新疆蒲公英属有性生殖与无融合生殖植物的调查研究

以新疆２２种蒲公英的１８６份腊叶标本为材料,观察统计了每份标本的花粉败育率、花粉粒大小变异的标准方差。标准方差小于３、败育率小于２０％的花粉ＰＲ变异型,是有性生殖植物的特征;标准方差大于３、败育率大于４０％的花粉ＰＳ变异型,是无融合或兼性无融合植物的特征;不产生花粉的ＰＡ变异型,则是雄性不育植物的特征。在所观察的标本中,有４种蒲公英植物出现了有性生殖个体,其余为无融合或兼性无融合类。此外,还就蒲公     

CONTRIBUTIONS OF SEXUAL AND ASEXUAL REPRODUCTION TO POPULATION STRUCTURE IN THE CLONAL SOFT CORAL,ALCYONIUM RUDYI

Numerous studies of population structure in sessile clonal marine invertebrates have demonstrated low genotypic diversity and nonequilibrium genotype frequencies within local populations that are monopolized by relatively few, highly replicated genets. All of the species studied to date produce planktonic sexual propagules capable of dispersing long distances; despite local genotypic disequilibria, populations are often panmictic over large geographic areas. The population structure paradigm these species represent may not be typical of the majority of clonal invertebrate groups, however, which are believed to produce highly philopatric sexual propagules. I used allozyme variation to examine the population structure of the temperate soft coral, Alcyonium rudyi, a typical clonal species whose sexually produced larvae and asexually produced ramets both have very low dispersal capabilities. Like other clonal plants and invertebrates, the local population dynamics of A. rudyi are dominated by asexual reproduction, and recruitment of new sexually produced genets occurs infrequently. As expected from its philopatric larval stage, estimates of genetic differentiation among populations of A. rudyi were highly significant at all spatial scales examined (mean θ = 0.300 among 20 populations spanning a 1100-km range), suggesting that genetic exchange seldom occurs among populations separated by as little as a few hundred meters. Mapping of multilocus allozyme genotypes within a dense aggregation of A. rudyi ramets confirmed that dispersal of asexual propagules is also very limited: members of the same genet usually remain within < 50 cm of one another on the same rock surface. Unlike most previously studied clonal invertebrates, populations of A. rudyi do not appear to be dominated by a few widespread genets: estimates of genotypic diversity (G_o) within 20 geographically distinct populations did not differ from expectations for outcrossing, sexual populations. Despite theoretical suggestions that philopatric dispersal combined with typically small effective population sizes should promote inbreeding in clonal species, inbreeding does not appear to contribute significantly to the population structure of A. rudyi. Genet genotype frequencies conformed to Hardy-Weinberg expectations in all populations, and inbreeding coefficients (f) were close to zero. In general, the population structure of A. rudyi did not differ significantly from that observed among outcrossing sexual species with philopatric larval dispersal. Age estimates suggest, however, that genets of A. rudyi live for many decades. Genet longevity may promote high genotypic diversity within A. rudyi populations and may be the most important evolutionary consequence of clonal reproduction in this species and the many others that share its dispersal characteristics.     

GENETIC POPULATION STRUCTURE AND LEVELS OF GENE FLOW IN THE STREAM DWELLING WATERSTRIDER,AQUARIUS (=GERRIS) REMIGIS (HEMIPTERA: GERRIDAE)

Gene flow, in combination with selection and drift, determines levels of differentiation among local populations. In this study we estimate gene flow in a stream dwelling, flightless waterstrider, Aquarius remigis. Twenty-eight Aquarius remigis populations from Quebec, Ontario, New Brunswick, Iowa, North Carolina, and California were genetically characterized at 15 loci using starch gel electrophoresis. Sampling over two years was designed for a hierarchical analysis of population structure incorporating variation among sites within streams, streams within watersheds, watersheds within regions, and regions within North America. Hierarchical F statistics indicated that only sites within streams maintained enough gene flow to prevent differentiation through drift (Nm = 27.5). Above the level of sites within streams gene flow is highly restricted (Nm ≤ 0.5) and no correlation is found between genetic and geographic distances. This agrees well with direct estimates of gene flow based on mark and recapture data, yielding an N_e of approximately 170 individuals. Previous assignment of subspecific status to Californian A. remigis is not supported by genetic distances between those populations and other populations in North America. Previous suggestion of specific status for south-eastern A. remigis is supported by genetic distances between North Carolina populations and other populations in North America, and a high proportion of region specific alleles in the North Carolina populations. However, because of the high degree of morphological and genetic variability throughout the range of this species, the assignment of specific or subspecific status to parts of the range may be premature.     

