Genetic diversity in three species of Forsythia (Oleaceae) endemic to Korea: Implications for population history,taxonomy, and conservation

As the mountains in temperate regions have been hypothesized as refugia for boreal and temperate plants during the Last Glacial Maximum, one would expect high within-population genetic variation and low or moderate among-population differentiation for those species occurring there. We selected Forsythia ovata and Forsythia saxatilis, endemic to limestone mountains of Korea, to test the scenario that limestone ranges would also have acted as refugia. We undertook a population genetic analysis using 14 putative allozyme loci of the two species and the widely cultivated Forsythia koreana. We found relatively high levels of genetic diversity and low to moderate among-population differentiation in F. ovata and F. saxatilis, which are consistent with the proposed scenario. Allozyme data revealed that F. ovata and F. saxatilis are likely conspecific. As many populations of F. ovata and F. saxatilis have experienced dramatic decreases in their size, a comprehensive conservation program is urgently needed.     

Large effective population sizes and high levels of gene flow between subpopulations of Lilium cernuum (Liliaceae)

The Baekdudaegan, a mountain range that runs north to south along the Korean Peninsula, has been suggested to harbor important glacial refugia for boreal and temperate plant species. A series of allozyme-based genetic studies supports this trend. A large effective population size (N_e) is suggested as one of major factors contributing to maintaining moderate or high levels of within-population genetic variation in these plant species. To test this hypothesis, we examined the levels and patterns of allozyme diversity, tested recent bottlenecks, and estimated recent migration rates in 10 subpopulations (collected within a distance of ca. 640 m) of the boreal Lilium cernuum at Mt. Deokhang, in the central part of the Baekdudaegan. We found high levels of within-population genetic variation as well as a low between-population genetic differentiation (H_e = 0.206 and F_ST = 0.019). Based on the F_ST estimate and mean recent migration rate, we approximately calculated a total effective population size of 508 across 10 subpopulations. Consistent with this, we found no evidence of recent bottlenecks in the subpopulations. This study reveals that subpopulations of L. cernuum at Mt. Deokhang are effectively large (on the order of hundreds), and that high levels of gene flow occur among them, probably due to the species' high potential for seed dispersal. These demographic and life-history traits, coupled with its high levels of genetic diversity, suggests that this cold-adapted species would have found large refugial areas in these mountains (i.e., macrorefugia) during the Last Glacial Maximum.     

Comparison of genetic diversity in the two arctic–alpine plants Diapensia lapponica var. obovata (Diapensiaceae) and Empetrum nigrum var. japonicum (Empetraceae) between Sakhalin in Russian Far East and Jeju Island in Korea, the southernmost edge of their distribution range

We compared allozyme variation in the two arctic–alpine plants Diapensia lapponica var. obovata and Empetrum nigrum var. japonicum between Sakhalin Island in Russian Far East, within their range core, and the Korean island of Jeju, their world’s southernmost distribution. For D. lapponica var. obovata, Sakhalin populations harbored moderate levels of within-population genetic variation and low among-population divergence, whereas extremely low levels of within-population genetic diversity and high among-population differentiation were found in Jeju Island populations. In contrast, we found moderate levels of within-population variation and low among-population differentiation in E. nigrum var. japonicum in both northern populations (those of Sakhalin and an additional population from northern Japan) and Jeju Island populations. Under a similar scenario of immigration history of arctic–alpine plants on Jeju Island during the glacial periods of the Pleistocene and local persistence through glacial/interglacial cycles, the contrasting genetic structure between D. lapponica var. obovata and E. nigrum var. japonicum is mainly attributable to their different life-history, ecological, and demographic traits: (1) hermaphroditic versus monoecious, dioecious or polygamous, (2) seeds with no adaptations for long-distance dispersal versus berry-like drupes dispersed by animals and birds, and (3) a very small patch near the peak of Mt. Halla with a few hundred individuals versus a relatively continuous distribution around the peak of Mt. Halla with numerous individuals. From a conservation perspective, in situ and ex situ conservation measures should be strengthened for D. lapponica var. obovata on Jeju Island given their extreme rarity there.     

