Trans-continental visible morph-frequency variation at homologous loci in two species of spider, Enoplognatha ovata s.s. & E. latimana

The spiders Enoplognatha ovata s.s. and E. latimana are sibling species which share a number of visible genetic polymorphisms. Data on colour and black-spotting morph frequencies in these species have been collected from 67 sites in western Europe. Sixty nine percent of the collections contained both species. In all adequately-sized samples, both species were polymorphic for colour and, in general, exhibited the same rank order of morphs lineata and redimita. (The top dominant morph, ovata, has not been found in E. latimana). Colour-morph frequencies are not correlated between species in sympatric populations from mainland Europe and from a previously studied area in Pembrokeshire, South Wales. Although associations with certain climatic variables are evident in E. ovata they are not consistent between transects, making their biological significance unclear. For black spotting, E. ovata s.s. is nearly fixed for spotting throughout mainland Europe but is highly variable in the Pembrokeshire populations. E. latimana is polymorphic in both areas. In Europe, spotting frequencies in E. latimana show significant associations with climatic factors but, again, their biological significance is not obvious. In E. ovata s.s. the variance in both colour and spotting frequencies among populations in Pembrokeshire is significantly greater than that in the whole of mainland Europe. The implications of these and previous results are considered in the context of the persistence of visible polymorphisms across species and the forces which determine morph frequencies in local populations.     

Long-term variation in colour-morph frequencies in the spider Enoplognatha ovata (Glerck) (Araneae: Theridiidae): natural selection, migration and intermittent drift

The spider Enoplognatha ovata is univoltine. It has three principal visible morphs, lineata, redimita and mala , whose genetics are understood. Phenotype frequencies and population sizes have been estimated at 44 sites in a single valley in Yorkshire for periods of up to 15 years. Geographically, recognizable areas of high and low frequencies of redimita and ovata are found, typically a few hundred metres across. Statistically significant frequency differences also exist on a scale of tens of metres between adjacent sites. Temporal variation in lineata allele frequency has been investigated using a weighted regression technique assuming either no deterministic change or a steady change at constant rate. There is little direct evidence for selection but variation within sites is less than would be expected through genetic drift. A negative relationship is demonstrated between initial allele frequency in a site and the direction and rate of allele frequency change with time, suggesting that populations are moving towards a global equilibrium. This conclusion is insensitive to varying assumptions about the relationship between estimated and effective population sizes. Evidence from transplantation experiments and from the most isolated site indicate that both migration and selection contribute to this trend. We suggest that populations were stochastically perturbed from the equilibrium during bottlenecks in population size which resulted from major habitat disturbance about 35 years ago.     

Genetic drift within a protected polymorphism: enigmatic variation in color-morph frequencies in the candy-stripe spider, Enoplognatha ovata

The candy-stripe spider, Enoplognatha ovata, exhibits a striking color polymorphism comprising three morphs. A number of lines of evidence strongly suggest that this polymorphism is maintained by natural selection: its presence in a sister species, E. latimana; the physical nature of the variation; the virtual lack of monomorphic populations; the highly consistent rank-order of morphs within populations; and the presence of large-scale clines associated with climatic variables. However, the absence of selection is equally strongly suggested by very local surveys of morph frequencies over space and time, perturbation experiments, and a variance in morph frequency between populations that is virtually independent of spatial scale. In addition, local spatial patterns in one study site (Nidderdale, Yorkshire, England) have been explained in terms of intermittent drift over half a century ago, a hypothesis supported here by the distributions of four other genetic markers (two allozyme and two visible polymorphisms). A heuristic model is suggested that reconciles these apparently contradictory messages regarding the importance of drift and selection in this system. It is proposed that when allele frequencies of the color morph redimita lie between approximately 0.05 and 0.3, the deltaq on q plot is very shallow, so that within this region, where the majority of populations lie, selection is weak and drift is the major force determining local morph frequencies. However, outside this range of frequencies, powerful selection acts to protect the polymorphism. This model may apply to polymorphisms in other species and explain why evidence of selection in natural populations is often elusive.     

