中国特有种四合木种群遗传多样性的RAPD分析

采用随机扩增多态性 DNA(RAPD)技术检测了西鄂尔多斯高原特有种四合木种群 5个斑块的遗传多样性。2 2个扩增引物产生 1 1 6条带 ,Shannon信息指数和 Nei指数对 RAPD数据的分析表明 :四合木种群存在较高的遗传多样性 ,其中千里山斑块的遗传多样性和多态位点比例较高 ,石嘴山斑块的最低。遗传多样性的 86.5 %存在于斑块内 ,斑块间的遗传变异为 1 3 .5 %。遗传距离与地理距离无直接相关关系。这些结果说明 ,遗传多样性反映了四合木种群基因组 DNA存在较高的变异性 ;同时各斑块间存在一定的基因流 ,四合木各斑块可看成是处于同一种群的半隔离状态 (meta-种群 ) ,对四合木应注意保护遗传多样性丰富的 meta-种群。     

蒙古沙葱萤叶甲种群遗传多样性的微卫星分析

【目的】沙葱萤叶甲 Galeruca daurica (Joannis)是一种近年来在内蒙古草原上猖獗成灾的新害虫,本研究旨在明确内蒙古沙葱萤叶甲不同地理种群间的遗传分化和基因交流程度。【方法】应用5对微卫星引物分析了沙葱萤叶甲8个地理种群的遗传多样性、基因流和遗传分化。【结果】5个位点等位基因数为10~18,有效等位基因数为9.2796~16.0388,多态信息含量值为0.6760~0.8985,期望杂合度为0.3430~0.5284,说明所选5个微卫星位点均为高度多态性位点。8个种群的期望杂合度为0.2216~0.3701,平均值为0.2680;种群间遗传分化系数为0.1244~0.4116,平均值为0.2521;种群间基因流为0.3574~1.7596,平均值为0.9622。8个地理种群根据遗传距离聚为3个分支,遗传距离与地理距离呈显著的正相关关系 (r=0.4854,P=0.0180)。【结论】沙葱萤叶甲种群遗传多样性低,不同地理种群之间基因流较小,遗传分化程度高;沙葱萤叶甲迁移能力弱和地理阻碍可能是限制其基因交流和导致遗传高度分化的主要原因。     

基于AFLP方法对不同地理种群棉褐带卷蛾遗传分化的研究

利用3对AFLP引物对我国棉褐带卷蛾Adoxophyes orana(Fischer von Rosslerstamm)4个地理种群进行了遗传分化研究。共扩增得到224个DNA条带,其中多态条带195个;遗传多样性分析表明各群体均具有较高的遗传多样性水平,Nei遗传多样性指数为0.1425⁰.1578,Shannon多样性指数为0.21370.1578,Shannon多样性指数为0.2137⁰.2383。群体间的遗传分化系数(Gst)和基因流(Nm)分析结果表明,北方的苹小卷叶蛾与南方茶小卷叶蛾之间产生了明显的遗传分化,而北方的苹小卷叶蛾不同种群间则存在着大量的基因交流,遗传分化程度很弱。AMOVA分析结果显示66.81%的变异来源于种群内。UMAGA聚类分析表明,北京种群、陕西种群和山东种群间遗传距离较小,聚在一起为一分支,福建的茶小卷蛾单独为一分支。推测棉褐带卷蛾种群间的遗传分化与寄主有关联,与地理距离不相关。     

中国特有种四合木种群遗传多样性的ＲＡＰＤ

采用随机扩增多态性ＤＮＡ（ＲＡＰＤ）技术检测了西鄂尔多斯高原特有种四合木种群５个斑块的遗传多样性。２２个扩增引物产生ｌｌ６条带,Ｓｈａｎｎｏｎ信息指数和Ｎｅｉ指数对ＲＡＰＤ数据的分析表明：四合木种群存在较高的遗传多样氓,其中千里山斑块的遗传多样性和多态位点比例较高,石嘴山斑块的最低。遗传多样性的８６．５％存在于斑块内,斑块间的遗传变异为１３．５％。遗传距离与地理距离无直接相关关系。这些结果说明,遗传多样性反映了四合木种群基因组ＤＮＡ存在较高的变异性;同时各斑块间存在一定的基因流,四合木各斑块可看成是处于同一种群的半隔离状态（ｍｅｔａ—种群）,对四合木应注意保护遗传多样性丰富的ｍｅｔａ—种群。     