Molecular phylogeography and intraspecific divergence of Spiraea alpina (Rosaceae) distributed in the Qinghai-Tibetan Plateau and adjacent regions inferred from nrDNA

The Qinghai-Tibetan plateau (QTP) uplift had a decisive effect on climatic and eco-environmental evolution in East Asia during the Quaternary. In the current study phylogeographic structure and diversification history of Spiraea alpina across the QTP were investigated for the first time based on nuclear internal transcribed spacer. The nuclear internal transcribed spacers (ITS_1a–ITS₄) were generated for a total of 284 individuals distributed within 31 natural populations. A clear phylogeographic structure was found for S. alpina. The results showed that this species colonized in three different glacial refugia during the Quaternary extensive glaciation and expanded during the Interglacial period. Analysis of molecular variance (AMOVA) showed 74.13% genetic diversity among populations and 25.87% genetic variation within populations with distinct phylogeographic structure (F_ST = 0.741*). The estimated divergence time revealed that the main lineages of S. alpina diversified during the Quaternary 1.2–0.6 million years ago. The study concluded that severe climatic oscillations during Quaternary and the uplift of QTP had a profound effect on intraspecific divergence of S. alpina.     

Phylogeography of Rhodiola kirilowii (Crassulaceae): A Story of Miocene Divergence and Quaternary Expansion

The evolution and current distribution of the Sino-Tibetan flora have been greatly affected by historical geological events, such as the uplift of the Qinghai-Tibetan Plateau (QTP), and Quaternary climatic oscillations. Rhodiola kirilowii, a perennial herb with its distribution ranging from the southeastern QTP and the Hengduan Mountains (HM) to adjacent northern China and central Asia, provides an excellent model to examine and disentangle the effect of both geological orogeny and climatic oscillation on the evolutionary history of species with such distribution patterns. We here conducted a phylogeographic study using sequences of two chloroplast fragments (trnL-F and trnS-G) and internal transcribed spacers in 29 populations of R. kirilowii. A total of 25 plastid haplotypes and 12 ITS ribotypes were found. Molecular clock estimation revealed deep divergence between the central Asian populations and other populations from the HM and northern China; this split occurred ca. 2.84 million year ago. The majority of populations from the mountains of northern China were dominated by a single haplotype or ribotype, while populations of the HM harbored both high genetic diversity and high haplotype diversity. This distribution pattern indicates that HM was either a diversification center or a refugium for R. kirilowii during the Quaternary climatic oscillations. The present distribution of this species on mountains in northern China may have resulted from a rapid glacial population expansion from the HM. This expansion was confirmed by the mismatch distribution analysis and negative Tajima''s D and Fu''s F _S values, and was dated to ca. 168 thousand years ago. High genetic diversity and population differentiation in both plastid and ITS sequences were revealed; these imply restricted gene flow between populations. A distinct isolation-by-distance pattern was suggested by the Mantel test. Our results show that in old lineages, populations may harbour divergent genetic forms that are sufficient to maintain or even increase overall genetic diversity despite fragmentation and low within-population variation.     

Population genetics and phylogeography of freshwater mussels in North America, Elliptio dilatata and Actinonaias ligamentina (Bivalvia: Unionidae)

Extrinsic and intrinsic forces combined shape the population structure of every species differently. Freshwater mussels are obligate parasites to a host fish during a juvenile stage (glochidia). Elliptio dilatata (ED) and Actinonaias ligamentina (AL) are co-occurring freshwater mussel taxa with similar North American distribution and share some potential host fish. Using mitochondrial DNA, we determined the genotypes of 190 + individuals from collection sites in at least two tributaries in the Lake Erie and Ohio River watersheds, along with the Ouachita and Strawberry rivers in the southeast. Both species had followed a stepping-stone model of dispersal, with greater pairwise genetic structure among collection sites of ED. Also, phylogeographical analysis for ED found significant geographical structuring of haplotype diversity. Overall, within-population variation increased significantly from north to south, with low genetic diversity in the Strawberry River. We calculated significant among-population structure for both species (ED: Phi(ST) = 0.62, P < 0.001; AL: Phi(ST) = 0.16, P < 0.001). Genetic analysis identified the Ouachita River as an area of significant reproductive isolation for both species. Results for AL indicated dispersal into northern areas from two genetically distinct glacial refugia, where results for ED indicated dispersal followed by low gene flow in northern areas. The conservation strategies for mussels that co-occur in the same 'bed' could be species specific. Species such as ED have management units on the population scale, where AL has a more homogeneous genetic structure across its range.     

