In contrast, no evidence of population differentiation was revealed by allozyme frequencies. F_ST-values were very small and Nm, an estimate of the relative strengths of gene flow and genetic drift, was large, indicating that genetic exchange among P. glaucus populations is relatively unrestricted. The contrasting patterns of spatial differentiation for host-use traits and lack of differentiation for electrophoretically detectable variation implies that differential selection among populations will be counterbalanced by gene flow, thereby maintaining genetic variation for host-use traits.     

APPARENT SELECTIVE NEUTRALITY OF MITOCHONDRIAL DNA SIZE VARIATION: A TEST IN THE DEEP-SEA SCALLOP PLACOPECTEN MAGELLANICUS

Animal mitochondrial DNA (mtDNA) is believed to have evolved under intense selection for economy of the size of the molecule. Among scallop species mtDNA size may vary by a factor of two and among conspecific individuals by as much as 25%. We have examined the possibility that large mtDNA size differences may be associated with fitness in the deep sea scallop Placopecten magellanicus by comparing shell lengths of individuals with different copy numbers of a large mtDNA repeated sequence. Among juvenile cohorts of same age, shell length is known to be a good index of overall fitness in marine bivalves and it is shown here to be affected by differences in nuclear genotype, expressed as the degree of enzyme heterozygosity. We have observed no correlation between shell length and mtDNA length and interpreted this to mean that variation in the size of animal mtDNA is effectively neutral to the forces of natural selection acting on the individual. This type of mtDNA variation must, therefore, be explained in terms of biases in the molecular mechanisms causing expansion or contraction of the molecule, differential replication rates of mtDNA molecules of different size, and the stochastic assortment of mtDNA size classes among individuals.     

INTROGRESSION IN THE DROSOPHILA SUBOBSCURA—D. MADEIRENSIS SISTER SPECIES: EVIDENCE OF GENE FLOW IN NUCLEAR GENES DESPITE MITOCHONDRIAL DIFFERENTIATION

Species hybridization, and thus the potential for gene flow, was once viewed as reproductive mistake. However, recent analysis based on large datasets and newly developed models suggest that gene exchange is not as rare as originally suspected. To investigate the history and speciation of the closely related species Drosophila subobscura, D. madeirensis, and D. guanche, we obtained polymorphism and divergence data for 26 regions throughout the genome, including the Y chromosome and mitochondrial DNA. We found that the D. subobscura X/autosome ratio of silent nucleotide diversity is significantly smaller than the 0.75 expected under neutrality. This pattern, if held genomewide, may reflect a faster accumulation of beneficial mutations on the X chromosome than on autosomes. We also detected evidence of gene flow in autosomal regions, while sex chromosomes remain distinct. This is consistent with the large X effect on hybrid male sterility seen in this system and the presence of two X chromosome inversions fixed between species. Overall, our data conform to chromosomal speciation models in which rearrangements are proposed to serve as gene flow barriers. Contrary to other observations in Drosophila, the mitochondrial genome appears resilient to gene flow in the presence of nuclear exchange.