Refugial isolation versus ecological gradients

Hypotheses for divergence and speciation in rainforests generally fall into two categories: those emphasizing the role of geographic isolation and those emphasizing the role of divergent selection along gradients. While a majority of studies have attempted to infer mechanisms based on the pattern of species richness and congruence of geographic boundaries, relatively few have tried to simultaneously test alternative hypotheses for diversification. Here we discuss four examples, taken from our work on diversification of tropical rainforest vertebrates, in which we examine patterns of genetic and morphological variation within and between biogeographic regions to address two alternative hypotheses. By estimating morphological divergence between geographically contiguous and isolated populations under similar and different ecological conditions, we attempt to evaluate the relative roles of geographic isolation and natural selection in population divergence. Results suggest that natural selection, even in the presence of appreciable gene flow, can result in morphological divergence that is greater than that found between populations isolated for millions of years and, in some cases, even greater than that found between congeneric, but distinct, species. The relatively small phenotypic divergence that occurs among long-term geographic isolates in similar habitats suggests that morphological divergence via drift may be negligible and/or that selection is acting to produce similar phenotypes in populations occupying similar habitats. Our results demonstrate that significant phenotypic divergence: (1) is not necessarily coupled with divergence in neutral molecular markers; and (2) can occur without geographic isolation in the presence of gene flow.     

GENE FLOW AND MITOCHONDRIAL DNA VARIATION IN THE KILLIFISH,FUNDULUS HETEROCLITUS

Two subspecies of killifish, Fundulus heteroclitus, inhabit the Atlantic coast of the United States. The contact zones between them are typified by morphological, physiological, nuclear gene, and mitochondrial DNA clines. Considerable debate exists about the importance of restricted gene flow and natural selection in maintaining these clines. To evaluate the relative importance of these two evolutionary forces we employed analysis of mtDNA as an independent measure of gene flow. Solutions of equilibrium diffusion equations suggested that killifish dispersal must be less than 2 km to maintain previously observed allozyme frequency clines in the absence of strong selection. To determine whether populations separated by distances greater than 2 km accumulate significant genetic differences we examined a total of N = 480 individuals from five killifish populations spanning a continuous shoreline distance of 8.4 km. Distribution of the 25 detected haplotypes indicated that most of the variation was within rather than between sampling locations. No evidence of geographic structure was detected, nor were there any significant genetic differences between killifish populations. With these data, gene flow was evaluated by estimating effective migration rate (N_em) between the populations. N_em was estimated from G_st(N_em = 49.4), from F_st (N_em = 24.1), by the private alleles approach (N_em = 18.5) and via four phylogenetic analyses (N_em ranged from 11.4 to 16.9). Regardless of the analysis, N_em was greater than one; the threshold level at which gene flow is strong enough to prevent differentiation due to genetic drift. This suggests that while the characteristics that distinguish Fundulus subspecies may have arisen in isolation, the current clinal distributions exhibited along the Atlantic coast are most likely maintained by selection.     

Genetic differentiation across a latitudinal gradient in two co-occurring butterfly species: revealing population differences in a context of climate change

Genetic differentiation within a species' range is determined by natural selection, genetic drift, and gene flow. Selection and drift enhance genetic differences if populations are sufficiently isolated, while gene flow precludes differentiation and local adaptation. Over large geographical areas, these processes can create a variety of scenarios, ranging from admixture to a high degree of population differentiation. Genetic differences among populations may signal functional differences within a species' range, potentially leading to population or ecotype-specific responses to global change. We investigated differentiation within the geographical range of two butterfly species along a broad latitudinal gradient. This gradient is the primary axis of climatic variation, and many ecologists expect populations at the poleward edge of this gradient to expand under climate change. Our study species inhabit a shared ecosystem and differ in body size and resource specialization; both also find their poleward range limit on an island. We find evidence for divergence of peripheral populations from the core in both taxa, suggesting the potential for genetic distinctiveness at the leading edge of climate change. We also find differences between the species in the extent of peripheral differentiation with the smaller and more specialized species showing greater population divergence (microsatellites and mtDNA) and reduced gene flow (mtDNA). Finally, gene flow estimates in both species differed strongly between two marker types. These findings suggest caution in assuming that populations are invariant across latitude and thus will respond as a single ecotype to climatic change.     

Genetic diversity and historical population structure in the New Zealand mayfly Acanthophlebia cruentata

1. Nucleotide sequences of a 280 base pair region of the cytochrome b gene were used to assess genetic diversity and to infer population histories in the New Zealand mayfly Acanthophlebia cruentata. 2. A hierarchial examination of populations from 19 streams at different spatial scales in the central and northern North Island of New Zealand found 34 haplotypes. A common haplotype was found in all central region streams and unique haplotypes in northern streams. Several central streams had region specific haplotypes with genetically differentiated populations at the 70–100 km scale. 3. Haplotype diversity was high (0.53–0.8) at most sites, but low (0–0.22) in some central sites. amova analyses found significant genetic diversity among regions (69%) and among catchments (58%). Most population pairwise F_ST tests were significant, with non‐significant pairwise tests among sites in the central region and pairs of sites between neighbouring streams. 4. The levels of sequence divergence are interpreted as the result of Pleistocene divergence in multiple refugia, leading to the evolution of regionally unique haplotypes. The low diversity in some central region populations may result from recent colonisation following local extinctions, associated with volcanic events.     