Ecological and developmental dynamics of a host-parasite system involving a sea anemone and two ctenophores

The lined sea anemone Edwardsiella lineata has evolved a derived parasitic life history that includes a novel body plan adapted for life inside its ctenophore hosts. Reputedly its sole host is the sea walnut, Mnemiopsis leidyi, a voracious planktivore and a seasonally abundant member of many pelagic ecosystems. However, we have observed substantially higher E. lineata prevalence in a second ctenophore species, the ctenophore predator Bero? ovata. The interplay among these 3 species has important conservation consequences as M. leidyi introductions are thought to be responsible for the severe depletion of numerous commercial fisheries in the Mediterranean basin, and both E. lineata and B. ovata have been proposed as biological controls for invasive M. leidyi. Over a 3-yr period (2004-2006), we collected 8,253 ctenophores from Woods Hole, Massachusetts, including M. leidyi, B. ovata, and a third ctenophore, Pleurobrachia pileus, and we recorded E. lineata infection frequencies, parasite load, and parasite location. We also conducted laboratory experiments to determine the likely mechanisms for parasite introduction and the effect of each host on parasite development. We observed peak E. lineata infection frequencies of 0% in P. pileus, 59% in M. leidyi, and 100% in B. ovata, suggesting that B. ovata could be an important natural host for E. lineata. However, in laboratory experiments, E. lineata larvae proved far more successful at infecting M. leidyi than B. ovata, and E. lineata parasites excised from M. leidyi exhibited greater developmental competence than parasites excised from B. ovata. Although we show that E. lineata is efficiently transferred from M. leidyi to B. ovata when the latter preys upon the former, we conclude that E. lineata larvae are not well adapted for parasitizing the latter species and that the E. lineata parasite is not well adapted for feeding in B. ovata; these developmental and ecological factors underlie the host specificity of this recently evolved parasite.     

Postglacial colonization history reflects in the genetic structure of natural populations of Festuca rubra in Europe

We conducted a large‐scale population genetic survey of genetic diversity of the host grass Festuca rubra s.l., which fitness can be highly dependent on its symbiotic fungus Epichloë festucae, to evaluate genetic variation and population structure across the European range. The 27 studied populations have previously been found to differ in frequencies of occurrence of the symbiotic fungus E. festucae and ploidy levels. As predicted, we found decreased genetic diversity in previously glaciated areas in comparison with nonglaciated regions and discovered three major maternal genetic groups: southern, northeastern, and northwestern Europe. Interestingly, host populations from Greenland were genetically similar to those from the Faroe Islands and Iceland, suggesting gene flow also between those areas. The level of variation among populations within regions is evidently highly dependent on the postglacial colonization history, in particular on the number of independent long‐distance seed colonization events. Yet, also anthropogenic effects may have affected the population structure in F. rubra. We did not observe higher fungal infection rates in grass populations with lower levels of genetic variability. In fact, the fungal infection rates of E. festucae in relation to genetic variability of the host populations varied widely among geographical areas, which indicate differences in population histories due to colonization events and possible costs of systemic fungi in harsh environmental conditions. We found that the plants of different ploidy levels are genetically closely related within geographic areas indicating independent formation of polyploids in different maternal lineages.     

Selective predation favouring cryptic individuals of marine snails (Littorina)

Many gastropods have inherited conspicuous shell colour polymorphisms. A challenging question is, are colour frequencies under selection or is polymorphism owing to random evolutionary processes? The intertidal species Littorina saxatilis (a rock‐dweller) and L. obtusata (confined to macroalgae) both have genetically determined shell colour variation. In Iceland, Littorina obtusata are mostly cryptic on brown macroalgae by having brown or yellow shells (~95% of the snails), while Littorina saxatilis often appears conspicuous to the background of dark rocks owing to non‐cryptic colours (15–20%). This difference may be due to selective elimination of conspicuously coloured L. obtusata by visual predators, while L. saxatilis, largely living in another habitat, is not under a similarly intense colour selection. To test this hypothesis we increased the frequencies of conspicuous L. saxatilis in experimental populations (from <12 to 55%) and placed these in the seaweed zone, the main habitat of L. obtusata. Fifteen populations were released on isolated spots of seaweed and three of these were covered by net cages to exclude bird predators. One month later, yellow snails had increased in frequency within the patches, and to our surprise the result did not differ between bare and caged patches. This suggests selection favouring a colour that matches the background of fucoid seaweeds by visual predators able to enter the cages. Birds acted as important predators by picking 16% of the experimental snails in the uncaged spots, but were unable to enter the caged spots. However, the bird predation was non‐selective with respect to snail colour. For various reasons the most likely predators able to enter the cages were intertidal fish, these were thus responsible for the selection of non‐cryptic snails. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society, 2002, 76 , 137–144.     