Genetic structure of Podocarpus nubigena (Podocarpaceae) provides evidence of Quaternary and ancient historical events

Evolution of ancient lineages inhabiting austral latitudes of the southern Andes has taken place in response to historical processes occurring at distinct spatial and temporal scales. We tested the hypothesis that distribution patterns of genetic polymorphisms within Podocarpus nubigena Lindl. (Podocarpaceae) were shaped by contemporary events acting at local scales such as Quaternary glaciations as well as pre-Quaternary palaeogeographical features of the landscape. We predict that such cold hardy species was able to survive locally in refugia during ice ages. As a result of local long-lasting persistence, its gene pool may also reflect the pre-Quaternary palaeogeography of southern South America. We collected fresh leaves from 30 randomly selected individuals at 14 populations. Leaf tissue was analyzed by isozyme electrophoresis using 12 putative loci most of which (92%) were polymorphic in at least one population. Elevated total genetic diversity was recorded H_T = 0.275 which was similarly high throughout populations with an overall significant mean among-population isolation (F_ST = 18%). Multivariate cluster and Bayesian analyses yielded significant latitudinal divergence at 43°S associated to a palaeobasin that produced a barrier for gene flow. Populations south of 43°S and those growing in lowland areas on Quaternary substrates of the North Patagonian Massif (Q NPM) present the highest genetic diversity suggesting long-lasting persistence. In contrast, northern mountain populations on pre-Quaternary substrates (pre-Q NPM) hold reduced genetic variation due to altitudinal movements in response to climate shifts. We suggest that palaeogeography of southern South America in addition to Neogene glaciations have shaped the genetic makeup of such ancient lineage as Podocarpus.     

Comparative biogeography of the congener lilies Lilium distichum and Lilium tsingtauense in Korea

The main Korean mountain range (the so-called “Baekdudaegan”), which stretches from north to south across most of the Peninsula, has been thought to harbor glacial refugia for boreal and temperate plant species, where they likely found relatively stable habitats and maintained large population sizes. Under this scenario, high levels of genetic variation for plant populations can be anticipated. To test this hypothesis, we examined levels of allozyme diversity in the boreal herb Lilium distichum, which in Korea occurs largely along the Baekdudaegan and in its closely related congener L. tsingtauense, a temperate species that can occur from the coastal islands to high-altitude mountains in the Baekdudaegan. As expected, L. distichum harbored high levels of genetic variation within populations, and we found significant correlations between percentage of polymorphic loci (and allelic richness) and elevation. For L. tsingtauense we found moderate genetic variation, with its populations showing a significantly positive correlation between intra-population expected heterozygosity and elevation. Our results suggest that populations/species that occur within or near the main ridge of the Baekdudaegan probably endured the glacial periods in macrorefugia, whereas those populations/species located at low elevations (far away from mid- and high-elevation mountains) did it in microrefugia. Palaeodistribution modelling is in agreement with genetic data, indicating that the Baekdudaegan was suitable habitat for both species during the glacial (LGM) and interglacial periods (like the present). This study as well as the data from other studies compiled here provides strong evidence that the Baekdudaegan was a major refugial area for the East Asian flora throughout the Quaternary and, thus, meriting high priority for conservation biogeography.     