基于COI基因全长序列的假眼小绿叶蝉地理种群遗传分化研究

假眼小绿叶蝉Empoasca vitis(Gthe)是我国大陆茶区分布最广、为害最重的茶树害虫,其种群世代重叠严重,数量大。从我国13个主要产茶省份各选出一个重点产茶县(市),采集假眼小绿叶蝉标本,首次扩增得到其线粒体CO I基因全长序列,并以此探讨了假眼小绿叶蝉13个地理种群间的遗传多样性、分子变异、遗传分化程度及基因流水平。在13个地理种群中共得到了176条CO I基因序列,发现了113个变异位点,形成了105个CO I单倍型。总群体单倍型多样性Hd为0.9720,种群内单倍型多样性在0.804—1.000范围内,总群体和各种群的Tajima's D检验结果均不显著,说明茶园假眼小绿叶蝉的进化符合中性模型,种群数量较为稳定。AMOVA分子变异分析结果表明,茶园假眼小绿叶蝉的遗传分化主要来自于种群内部。Mantel检验显示各地理种群的遗传距离与地理距离之间不具有显著的相关性。总群体的遗传分化系数Gst为0.03652,固定系数Fst为0.10876,基因流Nm为4.097,表明地理种群间存在较频繁的基因交流,遗传差异较小。推测,20世纪90年代以来的省际之间频繁的茶树鲜叶贩运、异地茶苗的大批量调拨等因素促进了假眼小绿叶蝉的长距离迁移,加强了不同种群间的基因交流。     

内蒙古小花棘豆遗传多样性的ISSR分析

应用ISSR分子标记对内蒙古西部地区小花棘豆（Oxytropis glabra DC.）6个地理种群进行了遗传多样性研究.结果表明：小花棘豆具有较高的遗传多样性;14个引物共扩增出327个位点,物种水平上Nei＇s基因多样性指数为0.1972,Shannon多样性指数为0.3287;种内总基因多样性为0.1976,种群内基因多样性为0.1653,83.67%的遗传变异存在于种群内,16.33%的遗传变异存在于种群间,种群间的遗传分化系数为0.1633,基因流为2.5613,6个种群间有一定遗传分化,但遗传漂变不会引起遗传分化.UPGMA遗传距离聚类结果表明,生态地理条件相似的种群优先聚集.     

内蒙古中部小驼嗡蜣螂七个种群的遗传多样性

采用RAPD分子标记技术分析了内蒙古中部地区小驼嗡蜣螂（Onthophagus gibbulus）7个种群的遗传多样性及遗传分化。16条引物共检测到363条带,总多态位点比率为100％,Nei基因多样度为0．280,总的Shannon信息指数为0．433,结果表明,小驼嗡蜣螂种群遗传多样性丰富。在总的遗传变异中,12．57％的变异存在于种群间,87．43％的变异存在于种群内,种群间的遗传分化程度较低,种群间遗传分化系数（Gst）在0．074～0．203,基因流（Nm）在4．569～9．791,表明种群间存在广泛的基因交流。在本研究涉及的地理范围内,小驼嗡蜣螂种群的遗传距离与地理距离不相关,说明小驼嗡蜣螂种群间的分化与其生境的异质性有关。     

高原鼠兔四个地理种群的遗传多样性与遗传分化

采用7 个多态微卫星DNA 标记分析了4 个高原鼠兔地理种群(海晏种群、西大滩东种群、西大滩西种群和昆仑山种群)的遗传多样性和遗传分化。采用多态信息含量(PIC)、期望杂合度(H_e )、观测杂合度(H_o)、基因流(N_m )和基因分化系数(F_st )对种群及物种的遗传多样性水平进行分析;计算各种群间的遗传距离(GD)、遗传相似度(GI)以及遗传距离与地理距离的相关系数,并进行UPGMA 聚类分析。研究结果显示,高原鼠兔的遗传多样性在小哺乳动物中处于较高水平, 其H_e 和H_o 的平均值分别为0. 5332 和0. 6003, 其F_st 为0. 0975,即有9. 75% 的变异存在于种群之间,即变异主要存在于种群内部。4 个种群间的杂合度与多态信息含量均无显著差异,说明它们的遗传多样性水平基本相当。海晏种群与其余3 个种群的遗传分化均达到了中度分化水平(F_st = 0.1043),而另3 个种群间几乎没有分化(F_st = 0. 0253)。Mantel test 结果表明地理距离与遗传距离间没有显著的相关性。     