Cytogenetic analysis of three Italian populations of Coregonus lavaretus (Pisces, Salmoniformes) with chromosomal localization of major and minor ribosomal genes, and telomeric repeats

The European whitefish, Coregonus lavaretus, widely distributed in freshwater of northern Europe and introduced into the major lakes of northern Italy, has been restocked in central Italian lakes. In accordance with current managing practices, a reduced number of spawners contribute to reproduction within each lake and a certain degree of isolation is to be expected between populations from different lakes, resulting in the rapid fixing of chromosomal changes. A detailed survey of three populations from different lakes was carried out using classical and molecular cytogenetic techniques, to verify if specific chromosomal markers are present in the distinct populations. The comparative analysis revealed intraspecific variability of NORs and fixed differences in their number in the three populations. A co-localization of major and minor rRNA genes on one chromosome site was also observed. The original data regarding the chromosome mapping of the (TTAGGG)(n) telomeric repeat obtained in this study, demonstrated their exclusively terminal distribution, and a conspicuous inter-chromosomal variation in the number of repeats. The results are compared with data available for populations from native geographic ranges.     

Heterozygosity and ethnic variation in Japanese platelet proteins

Summary Sixty-two polypeptides visualized on silver-stained two-dimensional polyacrylamide gel electrophoresis (2-D PAGE) preparations of platelets from 85 Japanese subjects, in total 5,252 spots, have been scored for genetic variation. Inherited variants of 11 of the polypeptides were recognized; the index of heterozygosity was 2.4%±0.2%. Thus far, 10 genetic polymorphisms identified by 2-D PAGE of plasma, erythrocytes, or platelets have been identified in both Japanese and Caucasian subjects. A comparison of allele frequencies reveals four significant ethnic differences. We also observed four polypeptides exhibiting a low frequency polymorphism in one group but not in the other, as well as three polymorphisms in Caucasians for which no counterpart polypeptide has thus far been recognized in the Japanese group and, vice versa, 11 such polymorphisms in Japanese. Although a similar comparison of 7 enzyme polymorphisms studied with one-dimensional electrophoresis (1-D E) in the same populations revealed a relatively higher number of significant ethnic differences, evidence is presented that this is due primarily to the greater number of 1-D E observations entering into the calculation. It is argued that this similarity in the frequency of ethnic differences among the polypeptides studied by 2-D PAGE and by 1-D E is further evidence that the proteins revealed by 2-D PAGE do not differ greatly in their response to the interplay of mutation, selection, and drift from those revealed by 1-D E studies of plasma proteins and erythrocyte enzymes.     

Genetic studies of human apolipoproteins. XVII: Population genetics of apolipoprotein polymorphisms in American Samoa

Variation in human apolipoprotein genes is a major source of phenotypic differences in human lipid metabolism. Data regarding genetic variation at apolipoprotein loci in various populations are only beginning to accumulate, and they suggest that different populations vary widely in distribution of apolipoprotein alleles. Using isoelectric focusing-immunoblotting techniques, we screened 67 serum samples from self-identified Samoan residents of American Samoa to investigate structural variation at six apolipoprotein loci: A-I, A-II, A-IV, C-II, E, and H. The APO A-I, A-II, and C-II loci were found to be monomorphic by isoelectrical focusing. In Samoans, the common three-allele polymorphism was observed for APO E, with no striking differences in frequencies from Caucasian populations. The three common alleles of the APO H locus also were identified; however, frequencies of the less common alleles (APO H*I and APO H*3) were different from those observed elsewhere for Caucasians.     

Variation and seasonal changes in hoverfly species: interactions between temperature, age and genotype

Eristalis tenax L. and E. intricarius L. are two hoverflies which vary considerably in colour pattern. Whilst much of the phenotypic variation in both species is due to genetic variation at major gene loci, there are interactions with pupal temperature and with age of adult. Low pupal temperatures produce, on average, darker abdominal patterns in E. tenax , although the effect is hard to pick out in natural populations. Changes of pattern with age convert a bimodal autumn population to a unimodal post-hibernation population in the following spring. Hair colour is also made darker by cold treatment. Pupal treatments have a strong effect on hair colouration in E. intricarius. High temperatures inhibit the production of black hairs on the thorax, but not all the genotypes are equally sensitive to temperature. Seasonal fluctuations in colour pattern frequencies are detectable in E. intricarius. In three Eristalis species that have been studied so far, the interactions between genotype, age and temperature operate in quite different ways. The temperature responses may be relevant to theories of thermal melanism, although the colour pattern polymorphisms are, more obviously, examples of Batesian mimicry.     