SPECIES-WIDE POPULATION STRUCTURE IN A SOUTHEASTERN U.S. FRESHWATER FISH,HETERANDRIA FORMOSA: GENE FLOW AND BIOGEOGRAPHY

The phylogeography of the freshwater fish fauna of the southeastern United States has almost achieved paradigm status in evolutionary biology (Avise 1992), and the major geographic features responsible for shaping species distributions are well-characterized. Nevertheless, variation among species in distributions of allele or haplotype frequencies suggests that species-specific processes (e.g., migration) may also play a role in establishing those distributions. There has also been relatively little investigation into how population structure may differ among subregions in the Southeast, for example, on the Florida peninsula versus the U.S. mainland to the northwest and/or northeast. The geology of the peninsula is such that both physical and biotic fluctuations may have been (and still be) particularly important in establishing the population structure of freshwater taxa. This possibility leads to two interesting questions in population genetics. (1) Does gene flow in freshwater species of the region better approximate a one- or two-dimensional pattern? (2) Are populations on the peninsula farther from migration-genetic drift equilibrium than their counterparts on the mainland? These questions were addressed by examining the population strucuture of a livebearing fish, Heterandria formosa; several features of the biology of the species make it particularly likely that recent gene flow has been important in its evolution. I surveyed electrophoretic variation in 34 populations distributed throughout the species range. The phylogeographic patterns observed are in general concordance with those found in other species, although with some differences. A two-dimensional hypothesis of gene flow on the Florida peninsula better explains the data than does a one-dimensional one. There is no evidence that populations on the peninsula are farther from migration-drift equilibrium than those to the northwest. Populations in the northeast have lower genetic diversity than those to the south and west and show no isolation by distance; those results are consistent with a recent range expansion into the northeast, although smaller historical effective population sizes could also explain the pattern.     

DYNAMICS OF POLYPLOID FORMATION IN TRAGOPOGON (ASTERACEAE): RECURRENT FORMATION,GENE FLOW,AND POPULATION STRUCTURE

Polyploidy is a major feature of angiosperm evolution and diversification. Most polyploid species have formed multiple times, yet we know little about the genetic consequences of recurrent formations. Among the clearest examples of recurrent polyploidy are Tragopogon mirus and T. miscellus (Asteraceae), each of which has formed repeatedly in the last ～80 years from known diploid progenitors in western North America. Here, we apply progenitor‐specific microsatellite markers to examine the genetic contributions to each tetraploid species and to assess gene flow among populations of independent formation. These data provide fine‐scale resolution of independent origins for both polyploid species. Importantly, multiple origins have resulted in considerable genetic variation within both polyploid species; however, the patterns of variation detected in the polyploids contrast with those observed in extant populations of the diploid progenitors. The genotypes detected in the two polyploid species appear to represent a snapshot of historical population structure in the diploid progenitors, rather than modern diploid genotypes. Our data also indicate a lack of gene flow among polyploid plants of independent origin, even when they co‐occur, suggesting potential reproductive barriers among separate lineages in both polyploid species.     