Population genomic evidence for plant glacial survival in Scandinavia

Quaternary glaciations have played a major role in shaping the genetic diversity and distribution of plant species. Strong palaeoecological and genetic evidence supports a postglacial recolonization of most plant species to northern Europe from southern, eastern and even western glacial refugia. Although highly controversial, the existence of small in situ glacial refugia in northern Europe has recently gained molecular support. We used genomic analyses to examine the phylogeography of a species that is critical in this debate. Carex scirpoidea Michx subsp. scirpoidea is a dioecious, amphi‐Atlantic arctic–alpine sedge that is widely distributed in North America, but absent from most of Eurasia, apart from three extremely disjunct populations in Norway, all well within the limits of the Weichselian ice sheet. Range‐wide population sampling and variation at 5,307 single nucleotide polymorphisms show that the three Norwegian populations comprise unique evolutionary lineages divergent from Greenland with high between‐population divergence. The Norwegian populations have low within‐population genetic diversity consistent with having experienced genetic bottlenecks in glacial refugia, and host private alleles that probably accumulated in long‐term isolated populations. Demographic analyses support a single, pre‐Weichselian colonization into Norway from East Greenland, and subsequent divergence of the three populations in separate refugia. Other refugial areas are identified in North‐east Greenland, Minnesota/Michigan, Colorado and Alaska. Admixed populations in British Columbia and West Greenland indicate postglacial contact. Taken together, evidence from this study strongly indicates in situ glacial survival in Scandinavia.     

Molecular evidence for glacial expansion and interglacial retreat during Quaternary climatic changes in a montane temperate pine (Pinus kwangtungensis Chun ex Tsiang) in southern China

It has been suggested that the eastern Asian temperate flora have responded to Quaternary climatic changes in a different way compared with temperate plants in Europe and Northern America. However, knowledge about their phylogeographic structure and evolutionary history is still limited. In this study we investigated mitochondrial DNA variation in 17 populations of Pinus kwangtungensis, a five-needled pine inhabiting in isolated mountains of southern China and northern Vietnam. A total of ten mitochondrial haplotypes (mitotypes) were characterized by the polymorphisms of two fragments (cox-1-2 and nad7 intron1), and total genetic diversity was high (h _T = 0.847). The construction of phylogenetic relationships of the ten mitotypes detected three major, well-distinct clades, largely corresponding to four population groups identified by SAMOVA analysis. SAMOVA also indicated that most genetic variance should be attributed to among-group differentiation (F _CT = 0.868), consistent with the substantial genetic structure found within P. kwangtungensis (G _ST = 0.751). The genetic distances of P. kwangtungensis weakly but significantly correlated with geographical distances (R = 0.228, P = 0.03), revealing a pattern of isolation-by-distance. Demographic analysis did not detect any bottleneck events in the recent history of P. kwangtungensis. These results clearly suggested that there were three major refugia for this montane temperate conifer during warm stages (along with other minor refugia), and that the species would have tracked Quaternary climatic changes by expanding to nearby lowlands in the glacial periods, and by retreating to mountaintops (the refugia) during interglacial warmer stages, experiencing fragmentation and isolation among refugia.     

Vicariance and Its Impact on the Molecular Ecology of a Chinese Ranid Frog Species-Complex (Odorrana schmackeri,Ranidae)

Paleogeological events and Pleistocene climatic fluctuations have had profound influences on the genetic patterns and phylogeographic structure of species in southern China. In this study, we investigated the population genetic structure and Phylogeography of the Odorrana schmackeri species complex, mountain stream-dwelling odorous frogs, endemic to southern China. We obtained mitochondrial sequences (1,151bp) of the complete ND2 gene and two flanking tRNAs of 511 individuals from 25 sites for phylogeographic analyses. Phylogenetic reconstruction revealed seven divergent evolutionary lineages, with mean pairwise (K2P) sequence distances from 7.8% to 21.1%, except for a closer ND2 distance (3.4%). The complex geological history of southern China drove matrilineal divergence in the O. schmackeri species complex into highly structured geographical units. The first divergence between lineage A+B and other lineages (C-G) had likely been influenced by the uplift of coastal mountains of Southeast China during the Mio-Pliocene period. The subsequent divergences between the lineages C-G may have followed the formation of the Three Gorges and the intensification of the East Asian summer monsoon during the late Pliocene and early Pleistocene. Demographic analyses indicated that major lineages A and C have been experienced recent population expansion (c. 0.045–0.245 Ma) from multiple refugia prior to the Last Glacial Maximum (LGM). Molecular analysis suggest that these seven lineages may represent seven different species, three described species and four cryptic species and should at least be separated into seven management units corresponding to these seven geographic lineages for conservation.     