The role of changes in environmental quality in multitrait plastic responses to environmental and social change in the model microalga Chlamydomonas reinhardtii

Intraspecific variation plays a key role in species'' responses to environmental change; however, little is known about the role of changes in environmental quality (the population growth rate an environment supports) on intraspecific trait variation. Here, we hypothesize that intraspecific trait variation will be higher in ameliorated environments than in degraded ones. We first measure the range of multitrait phenotypes over a range of environmental qualities for three strains and two evolutionary histories of Chlamydomonas reinhardtii in laboratory conditions. We then explore how environmental quality and trait variation affect the predictability of lineage frequencies when lineage pairs are grown in indirect co‐culture. Our results show that environmental quality has the potential to affect intraspecific variability both in terms of the variation in expressed trait values, and in terms of the genotype composition of rapidly growing populations. We found low phenotypic variability in degraded or same‐quality environments and high phenotypic variability in ameliorated conditions. This variation can affect population composition, as monoculture growth rate is a less reliable predictor of lineage frequencies in ameliorated environments. Our study highlights that understanding whether populations experience environmental change as an increase or a decrease in quality relative to their recent history affects the changes in trait variation during plastic responses, including growth responses to the presence of conspecifics. This points toward a fundamental role for changes in overall environmental quality in driving phenotypic variation within closely related populations, with implications for microevolution.     

A countrywide survey of colour morph frequencies in the spider Enoplognatha ovata (Glerck) (Araneae: Theridiidae): evidence for natural selection

Data on the frequencies of the main colour morphs ( lineata, redimita and ovata ) of Enoplognatha ovata have been collected from a total of 454 Ordnance Survey 10 km squares distributed throughout Great Britain. Only c. 0.5% of samples were monomorphic, for lineata in each case. Multiple regression analysis has been used to assess possible associations between morph frequencies and principal components derived from nine environmental variables. The distribution of morph frequencies is not random but shows weak clines associated with certain climatic factors. These large scale clines indicate the action of natural selection although very local variations in morph frequencies may result from selection and/or drift.     

Population Genetics of Euphydryas Butterflies. I. Genetic Variation and the Neutrality Hypothesis

Twenty-one populations of the checkerspot butterfly, Euphydryas editha, and ten populations of Euphydryas chalcedona were sampled for genetic variation at eight polymorphic enzyme loci. Both species possessed loci that were highly variable from population to population and loci that were virtually identical across all populations sampled. Our data indicate that the neutrality hypothesis is untenable for the loci studied, and therefore selection is indicated as the major factor responsible for producing these patterns. Thorough ecological work allowed gene flow to be ruled out (in almost all instances) as a factor maintaining similar gene frequencies across populations. The Lewontin-Krakauer test indicated magnitudes of heterogeneity among standardized variances of gene frequencies inconsistent with the neutrality hypothesis. The question of whether or not to correct this statistic for sample size is discussed. Observed equitability of gene frequencies of multiple allelic loci was found to be greater than that predicted under the neutrality hypothesis. Genetic differentiation persisting through two generations was found between the one pair of populations known to exchange significant numbers of individuals per generation. Two matrices of genetic distance between populations, based on the eight loci sampled, were found to be significantly correlated with a matrix of environmental distance, based on measures of fourteen environmental parameters. Correlations between gene frequencies and environmental parameters, results of multiple regression analysis, and results of principle component analysis showed strong patterns of association and of "explained" variation. The correlation analyses suggest which factors might be further investigated as proximate selective agents.     

Countergradient variation in the sexual coloration of guppies (Poecilia reticulata): drosopterin synthesis balances carotenoid availability

Trinidad guppies (Poecilia reticulata) are distributed along an environmental gradient in carotenoid availability that limits the carotenoid content of the orange spots of males. The amount of synthetic red pteridines (drosopterins) in the orange spots covaries with the carotenoid content, such that the ratio of the two types of pigments is roughly conserved across streams. Carotenoids and drosopterins have different spectral properties and thus the ratio of the two types of pigments affects the shape of the orange spot reflectance spectrum. Geographic conservation of the carotenoid:drosopterin ratio suggests that males may be under selection to maintain a particular hue. We tested this hypothesis by comparing the pigmentation and coloration of guppies from six streams in the field to that of second-generation descendants of the same populations raised on three dietary carotenoid levels in the laboratory. The results show clearly that the geographic variation in drosopterin production is largely genetic and that the hue of the orange spots is conserved among populations in the field, relative to the laboratory diet groups. This is a countergradient pattern because genetic differences between populations in drosopterin production mask the effect of carotenoid availability on the hue of the orange spots. The potential for countergradient sexual selection to contribute to reproductive isolation between populations is discussed.     