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.     

Genetic differentiation in pikeperch (Sander lucioperca): the relative importance of gene flow, drift and common history

To investigate the relative importance of homogenizing factors, such as gene flow, and diversifying factors, such as drift, genetic variation in pikeperch ( Sander lucioperca ) in two Fennoscandian regions (North and South) was analysed with microsatellites. Allelic richness and the degree of differentiation were significantly higher in the North ( F _ST= 0·20) than in the South ( F _ST= 0·064). In northern areas, assignments of genotypes were almost exclusively to the population of origin, but in southern areas, the proportion of correct assignments was significantly lower. Most samples exhibited significant heterozygote deficits, and the level of relatedness was higher than expected from randomness. These combined results suggest that there has been more gene flow between populations in southern areas than in northern areas, where the importance of genetic drift has been greater. Effective population sizes were small ( c. 100) and did not differ between areas. The effect of a common history appears minor, and thus processes such as genetic drift and gene flow have been more influential in shaping the patterns of genetic diversity in this species.     

Passive rafting is a powerful driver of transoceanic gene flow

Dispersal by passive oceanic rafting is considered important for the assembly of biotic communities on islands. However, not much is known about levels of population genetic connectivity maintained by rafting over transoceanic distances. We assess the evolutionary impact of kelp-rafting by estimating population genetic differentiation in three kelp-associated invertebrate species across a system of islands isolated by oceanic gaps for over 5 million years, using mtDNA and AFLP markers. The species occur throughout New Zealand''s subantarctic islands, but lack pelagic stages and any opportunity for anthropogenic transportation, and hence must rely on passive rafting for long-distance dispersal. They all have been directly observed to survive transoceanic kelp-rafting journeys in this region. Our analyses indicate that regular gene flow occurs among populations of all three species between all of the islands, especially those on either side of the subtropical front oceanographic boundary. Notwithstanding its perceived sporadic nature, long-distance kelp-rafting appears to enable significant gene flow among island populations separated by hundreds of kilometres of open ocean.     

Population structure and gene flow of the Atlantic walrus (Odobenus rosmarus rosmarus) in the eastern Atlantic Arctic based on mitochondrial DNA and microsatellite variation

The population structure of the Atlantic walrus, Odobenus rosmarus rosmarus , was studied using 11 polymorphic microsatellites and restriction fragment length polymorphism detected in the NADH-dehydrogenase ND1, ND2 and ND3/4 segments in mtDNA. A total of 105 walrus samples were analysed from northwest (NW) Greenland, east (E) Greenland, Svalbard and Franz Joseph Land. Two of the 10 haplotypes detected in the four samples were diagnostic for the NW Greenland sample, which implied that the group of walruses in this area is evolutionary distinct from walruses in the other three areas. One individual sampled in E Greenland exhibited a Pacific haplotype, which proved a connection between the Pacific walrus and walruses in eastern Greenland. The Franz Joseph Land, Svalbard and E Greenland samples shared the most common haplotype, indicating very little differentiation at the mtDNA level. Gene flow ( Nm ) estimates among the four areas indicated a very restricted exchange of female genes between NW Greenland and the more eastern Atlantic Arctic samples, and a closer relationship between the three samples composing the eastern Atlantic Arctic. The genetic variation at 11 polymorphic microsatellite loci grouped individuals into three populations, NW Greenland, E Greenland and a common Franz Joseph Land–Svalbard population, which were connected by moderate gene flow.     

利用SSR分子标记分析秦岭冷杉自然居群的遗传多样性

利用10对SSR引物对濒危植物秦岭冷杉(Abies chensiensis)6个自然居群的120个个体进行了遗传多样性研究,旨在分析秦岭冷杉6个自然居群的遗传多样性、遗传结构及基因流变化.研究结果表明,120个个体在10个位点上共检测到149个等位基因,平均每个位点的等位基因数(A)为14.9,每个位点的有效等位基因数(e)为7.7,每个位点的平均预期杂合度(He)和平均观察杂合度(Ho)分别为0.841和0.243,Shannon多样性指数(Ⅰ)为2.13,自然居群杂合性基因多样度的比率(FsT)为6.7％,居群间的基因流(Nm)为3.45.利用Mantel检测到自然居群的遗传距离与地理距离间无显著相关性(r=0.490 6,P＞0.05).秦岭冷杉自然居群的遗传多样性水平较低,遗传变异主要存在于居群内部.结合研究数据、实地调查及相关资料,推测秦岭冷杉自然居群间基因流较原来增大可能是因为居群间发生了远交衰退.     