西南地区麻疯树天然种群遗传多样性的等位酶变异

为了揭示麻疯树(Jatropha curcas)天然种群的遗传多样性和遗传结构, 采用聚苯烯酰胺凝胶垂直平板电泳技术, 对采自四川、云南、贵州3个省的10个麻疯树天然种群的叶片样本进行了同工酶分析。7个酶系统10个位点的检测结果表明: 麻疯树种群水平上的遗传多样性较高, 每位点平均等位基因数为2.428 6, 多态位点百分率为97.14%, 平均期望杂合度为0.396 4。种群间遗传分化系数为0.041 3, 种群间总的基因流较高, 为5.808 9, 种群间遗传一致度较高(Shannon信息指数为0.921 7- 0.995 3)。非加权类平均法(UPGMA)聚类结果显示, 10个种群的遗传距离与地理距离相关性不显著。麻疯树天然种群具有较低程度的遗传分化、较高的基因流, 种内及种群内多样性丰富, 这为麻疯树优良品种的选育提供了良好的遗传基础。     

中国南方茶棍蓟马地理种群遗传分化分析

【目的】基于mtDNA COI和COⅡ的联合序列,探讨中国南方茶棍蓟马Dendrothrips minowai地理种群的遗传多样性与遗传分化。【方法】利用MEGA 6.0, DnaSP 5.10和Arlequin 3.5.2.2等软件对中国南方茶棍蓟马8个地理种群遗传多样性、遗传分化、基因流、分子变异及地理距离与遗传距离的相关性进行分析,并推断群体演化历史。【结果】茶棍蓟马mtDNA COI和COⅡ序列均具有明显的AT偏好性,COI和COⅡ联合序列的长度为1 146 bp,共有22个单倍型。中国南方茶棍蓟马总群体遗传多样性较高,表现出高单倍型多样性(Hd=0.924)和低核苷酸多样性(π=0.00600)。8个地理种群总群体的遗传分化程度高(F_ST=0.84830),基因交流水平低(Nm=0.040),群体可能由于遗传漂变而发生明显分化。AMOVA分析显示,茶棍蓟马种群的遗传变异主要来自组间种群(F_CT=0.84922);Mantel检测显示地理距离与遗传距离存在显著的正相关(r=0.5029, P<0.01)。中性检验结果表明,除云南两个地理种群外的其他地理种群近期可能经历了种群扩张。【结论】中国南方茶棍蓟马地理种群遗传多样性较高,具有明显的遗传分化,基因交流较少;地理距离可能是影响茶棍蓟马地理种群遗传分化的主要因素之一。     

Effects of clonality on the genetic variability of rare,insular species: the case of Ruta microcarpa from the Canary Islands

Many plant species combine sexual and clonal reproduction. Clonal propagation has ecological costs mainly related to inbreeding depression and pollen discounting; at the same time, species able to reproduce clonally have ecological and evolutionary advantages being able to persist when conditions are not favorable for sexual reproduction. The presence of clonality has profound consequences on the genetic structure of populations, especially when it represents the predominant reproductive strategy in a population. Theoretical studies suggest that high rate of clonal propagation should increase the effective number of alleles and heterozygosity in a population, while an opposite effect is expected on genetic differentiation among populations and on genotypic diversity. In this study, we ask how clonal propagation affects the genetic diversity of rare insular species, which are often characterized by low levels of genetic diversity, hence at risk of extinction. We used eight polymorphic microsatellite markers to study the genetic structure of the critically endangered insular endemic Ruta microcarpa. We found that clonality appears to positively affect the genetic diversity of R. microcarpa by increasing allelic diversity, polymorphism, and heterozygosity. Moreover, clonal propagation seems to be a more successful reproductive strategy in small, isolated population subjected to environmental stress. Our results suggest that clonal propagation may benefit rare species. However, the advantage of clonal growth may be only short‐lived for prolonged clonal growth could ultimately lead to monoclonal populations. Some degree of sexual reproduction may be needed in a predominantly clonal species to ensure long‐term viability.     