THE RELATIVE IMPORTANCE OF HISTORICAL EVENTS AND GENE FLOW ON THE POPULATION STRUCTURE OF A MEDITERRANEAN RAGWORT,SENECIO GALLICUS (ASTERACEAE)

Comparisons of cytoplasmic and nuclear diversity within and among natural plant populations have the potential to distinguish the relative influences of seed and pollen dispersal on contemporary gene flow, or alternatively, may permit inferences of the colonization history of a species via seed. We examined patterns of cpDNA and allozyme variation in Senecio gallicus, a diploid, annual plant that occurs in both coastal and ruderal inland areas of the Iberian Peninsula and southern France. The species appears to have a strong propensity for long-distance seed dispersal. Five cpDNA haplotypes were found by RFLP analysis among a sample of 111 individuals derived from 11 populations. Differences in haplotype frequencies across populations were most evident with respect to a dramatic increase in the frequency of a derived haplotype from coastal to inland localities. The level of cpDNA differentiation among populations within the inland group (θ₀ = 0.07) was significantly less than that seen within the coastal group (θ₀ = 0.41). In contrast, for allozymes, no significant difference in population structure was evident between collections from coastal and inland habitats. At the rangewide geographic scale, there was only a very weak association between inferred levels of gene flow and geographic distance for cpDNA, and no such association was found for allozymes. It appears that while seed movement in the species might be sufficiently great to disturb the pattern of isolation by distance for cpDNA, it cannot fully account for the nearly randomized spatial structure at polymorphic allozyme loci. It is suggested that isolation of populations in Atlantic-Mediterranean coastal refugia during previous glacial maxima, and the effects of subsequent colonization events in inland areas, have had an important effect on molding the present genetic structure of the species.     

POPULATION STRUCTURE AND LEVELS OF GENE FLOW IN THE MEDITERRANEAN LAND SNAIL ALBINARIA CORRUGATA (PULMONATA: CLAUSILIIDAE)

The amount of gene flow among local populations partly determines the relative importance of genetic drift and natural selection in the differentiation of such populations. Land snails, because of their limited powers for dispersal, may be particularly likely to show such differentiation. In this study, we directly estimate gene flow in Albinaria corrugata, a sedentary, rock-dwelling gastropod from Crete, by mark-recapture studies. In the same area, 23 samples were taken and studied electrophoretically for six polymorphic enzyme loci. The field studies indicate that the population structure corresponds closely to the stepping-stone model: demes are present on limestone boulders that are a few meters apart, and dispersal takes place mainly between adjacent demes. Average deme size (N) is estimated at 29 breeding individuals and the proportion of migrants per generation at 0.195 (Nm = 5.7). We find no reason to assume long-distance dispersal, apart from dispersal along occasional stretches of suitable habitat. Genetic subdivision of the population, as derived from F_ST values, corresponds to the direct estimate only at the lowest spatial level (distance between sample sites < 10 m), where values for Nm of 5.4 and 17.6 were obtained. In contrast, at the larger spatial scales, F_ST values give gene-flow estimates that are incompatible with the expected amount of gene flow at these scales. We explain these discrepancies by arguing that gene flow is in fact extremely limited, making correct estimates of Nm from F_ST impossible at the larger spatial scales. In view of these low levels of gene flow, it is concluded that both genetic drift and natural selection may play important roles in the genetic differentiation of this species, even at the lowest spatial scales.     

THE INFLUENCE OF POPULATION SIZE AND ISOLATION ON GENE FLOW BY POLLEN IN SILENE ALBA

In a series of experiments conducted over two seasons, we used arrays of experimental populations to examine the effects of flower number and distance between patches on gene flow by pollen. For this study we used the dioecious, short-lived perennial plant Silene alba (Caryophyllaceae). This species lives in disturbed roadside and agricultural habitats and displays a weedy population dynamic with high colonization and extinction rates. The motivation for the study was to understand what factors may be influencing genetic connectedness among newly colonized populations within a regional metapopulation. By using experimental populations composed of genotypes homozygous at a diagnostic locus, it was possible to identify explicitly pollen movement into a focal patch as a function of flower number and distance to the nearest neighboring patch. Overall, the mean immigration rate (measured as the fraction of seeds sired by males outside the focal patch) at 20 m was just over 47%, whereas at 80 m immigration rates were less than 6%. In addition, by knowing the context in which each of these gene-flow events occurred, it was possible to understand some of the factors that influenced the exchange of genes. Both the number of flowers in the focal population (target) and in the neighboring populations (source) had a significant effect on the frequency of gene flow. Our experimental data also demonstrate that factors that influence gene flow at one spatial scale may not act in the same way at another. Specifically, the influence of target size and the relative size of the target and source patches on rates of gene flow depended on whether the patches were separated by 20 m or 80 m. These data suggest that the patterns of gene flow within a metapopulation system can be complex and may vary within a growing season.     