Empirical evaluation of cytonuclear models incorporating genetic drift and tests for neutrality of mtDNA variants: data from experimental Gambusia hybrid zones

Statistical tests of genetic drift and of the neutrality of mtDNA are presented using empirical time‐series data on multi‐generational changes in cytonuclear disequilibria within replicated experimental hybrid populations of two species of live‐bearing Poeciliid fishes (Gambusia holbrooki and G.affinis) which were monitored over a period of two years (three generations). Cytonuclear disequilibria D and D (which measure departures from random associations of cytoplasmic and nuclear genotypes) over the three generations of the experiment were non‐zero for all replicate populations. For each of five nuclear loci, the observed measures of D and D were highly concordant between replicates during each generation. Significant departures from expectations were observed after one and two generations. A statistical measure of goodness of fit of observed changes in cytonuclear disequilibria (and implicitly of the neutrality of the mtDNA markers) was calculated for each nuclear locus. When the results for the replicates were combined into an overall test of neutrality, the fit to the random union of zygotes (RUZ) model was rejected for four of the five nuclear loci (P < 0.05). A simple genetic drift model does not explain the temporal changes in composite cytonuclear genotypic frequencies. Frequencies of parental G. holbrooki mitochondrial alleles and nuclear genotypes exceeded expected values during most time periods, implying some selective advantage of offspring produced by G. holbrooki females. Expansion of cytonuclear models to explicitly address questions of genetic drift and neutrality have general relevance to studies of natural populations. This revised version was published online in July 2006 with corrections to the Cover Date.     

Short-term impact of disturbance on genetic diversity and structure of Indonesian populations of the butterfly Drupadia theda in East Kalimantan

We investigated the short‐term impact of disturbance on genetic diversity and structure of the tropical butterfly Drupadia theda Felder (Lepidoptera: Lycaenidae). Populations were sampled from five landscapes in East Kalimantan (Borneo, Indonesia) which were differentially disturbed by selective logging and the 1997/1998 El Niño Southern Oscillation (ENSO)‐induced drought and fires. Sampling occurred before (in 1997) and after the forest fires (in 1998, 1999, 2000, and 2004). Drupadia theda populations underwent serious population size reductions following the 1997/1998 ENSO event. For a total of 208 individuals, we sequenced a 509‐bp segment of mtDNA containing the control region plus the 5’ end of the 12S rDNA gene. Haplotype diversity in D. theda populations ranged from 0.468 to 0.953. Just after the 1997/1998 ENSO event, number of recorded individuals and genetic diversity were very low in D. theda populations sampled in the two severely burned areas and in a small pristine forest fragment that was surrounded by burned forest and thereby affected by drought. Interestingly, higher levels of genetic diversity were observed in logged forest compared to proximate pristine forest. After 1998, the genetic composition within the three ENSO‐disturbed areas diverged. In the twice‐burned forest, the genetic diversity in 1999 already approached pre‐fire levels, while it remained nearly unchanged in proximate once‐burned forest. Our data suggest that the 1997/1998 ENSO‐induced drought and fires caused massive reductions in the genetic diversity of D. theda and that population recoveries were linked to their geographical position relative to patches of unburned forest (and thus to source populations).     

Population Genetic Structure of the Yangtze Finless Porpoise (Neophocaena phocaenoides asiaeorientalis): Implications for Management and Conservation

Understanding the population genetic structure is a prerequisite for conservation of a species. The degree of genetic variability characteristic of the mitochondrial DNA control region has been widely exploited in studies of population genetic structure and can be useful in identifying meaningful population subdivisions. To estimate the genetic profile of the Yangtze finless porpoise (Neophocaena phocaenoides asiaeorientalis), an endangered freshwater population endemic to China, the complete mtDNA control region was examined in 39 individuals belonging to seven different stocks inhabiting the middle and lower reaches of the Yangtze River. Very low genetic diversity was found (nucleotide diversity 0.0011± 0.0002 and haplotypic diversity 0.65± 0.05). The mtDNA genetic pattern of the Yangtze population appears to indicate a founder event in its evolutionary history and to support the marine origin for this population. Analyses by F_st and Φ_st yielded statistically significant population genetic structure (F_st = 0.44, P < 0.05; Φ_st = 0.36, P < 0.05). These results may have significant implications for the management and conservation of the Yangtze finless porpoise in the future.     

Ecology and genetics of interspecific hybridization in the swallowtails, Papilio hospiton Gene and P. machaon L., in Corsica (Lepidoptera: Papiliomdae)

The Corsican swallowtail butterfly, Papilio hospiton , is endemic to Corsica and Sardinia (France and Italy) and included in the list of endangered species by the Washington Convention, It is spread all over Corsica in scattered populations linked to diverse habitats. A study by enzyme electrophoresis showed that the genetic diversity of the species is of the same order of magnitude as that of Papilio machaon from continental France and Corsica. The differentiation between populations is rather low, which is consistent with the high vagility of the adults. Natural hybridization between P. hospiton and P. machaon is frequent, and laboratory crosses show that the hybrids are not sterile. However, developmental perturbations impair the viability of further hybrid progenies. Although limited introgression between the two species is likely to take place, enzyme electrophoresis and PCR-RFLP analysis of mitochondrial DNA show that their gene pools remain distincfTtienetic assimilation by P. machaon therefore does not seem to be a threat for P. hospiton.