Refugia of Potentilla matsumurae (Rosaceae) located at high mountains in the Japanese archipelago

Molecular phylogeographic studies have revealed the genetic patterns and glacial-interglacial history of many plant and animal species. To infer the Quaternary history of alpine plants in the Japanese archipelago, which is poorly known, we investigated 203 individuals of Potentilla matsumurae and its varieties from 22 populations. We found 11 haplotypes based on approximately 1400 bp of two intergenetic spacers in chloroplast DNA (trnT-L and rpl20-rps20). The distribution of these haplotypes was geographically structured, which was supported by haplotype composition, principal component analysis, and unweighted pair group method with arithmetic mean (UPGMA), and N(ST) (0.71) was significantly greater than G(ST) (0.68). In addition to the positive correlation between genetic and geographic distance (Mantel test, r = 0.497, P < 0.001), an abrupt genetic change was detected between mountains in central Honshu and the Tohoku region. This genetic boundary was further supported by analysis of molecular variance (AMOVA), and high variation (54.0%) was explained by differences on either side of this boundary. Moreover, haplotypes in central Honshu were thought to have diverged, based on an outgroup comparison. These results suggest that mountains in central Honshu served as refugia during the Quaternary climatic oscillation, although the results could not reveal the history of most mountains in the Tohoku region and Hokkaido. Nevertheless, following floristic studies, our results indicate that alpine plants in Japan experienced a history different from that in Europe; i.e. they retreated into refugia during warm periods to avoid forest development, rather than glaciers.     

Population genetics of the hispid cotton rat (Sigmodon hispidus): patterns of genetic diversity at the major histocompatibility complex

The hispid cotton rat, Sigmodon hispidus, is a common rodent widely distributed across the southern USA and south into South America. To characterize major histocompatibility complex (MHC) diversity in this species and to elucidate large-scale patterns of genetic partitioning, we examined MHC genetic variability within and among 13 localities, including a disjunct population in Arizona and a population from Costa Rica that may represent an undescribed species. We also tested the hypothesis that populations within the USA are at equilibrium with regard to gene flow and genetic drift, resulting in isolation-by-distance. Using single-strand conformation polymorphism (SSCP) analysis we identified 25 alleles from 246 individuals. Gene diversity within populations ranged from 0.000 to 0.908. Analysis of molecular variance (AMOVA) revealed that 83.7% of observed variation was accounted for by within-population diversity and 16.3% was accounted for by among-population divergence. The disjunct population in Arizona was fixed for a single allele. The Costa Rican population was quite divergent based on allelic composition and was the only population with unique alleles. Within the main portion of the geographical distribution of S. hispidus in the USA there was considerable divergence among some populations; however, there was no significant pattern of isolation-by-distance overall (P = 0.090). Based on the significant divergence of the only sampled population to its east, the Mississippi River appears to represent a substantial barrier to gene flow.     