POPULATION GENETIC STRUCTURE AND GENE FLOW ACROSS ARID VERSUS MESIC ENVIRONMENTS: A COMPARATIVE STUDY OF TWO PARAPATRIC SENECIO SPECIES FROM THE NEAR EAST

To shed light on the potential effects of xeric/arid versus mesic environments on plant population genetic structure and patterns of gene flow, we have compared allozyme and cpDNA haplotype variation in populations of two closely related, highly outcrossed, and largely wind-dispersed winter annuals of Senecio (Asteraceae). The species form a distinctive zone of parapatric distribution in the Near East by differing in their ecogeographical regimes. Senecio vernalis mainly thrives in the mesic Mediterranean life zone of Israel, whereas S. glaucus inhabits either xeric maritime or arid (semi-) desert sites. Significant differences in allozymic population subdivision among S. vernalis (θ_n = 0.04; Nm_n = 5.85) and S. glaucus (θ_n = 0.12; Nm_n = 1.85) largely resulted from topogeographical substructuring present within the latter species. Because of the similarity of within-region estimates of population structure for S. glaucus with those measured among populations of S. vernalis, it appears unlikely that ecological “aridity” factors per se are important in influencing levels of population differentiation in these species. Based on hierarchical F-statistics and tests of isolation by distance, we further conclude that geographical topologies influence the level and mode of nuclear gene flow (via pollen and/or seed) among and within subsets of S. glaucus populations, although without providing a complete barrier to interregional dispersal (dNm_reg = 2.16) and without promoting allopatric differentiation via drift. The allozymic data further suggested that S. vernalis and S. glaucus form a zone of secondary contact in the Near East, accompanied by an almost complete interspecific barrier to nuclear gene flow (dnNm_sp = 0.253). However, to account for the considerable sharing of cpDNA haplotypes, both at the intra- and interspecific level, it is necessary to invoke either (1) selection acting against alien nuclear but not cytoplasmic DNA; or (2) the sporadic immigration of cpDNA via seed with large homogenizing effects on cytoplasmic population structure over time.     

ECOLOGICAL AND HISTORICAL ASSOCIATIONS OF GENE FLOW IN DARTERS (TELEOSTEI: PERCIDAE)

Life history should relate to gene flow (Nm) through its influence on dispersal and effective population size. Comparative studies designed to elucidate this relationship must contend with historical events that can yield misleading estimates of gene flow and statistical problems associated with inclusion of life-history traits correlated with phylogeny. We studied the relationships of life-history characters and gene flow in 15 species of darters, a monophyletic group of stream fishes. Populations of coexisting species were sampled in three geographic regions with different Pleistocene glaciation histories. Gene flow was estimated indirectly from allozymes using two methods, 8 and private alleles. Isolation-by-distance was also tested using regression of pairwise estimates of gene flow (M?) on distance. Theta and private-alleles methods produced congruent estimates of Nm, except in a study region hypothesized to have been historically fragmented and then united following Pleistocene glaciation. A relatively weak association between life-history traits and Nm (based on θ) was observed when species from the historically fragmented region were included in stepwise regression analysis, because Nm was low despite life-history differences among taxa in this region. Excluding observations from this region produced stronger associations between clutch size and Nm (r² = 0.57), and between female size, egg size, and Nm (r² = 0.95). Additional analyses that corrected for female body size and phylogenetic nonindependence agreed that darters with high fecundity and small eggs exhibited high gene flow, whereas darters with small clutches and large eggs had low gene flow. The latter combination of life-history traits primarily is exhibited in species from headwater habitats where parental investment presumably confers survivorship on offspring. Reduced gene flow and genetic divergence among demes appear to be evolutionary consequences of this strategy.     