Microsatellite and major histocompatibility complex variation in an endangered rattlesnake,the Eastern Massasauga (Sistrurus catenatus)

Genetic diversity is fundamental to maintaining the long‐term viability of populations, yet reduced genetic variation is often associated with small, isolated populations. To examine the relationship between demography and genetic variation, variation at hypervariable loci (e.g., microsatellite DNA loci) is often measured. However, these loci are selectively neutral (or near neutral) and may not accurately reflect genomewide variation. Variation at functional trait loci, such as the major histocompatibility complex (MHC), can provide a better assessment of adaptive genetic variation in fragmented populations. We compared patterns of microsatellite and MHC variation across three Eastern Massasauga (Sistrurus catenatus) populations representing a gradient of demographic histories to assess the relative roles of natural selection and genetic drift. Using 454 deep amplicon sequencing, we identified 24 putatively functional MHC IIB exon 2 alleles belonging to a minimum of six loci. Analysis of synonymous and nonsynonymous substitution rates provided evidence of historical positive selection at the nucleotide level, and Tajima's D provided support for balancing selection in each population. As predicted, estimates of microsatellite allelic richness, observed, heterozygosity, and expected heterozygosity varied among populations in a pattern qualitatively consistent with demographic history and abundance. While MHC allelic richness at the population and individual levels revealed similar trends, MHC nucleotide diversity was unexpectedly high in the smallest population. Overall, these results suggest that genetic variation in the Eastern Massasauga populations in Illinois has been shaped by multiple evolutionary mechanisms. Thus, conservation efforts should consider both neutral and functional genetic variation when managing captive and wild Eastern Massasauga populations.     

Polyploidy and microsatellite variation in the relict tree Prunus lusitanica L.: how effective are refugia in preserving genotypic diversity of clonal taxa?

Refugia are expected to preserve genetic variation of relict taxa, especially in polyploids, because high gene dosages could prevent genetic erosion in small isolated populations. However, other attributes linked to polyploidy, such as asexual reproduction, may strongly limit the levels of genetic variability in relict populations. Here, ploidy levels and patterns of genetic variation at nuclear microsatellite loci were analysed in Prunus lusitanica, a polyploid species with clonal reproduction that is considered a paradigmatic example of a Tertiary relict. Sampling in this study considered a total of 20 populations of three subspecies: mainland lusitanica (Iberian Peninsula and Morocco), and island azorica (Azores) and hixa (Canary Islands and Madeira). Flow cytometry results supported an octoploid genome for lusitanica and hixa, whereas a 16‐ploid level was inferred for azorica. Fixed heterozygosity of a few allele variants at most microsatellite loci resulted in levels of allelic diversity much lower than those expected for a high‐order polyploid. Islands as a whole did not contain higher levels of genetic variation (allelic or genotypic) than mainland refuges, but island populations displayed more private alleles and higher genotypic diversity in old volcanic areas. Patterns of microsatellite variation were compatible with the occurrence of clonal individuals in all but two island populations, and the incidence of clonality within populations negatively correlated with the estimated timing of colonization. Our results also suggest that gene flow has been very rare among populations, and thus population growth following founder events was apparently mediated by clonality rather than seed recruitment, especially in mainland areas. This study extends to clonal taxa the idea of oceanic islands as important refugia for biodiversity, since the conditions for generation and maintenance of clonal diversity (i.e. occasional events of sexual reproduction, mutation and/or seed immigration) appear to have been more frequent in these enclaves than in mainland areas.     

Genetic variation and clonal structure of the rare,riparian shrub <Emphasis Type="Italic">Spiraea virginiana</Emphasis> (Rosaceae)