Phylogeography of the Chinese endemic freshwater crab Sinopotamon acutum (Brachyura,Potamidae)

Phylogeographical inferences based on multiple molecular markers from different modes of inheritance (i.e. mtDNA and nucDNA) can help to explore drivers of current species divergence over different timescales and allow a deeper understanding of evolutionary processes. River systems and mountains, owing to their drainage networks and complex topography, are often a high priority when inferring external causes of phylogeographical patterns. Here, we selected the Chinese endemic freshwater crab Sinopotamon acutum inhabiting drainage networks across the Qinling–Daba Mountains, a watershed that divides northern from southern fauna in China, as a model species (1) to investigate whether river networks and mountain systems act as barriers in shaping the phylogeography of freshwater crabs and (2) to test the impact of historic environments and ecological habitats on crab distribution. Phylogenetic tree and network analyses based on mtDNA revealed divergence between eastern and western populations, but microsatellite DNA did not recover this. Frequent and male‐biased gene flow is evident. The adjacent tributaries, such as Weihe River 1 and Hanjiang River 1, although isolated by Qinling Mts, have low genetic variation in mtDNA. In addition, microsatellite DNA showed low variation between all rivers. Frequent gene flow, genetic variation and mitochondrial genetic structure in S. acutum indicated that the Qinling–Daba Mountains have had no effect on divergence in this species; instead, hydrological networks across these mountains serve as a continuous migration corridor for these crabs. Glacial refugia are probably responsible for the initial isolation of the eastern and western clades, and local adaptations after dispersal have further driven divergence. Cytonuclear discordance in phylogeographical patterns may be attributable to ongoing male‐biased gene flow.     

Phylogeographical patterns of two closely related desert shrubs,Nitraria roborowskii and N. sphaerocarpa (Nitrariaceae), from arid north‐western China

To investigate the influence of climate aridification and oscillations on the genetic diversity and evolutionary processes of organisms in the Quaternary in north‐western China, we selected Nitraria roborowskii and N. sphaerocarpa and examined the geographical apportionment of genetic variation in their vast range. In the study, 21 plastid haplotypes were identified in N. roborowskii based on two plastid DNA regions. We found significant genetic differentiation between populations in both N. roborowskii and N. sphaerocarpa. We also found similar phylogeographical patterns of allopatric divergence and regional range expansion in both species, but the degree of allopatric divergence in N. roborowskii was lower. Finally, areas to the south of the Alxa Desert, the Hexi Corridor and Ningxia Province were identified as possible refugia for N. roborowskii.     

横断山区高山绣线菊的谱系地理学研究

横断山区作为青藏高原东南部主要的一个冰期避难所,第四纪冰期气候的变化对该地区的植物地理分布和居群遗传结构都产生了重要的影响。为了揭示该地区物种分布的分子系统地理学结构,选取在该地区广泛分布的一种高山灌木-高山绣线菊的叶绿体trnL-trnF序列进行研究。采集了15个居群182个个体进行测序,共发现7个单倍型。总的遗传多样性较高(HT=0.809),但居群内遗传多样性较低(HS=0.236)。分子变异分析(AMOVA)结果表明分布区内高山绣线菊的遗传变异主要存在于居群间(84.48%),且居群间的遗传分化很高(GST=0.708,FST=0.84476,NST=0.863),有着显著的谱系地理学结构(NST>GST,P<0.01)和较低的居群间平均基因流(Nm=0.09)。单倍型的系统进化树和进化分支网络分析得到了相似的拓扑结构,7种单倍型都按照地理分布聚为三支:横断山区西部、横断山区东部以及两者的交接地带。本研究推测该物种在横断山区存在多个冰期避难所,而没有表现出大规模的种群集体扩张和迁移的现象。青藏高原隆升、第四纪气候的反复波动以及横断山区特殊的地理环境使得原来连续的居群片段化,并发生范围扩张,从而塑造了高山绣线菊的现代生物地理分布格局。     

Population divergence and morphometric integration in the greenfinch (Carduelis chloris) – evolution against the trajectory of least resistance?