GENETIC STRUCTURE AND MALE-MEDIATED GENE FLOW IN THE GHOST BAT (MACRODERMA GIGAS)

The Australian ghost bat is a large, opportunistic carnivorous species that has undergone a marked range contraction toward more mesic, tropical sites over the past century. Comparison of mitochondrial DNA (mtDNA) control region sequences and six nuclear microsatellite loci in 217 ghost bats from nine populations across subtropical and tropical Australia revealed strong population subdivision (mtDNA φ_ST = 0.80; microsatellites UR_ST = 0.337). Low-latitude (tropical) populations had higher heterozygosity and less marked phylogeographic structure and lower subdivision among sites within regions (within Northern Territory [NT] and within North Queensland [NQ]) than did populations at higher latitudes (subtropical sites; central Queensland [CQ]), although sampling of geographically proximal breeding sites is unavoidably restricted for the latter. Gene flow among populations within each of the northern regions appears to be male biased in that the difference in population subdivision for mtDNA and microsatellites (NT φ_ST = 0.39, UR_ST = 0.02; NQ φ_ST = 0.60, UR_ST = ?0.03) is greater than expected from differences in the effective population size of haploid versus diploid loci. The high level of population subdivision across the range of the ghost bat contrasts with evidence for high gene flow in other chiropteran species and may be due to narrow physiological tolerances and consequent limited availability of roosts for ghost bats, particularly across the subtropical and relatively arid regions. This observation is consistent with the hypothesis that the contraction of the species' range is associated with late Holocene climate change. The extreme isolation among higher-latitude populations may predispose them to additional local extinctions if the processes responsible for the range contraction continue to operate.     

LONG-DISTANCE GENE FLOW IN THE SEDENTARY BUTTERFLY,EUPHILOTES ENOPTES (LEPIDOPTERA: LYCAENIDAE)

The relationship between gene flow and geographic proximity has been assessed for many insect species, but dispersal distances are poorly known for most of these. Thus, we are able to assess the concordance between vagility and gene flow for only a few species. In this study, I documented variation at six allozyme loci among Washington and Oregon populations of the sedentary, patchily distributed, lycaenid butterfly, Euphilotes enoptes (Boisduval) to assess whether the relationship between gene flow and geographic distance is consistent with the dispersal biology of this species. Both a phenogram based on genetic distances between populations and a regression analysis of gene flow estimates on geographic distances showed a pattern consistent with genetic isolation by distance. Many estimates of gene flow among pairs of populations separated by more than 100 km exceeded the equivalent of 10 individuals exchanged per generation, a value much greater than would be predicted from the limited dispersal ability of this species. However, based on the allozyme data, genetic neighborhood size was estimated to be approximately 39 individuals, a value that is consistent with poor vagility. The results of this study speak to the power of stepping-stone gene flow among populations and are compared to the results of other studies that have examined the relationship between dispersal and gene flow in sedentary insects.     

FREQUENCY-DEPENDENT SEED PRODUCTION AND HYBRIDIZATION RATES: IMPLICATIONS FOR GENE FLOW BETWEEN LOCALLY ADAPTED PLANT POPULATIONS