Genetic diversity is often considered important for species that inhabit highly disturbed environments to allow for adaptation. Many variables affect levels of genetic variation; however, the two most influential variables are population size and type of reproduction. When analyzed separately, both small population size and asexual reproduction can lead to reductions in genetic variation, although the exact nature of which can be contrasting. Genetic variables such as allelic richness, heterozygosity, inbreeding coefficient, and population differentiation have opposite predictions depending upon the trait (rarity or clonality) examined. The goal of this study was to quantify genetic variation and population differentiation in a species that resides in a highly stochastic environment and is both rare and highly clonal, Spiraea virginiana, and to determine if one trait is more influential genetically than the other. From populations sampled throughout the natural range of S. virginiana, we used microsatellite loci to estimate overall genetic variation. We also calculated clonal structure within populations, which included genotypic richness, evenness, and diversity. Gene flow was investigated by quantifying the relationship between genetic and geographic distances, and population differentiation (θ) among populations. Observed heterozygosity, genotypic richness, and inbreeding coefficients were found to be representative of high clonal reproduction (averaging 0.505, 0.1, and –0.356, respectively) and the number of alleles within populations was low (range = 2.0–3.6), being more indicative of rarity. Population differentiation (θ) among populations was high (average = 0.302) and there was no relationship between genetic and geographic distances. By examining a species that exhibits two traits that both can lead to reduced genetic variation, we may find an enhanced urgency for conservation. Accurate demographic counts of clonal species are rarely, if ever, possible and genetic exploration for every species is not feasible. Therefore, the conclusions in this study can be potentially extrapolated to other riparian, clonal shrubs that share similar biology as S. virginiana.     

Geographic heterogeneity in natural selection on an MHC locus in sockeye salmon

Balancing selection maintains high levels of polymorphism and heterozygosity in genes of the MHC (major histocompatibility complex) of vertebrate organisms, and promotes long evolutionary persistence of individual alleles and strongly differentiated allelic lineages. In this study, genetic variation at the MHC class II DAB-beta1 locus was examined in 31 populations of sockeye salmon (Oncorhynchus nerka) inhabiting the Fraser River drainage of British Columbia, Canada. Twenty-five percent of variation at the locus was partitioned among sockeye populations, as compared with 5% at neutral genetic markers. Geographic heterogeneity of balancing selection was detected among four regions in the Fraser River drainage and among lake systems within regions. High levels of beta1 allelic diversity and heterozygosity, as well as distributions of alleles and allelic lineages that were more even than expected for a neutral locus, indicated the presence of balancing selection in populations throughout much of the interior Fraser drainage. However, proximate populations in the upper Fraser region, and four of six populations from the lower Fraser drainage, exhibited much lower levels of genetic diversity and had beta1 allele frequency distributions in conformance with those expected for a neutral locus, or a locus under directional selection. Pair-wise FST values for beta1 averaged 0.19 and tended to exceed the corresponding values estimated for neutral loci at all levels of population structure, although they were lower among populations experiencing balancing selection than among other populations. The apparent heterogeneity in selection resulted in strong genetic differentiation between geographically proximate populations with and without detectable levels of balancing selection, in stark contrast to observations at neutral loci. The strong partitioning and complex structure of beta1 diversity within and among sockeye populations on a small geographic scale illustrates the value of incorporating adaptive variation into conservation planning for the species.     

Genetic diversity and hierarchical population structure of a rare autotetraploid plant,Aster kantoensis (Asteraceae)

We sampled 17 populations of a rare autotetraploid Aster kantoensis (Asteraceae) from three river systems located in central Japan, and studied them for allelic variation at 22 enzyme loci. There was no significant correlation between the actual population size and three genetic diversity parameters, suggesting that the effective population size was very small even for the large populations, i.e., even large populations may still have a high probability of being of recent origin and remain influenced by the founder effect. Compared to other autotetraploid species, the total genetic variation of A. kantoensis is small. The number of alleles and gene diversity of a population were not significantly different among the river systems, although the percentage of polymorphic loci was different. Genetic differentiation among river systems was larger than between populations within the river systems, thereby indicating that gene flow between river systems is small, especially between the Kinu River system and Tama or Sagami River systems.     

Genetic diversity and population structure of the serpentine endemic Calystegia collina (Convolvulaceae) in northern California

We used enzyme electrophoresis to evaluate genetic diversity in 32 populations of Calystegia collina, a clonal plant species endemic to serpentine outcrops in northern California (USA). Of 34 loci examined 56% were polymorphic, but on average only 17% were polymorphic within local populations. Neither the total number of alleles nor the number of multilocus genotypes differed significantly between populations in small vs. large serpentine outcrops. Genetic and geographic distances between populations were positively correlated, but this relationship was not significantly affected by the isolation of serpentine outcrops. Populations were highly differentiated (F(st) = 0.417) and little genetic variation was explained by geographic region or serpentine outcrop.Observed heterozygosity within populations almost always exceeded Hardy-Weinberg expectations. In many populations, all 30 sample ramets were uniformly heterozygous at one or more loci yet were genetically variable at other loci. These results imply that many C. collina populations originate from one or a few genetic founders, with little recruitment from seeds. Genetic variation within uniformly heterozygous populations must be the product of multiple, closely related founders or somatic mutations within the population. We conclude that vegetative reproduction, perhaps coupled with somatic mutation, helps maintain genetic diversity in these isolated but long-lived populations.     