Theory predicts that genetic and phenotypic correlations among traits may direct the process of short-term evolution by limiting the directions of variation available to natural selection to act on. We studied correlations between 14 skeletal traits in 10 geographically distinct and relatively young greenfinch (Carduelis chloris) populations to unravel whether the divergence among populations has occurred into directions predicted by the within-population correlations (cf. drift/correlated responses models), or whether it is better explained by ‘adaptive’ models, which predict no necessary association between within- and among-population correlations (allometries). We found that the within-population character correlations (or covariances) did not predict character divergence between populations. This was because the first eigenvector of the among-population correlation/covariance matrix, summarizing the major dimension of divergence, was a bipolar body:beak dimension, and distinct from the (≈ isometric) first eigenvector of within-population matrix. Hence, as the divergence among greenfinch populations cannot be satisfactorily accommodated by drift/correlated response models, an adaptive basis for divergence is suggested. The second major axis of within-population variation was a classical ‘group size’ factor revealing that beak size was more or less free to vary independently of body size. Consequently, even if the divergence among populations cannot be simply accommodated to expectations of drift and correlated response models, it is striking that the most pronounced size-independent (nonallometric) changes had occurred along the second largest dimension of variance. This could mean that selection pressures which shape integration within populations are the same as those that cause divergence among populations. A relaxed beak:body integration could also occur as a result of species level selection favouring taxa in which independent evolution of beak and body is made possible.     

Genetic Variation in the Endangered Endemic Species Cycas fairylakea (Cycadaceae) in China and Implications for Conservation

Cycas fairylakea is an endangered endemic species in China. Genetic diversity within and among four natural populations of this species in China was investigated using amplified fragments length polymorphism (AFLP). A moderate to low level of intraspecific genetic diversity was detected in this species (at population level: P = 39.57 %, H₀ = 0.244; at species level: P = 60.22%, H₀ = 0.356). The among-population component accounted for, respectively, 25.7 and 31.5% of the genetic variation, according to AMOVA and Shannon’s index, indicating most of the genetic variation was found between individuals within populations. All four populations have opposite pyramid age structure, and few coning individuals, which is still decreasing. Possibly because of habitat degradation and environmental pollution, plant diseases and insect pests in the populations were extremely serious, suggesting that the main factors threatening the survival of C. fairylakea populations were not genetic variation, but human activities and the breeding system of this species.     

The evolution of sexual dimorphism in the house finch. I. Population divergence in morphological covariance structure

Abstract Patterns of genetic variation and covariation strongly affect the rate and direction of evolutionary change by limiting the amount and form of genetic variation available to natural selection. We studied evolution of morphological variance-covariance structure among seven populations of house finches (Carpodacus mexicanus) with a known phylogenetic history. We examined the relationship between within- and among-population covariance structure and, in particular, tested the concordance between hierarchical changes in morphological variance-covariance structure and phylogenetic history of this species. We found that among-population morphological divergence in either males or females did not follow the within-population covariance patterns. Hierarchical patterns of similarity in morphological covariance matrices were not congruent with a priori defined historical pattern of population divergence. Both of these results point to the lack of proportionality in morphological covariance structure of finch populations, suggesting that random drift alone is unlikely to account for observed divergence. Furthermore, drift alone cannot explain the sex differences in within- and among-population covariance patterns or sex-specific patterns of evolution of covariance structure. Our results suggest that extensive among-population variation in sexual dimorphism in morphological covariance structure was produced by population differences in local selection pressures acting on each sex.     

Genetic variation and population structure ofCarex breviculmis (Cyperaceae) in Korea

We determined the genetic diversity and population structures ofCarex breviculmis (Cyperaceae) populations in Korea, using genetic variations at 23 allozyme loci.C. breviculmis is a long-lived herbaceous species that is widely distributed in eastern Asia. A high level of genetic variation was found in 15 populations. Twelve enzymes revealed 23 loci, of which 11 were polymorphic (47.8%). Genetic diversity at the speciesand population levels were 0.174 and 0.146, respectively. Total genetic diversity (H_T = 0.363) and within-population genetic diversity (H_s = 0.346) were high, whereas the extent of the population divergence was relatively low (G_ST = 0.063). Deviation from random mating (Fis) within the 15 populations was 0.206. An indirect estimate of the number of migrants per generation(Nm = 3.69) indicated that gene flow was extensive among Korean populations of this species. Analysis of fixation indices revealed a substantial heterozygote deficiency in some populations and at some loci. Genetic identity between popu-lations was high, exceeding 0.956.