Gene flow between genetically distinct plant populations can have significant evolutionary consequences. It can increase genetic diversity, create novel gene combinations, and transfer adaptations from one population to another. This study addresses the roles of frequency-dependent selection and mating system in gene exchange between two subspecies of Gilia capitata (Polemoniaceae). Long-distance migrants are likely to be rare in new habitats, and the importance of immigrant frequency to fitness, gene exchange, and ultimately introgression, has not been explored. To test for the importance of frequency in migration, a field experiment was conducted in which artificial populations (arrays) composed of different mixtures of the two subspecies were placed in the home habitats of both. Female function (seed production) and a portion of male function (hybridization rate) were compared for the two subspecies to assess the potential for gene exchange and introgression between them. Individual fitness (through both hybridization and seed production) for the inland subspecies varied with its frequency as an immigrant at the coastal site. Rare immigrants produced fewer seeds and fathered fewer hybrid offspring. In contrast, both forms of reproductive function were frequency independent for the coastal subspecies when it was an immigrant at the inland site. Seed production was high and insensitive to frequency, and immigrants from the coast never successfully fertilized the inland subspecies' seeds. To control for the effects of frequency-dependent pollinator behavior in the field, hand crosses were performed in the greenhouse using a range of pollen mixtures. The greenhouse experiment demonstrated that cross-fertilization is possible in only one direction, that cross-pollination in the other direction is only partially successful, and that pollen from the coastal subspecies has a strong negative effect on seed production by the inland subspecies. Experimental pollen supplementation in the field verified both the unilateral incompatibility and the negative effect of coastal pollen on inland plant seed production observed in the greenhouse. Contrasts between field array and greenhouse results suggest that pollinator behavior and other ecological factors act to exaggerate reproductive barriers between the two subspecies. In this system, immigrant frequency interacts with reproductive biology and pollinator ecology to enhance gene flow between the populations in one direction, while restricting gene establishment and introgression in the other direction.     

UNDERSTANDING THE POPULATION GENETIC STRUCTURE OF GLEDITSIA TRIACANTHOS L.: THE SCALE AND PATTERN OF POLLEN GENE FLOW

Spatial and temporal patterns of gene flow determine the extent to which populations can differentiate from one another as a result of natural selection or genetic drift. In this study, we investigated pollen-mediated gene flow in two eastern Kansas populations of the subdioecious tree species, Gleditsia triacanthos L. (Leguminosae), or honeylocust. In 2 yr at each site, we used paternity-exclusion analysis to estimate the proportion of seeds sired by immigrant pollen. We also used a single-parent and parent-pair exclusion analysis on naturally established seedlings and saplings to estimate gene flow into one site over a 12-yr period and into the second site over a 22-yr period. Results of both analyses showed high minimum estimates of pollen gene flow into each site (17%–30%). In each population, we found significantly less gene flow in years of high fruit production than in years of low fruit production, but in one population, we observed little variation in gene-flow rates among age classes of seedlings and saplings. The level of pollen gene flow showed weak negative dependence on the relative isolation distances of the maternal trees sampled (140–240 m at one site vs. 85–120 m at the second site), and gene-flow estimates from naturally established juveniles were very similar at the two sites. Within populations, a multiple regression model showed that maximum-likelihood estimates of male fertility were negatively associated with distances between mates and positively associated with male size as measured by stem diameter. In neither population, however, did the regression explain more than 16% of the total variation in male fertilities.     

PHENOLOGICAL ISOLATION,GENE FLOW AND DEVELOPMENTAL DIFFERENCES AMONG LOW- AND HIGH-ELEVATION POPULATIONS OF EUPHILOTES ENOPTES (LEPIDOPTERA: LYCAENIDAE)

Populations of the specialist herbivore, Euphilotes enoptes (Lepidoptera: Lycaenidae), along three elevational transects in the mountains of central Washington state, differed markedly in the phenology of adult flight. In spite of this apparent limitation to gene flow, six allozyme loci revealed substantial gene exchange among populations along these gradients. The elevational difference, and thus the phenological difference, between populations has not influenced the extent of gene flow between them. Because the direct exchange of genes between low- and high-elevation populations is very unlikely, gene flow between them has probably occurred in a stepwise fashion via intermediate populations. It is hypothesized that such gene flow has been biased in an uphill direction due to the combined effects of source size and oviposition behavior. Adult emergence times of populations in the same region are positively correlated with elevation in a nonlinear fashion, consistent with the hypothesis that gene flow from low-elevation populations has been swamping selection at higher altitudes.     