Genetic diversity of Vietnamese domestic chicken populations as decision‐making support for conservation strategies

The aims of this study were to assess the genetic diversity of 17 populations of Vietnamese local chickens (VNN) and one Red Jungle Fowl population, together with six chicken populations of Chinese origin (CNO), and to provide priorities supporting the conservation of genetic resources using 20 microsatellites. Consequently, the VNN populations exhibited a higher diversity than did CNO populations in terms of number of alleles but showed a slightly lower observed heterozygosity. The VNN populations showed in total seven private alleles, whereas no CNO private alleles were found. The expected heterozygosity of 0.576 in the VNN populations was higher than the observed heterozygosity of 0.490, leading to heterozygote deficiency within populations. This issue could be partly explained by the Wahlund effect due to fragmentation of several populations between chicken flocks. Molecular analysis of variance showed that most of genetic variation was found within VNN populations. The Bayesian clustering analysis showed that VNN and CNO chickens were separated into two distinct groups with little evidence for gene flow between them. Among the 24 populations, 13 were successfully assigned to their own cluster, whereas the structuring was not clear for the remaining 11 chicken populations. The contributions of 24 populations to the total genetic diversity were mostly consistent across two approaches, taking into account the within‐ and between‐populations genetic diversity and allelic richness. The black H'mong, Lien Minh, Luong Phuong and Red Jungle Fowl were ranked with the highest priorities for conservation according to Caballero and Toro's and Petit's approaches. In conclusion, a national strategy needs to be set up for Vietnamese chicken populations, with three main components: conservation of high‐priority breeds, within‐breed management with animal exchanges between flocks to avoid Wahlund effect and monitoring of inbreeding rate.     

Population demography and heterozygosity–fitness correlations in natural guppy populations: An examination using sexually selected fitness traits

Heterozygosity–fitness correlations (HFCs) have been examined in a wide diversity of contexts, and the results are often used to infer the role of inbreeding in natural populations. Although population demography, reflected in population‐level genetic parameters such as allelic diversity or identity disequilibrium, is expected to play a role in the emergence and detectability of HFCs, direct comparisons of variation in HFCs across many populations of the same species, with different genetic histories, are rare. Here, we examined the relationship between individual microsatellite heterozygosity and a range of sexually selected traits in 660 male guppies from 22 natural populations in Trinidad. Similar to previous studies, observed HFCs were weak overall. However, variation in HFCs among populations was high for some traits (although these variances were not statistically different from zero). Population‐level genetic parameters, specifically genetic diversity levels (number of alleles, observed/expected heterozygosity) and measures of identity disequilibrium (g2 and heterozygosity–heterozygosity correlations), were not associated with variation in population‐level HFCs. This latter result indicates that these metrics do not necessarily provide a reliable predictor of HFC effect sizes across populations. Importantly, diversity and identity disequilibrium statistics were not correlated, providing empirical evidence that these metrics capture different essential characteristics of populations. A complex genetic architecture likely underpins multiple fitness traits, including those associated with male fitness, which may have reduced our ability to detect HFCs in guppy populations. Further advances in this field would benefit from additional research to determine the demographic contexts in which HFCs are most likely to occur.     

Maintenance of genetic diversity in cyclic populations-a longitudinal analysis in Myodes glareolus

Conspicuous cyclic changes in population density characterize many populations of small northern rodents. The extreme crashes in individual number are expected to reduce the amount of genetic variation within a population during the crash phases of the population cycle. By long-term monitoring of a bank vole (Myodes glareolus) population, we show that despite the substantial and repetitive crashes in the population size, high heterozygosity is maintained throughout the population cycle. The striking population density fluctuation in fact only slightly reduced the allelic richness of the population during the crash phases. Effective population sizes of vole populations remained also relatively high even during the crash phases. We further evaluated potential mechanisms contributing to the genetic diversity of the population and found that the peak phases are characterized by both a change in spatial pattern of individuals and a rapid accession of new alleles probably due to migration. We propose that these events act together in maintaining the high genetic diversity within cyclical populations.