HIERARCHICAL GENETIC STRUCTURE AND GENE FLOW IN MACROGEOGRAPHIC POPULATIONS OF THE EASTERN TENT CATERPILLAR (MALACOSOMA AMERICANUM)

Genetic structure and inferred rates of gene flow in macrogeographic populations of the eastern tent caterpillar Malacosoma americanum were analyzed at two hierarchical scales: local demes and regional subpopulations. Wright's F-statistics were used to estimate population genetic structure using multilocus genotypic data generated electrophoretically. Estimated values of F_ST and the distribution of private alleles were then used to obtain indirect estimates of gene flow. We found modest, though significant, genetic structure at both spatial scales, a pattern consistent with high rates of gene flow over the large distances involved. Modest values obtained for Nei's genetic distance also suggested high levels of gene flow across the range of this species, although some gene-flow restriction resulting from isolation by distance was suggested by a positive regression of genetic distance on geographic distance. The observed homogeneity at enzyme loci across the range of M. americanum parallels the reported uniformity in morphology, suggesting a general absence of local genetic differentiation in this widely distributed species. The genetic homogeneity observed in this wide-ranging insect is discussed in terms of organism-specific environmental experience at different spatial scales. Some organisms occupying apparently heterogeneous environments may ameliorate unsuitable local conditions through microhabitat selection or behavioral modification of their microenvironment. This may be accomplished in M. americanum through group shelter construction and behavioral thermoregulation, closely tying thermoregulation to social biology in this species. If in this way the tent helps produce an effectively homogeneous environment for this species across its extensive range, this system may provide a unique example of how social behavior can influence the distribution of genetic variation in a population.     

ECOLOGICAL BARRIERS TO GENE FLOW BETWEEN RIPARIAN AND FOREST SPECIES OF AINSLIAEA (ASTERACEAE)

Understanding the role of habitat‐associated adaptation in reducing gene flow resulting in population differentiation and speciation is a major issue in evolutionary biology. We demonstrate a significant role for habitat divergence in species isolation between two naturally hybridizing riparian and nonriparian plants, Ainsliaea faurieana and A. apiculata (Asteraceae), on Yakushima Island, Japan. By analyzing the fine‐scale population structure at six sympatric sites, we found that variations in leaf shape, geography, light conditions, and genotype were strongly correlated across riverbank–forest transitions. No evidence of effective gene flow was found between the two species across the majority of the transition zones, although the NewHybrid clustering analysis confirmed interspecific hybridization. However, a relatively high level of gene flow was observed across one zone with a more diffuse ecotone and intermediate flooding and light conditions, possibly generated by human disturbances. These results suggest that the barriers to gene flow between the riparian and forest species are primarily ecological. Additional common garden experiments indicated that the two species are adaptively differentiated to contrasting flooding and light environments. Overall, our study suggests that adaptations to different habitats can lead to the formation of reproductive isolating barriers and the maintenance of distinct species boundaries.     

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.     

SEXUAL SELECTION AND FITNESS VARIATION IN A POPULATION OF SMALLMOUTH BASS,MICROPTERUS DOLOMIEUI (PISCES: CENTRARCHIDAE)

Monogamy is often presumed to constrain mating variance and restrict the action of sexual selection. We examined the reproductive patterns of a monogamous population of smallmouth bass (Micropterus dolomieui), and attempted to identify sources of within-season fitness variation among females and known-age males. Many males did not acquire a nest site, and many territorial males were unsuccessful in acquiring a mate. The likelihood that territorial males mated depended on several aspects of nest sites. Mated males of age three were larger than the average size of age-three males in the population. The mean sizes of age-four and age-five mated males were not different from the average of same-age males in the population. Thus, selection resulting from the acquisition of a mate favored large size among only age-three males. Timing of nest construction and breeding among territorial males was negatively related to male size and did not depend on male age after taking male size into account. Indirect evidence (numbers of eggs deposited in nests) suggests that the timing of spawning among females was also negatively related to female size. Fertility selection favored early reproduction within the season by males of all ages, but large male size was favored among only age-four males. The combined early breeding of fecund females and female mate choice of large males may explain the positive correlation between the size of age-four males and the number of eggs acquired. Despite large differences of female fecundity, however, the variance of relative mate number contributed about two times more than the variance of relative fertility among females to the total variance of relative fitness within each sex.