彝族、藏族与哈尼族血清胆硷酯酶的遗传多态性

按照Kalow等的方法对中国3个人群的血清胆硷酯酶(简称ChE)的遗传多态性进行了研究。3个人群是四川省布拖县彝族(197人)、西藏自治区拉萨市藏族(100人)及云南省元江县哈尼族(170人)。根据每一个体的地布卡因值和氟化钠值,按Motulsky的分型标准进行分型。结果表明,在彝族中有3人属一般型+抗氟化物型(UF);在藏族中有2名UF;在哈尼族中也有3名UF,未发现不典型型(即抗地布卡因型的纯合子与杂合子),也未发现抗氟化物型的纯合子。因此,E_1~f基因频率在彝族、藏族与哈尼族中分别为0.0076、0.0100及0.0088。正常的常见等位基因E_1~(?)的频率分别为0.9924、0.9900及0.9912。这3个人群ChE表型分布均符合Hardy-Weinberg平衡法则。他们的基因频率与蒙古人种其他人群是一致的。     

Studies in the Melica ciliata – complex: 1. Distribution of allozyme variation within and among individuals, populations and geographic regions

Allozyme variation in 83 European populations of the Melica ciliata/ transsilvanica species complex has been investigated. In spite of a diploid chromosome number, most loci were duplicated and patterns of within-population variation were not explainable unless apomixis or uncommon patterns of chromosomal segregation were assumed. Allelic richness and genetic diversity were very high but most of the variation (55-98%) was due to population differentiation. Geographic patterns were analysed both using phenetic, based on allele frequencies within populations, and cladistic, based on multi-locus genotypes, methods. Both methods congruently recognised three large groups of populations with discrete geographic distributions, in addition to a number of smaller groups. Of the three main groups, one was distributed in eastern and central Europe, one was restricted to western Europe north of the Alps and one was distributed in Iberia, Brittany and along the western shores of the Mediterranean. These groups are likely to have had separate histories but there appears to have been significant gene-flow among them. Among the less distinct groups, it was suggestive that some, e.g. the group containing all populations from the disjunct distribution in the Baltic basin, have been formed by introgression between the three major groups, whereas other deviating populations may represent additional distinct entities.     

Isozyme Frequency Patterns in DROSOPHILA PAVANI Associated with Geographical and Seasonal Variables

Fourteen population samples of Drosophila pavani were obtained from a number of localities in Chile. The populations sampled were dispersed over 7 degrees of latitude and 1800 meters of elevation, and were drawn at three different times. Sixteen electrophoretic loci were assayed for each population; eight of the loci were analyzed statistically for geographic variation; the other eight were essentially monomorphic. For all eight variable loci, variation in allelic frequencies among populations was highly significant. In all cases, a significant portion of the variation among populations was associated with variation in gross environmental variables (latitude, elevation, month of collection). The implications of the evidence were discussed, and the authors concluded that there was suggestive evidence for selection.     

三种海桑属濒危红树植物的资源分布与濒危现状

通过文献查阅和访问咨询，结合野外调查，对3种海桑属(Sonneratia)濒危红树植物的资源分布和濒危现状进行了初步研究。结果显示：卵叶海桑(Sonneratia ovata Backer)有2个自然居群和3个人工居群，个体数量分别为231和38棵；海南海桑(S. × hainanensis W. C. Ko, E.Y. Chen & W.Y. Chen)有4个自然居群和2个人工居群，个体数量分别为139和11棵；拟海桑(S. × gulngai N. C. Duke & B. R. Jackes)有4个自然居群和3个人工居群，个体数量分别为345和12棵；它们的自然居群更新能力都极差，人工种群也均未见新生个体。气候变化和人为干扰是种群规模急剧减小的直接原因，自身更新困难是种群规模难以恢复的制约因素。因此，建议在天然种群就地保护的基础上，优化迁地保护策略，加强人工辅助育种研究，加快突破育种难题，并科学抚育野外回归种群，以保证其种群的正常更新。     

Reproduction in Laboratory Stocks of Pleuroxus (Chydoridae,Cladocera) Under the Influence of Photoperiod and Light Intensity

The effects of photoperiod and light intensity on the reproduction of laboratory populations of three species in Pleuroxus from different geographic origins were determined statistically. There were highly significant differences in the numbers of gamogenetic individuals produced between the populations at different photoperiods but not at different light intensities. More males than ephippial females appeared at long-day photoperiods. Pleuroxus denticulatus stocks showed greater variation of gamogenetic response to photoperiod and geographic origin suggesting that local populations have evolved different reproductive patterns to meet the major environmental stresses of the region.     

Nuclear RFLP variation in Eucalyptus camaldulensis Dehnh. from northern Australia

Eucalyptus camaldulensis Dehnh. is the most widely planted eucalypt in the tropics. Natural populations are riparian and sampling strategies for breeding programmes have assumed that gene flow among drainage basins is limited. RFLP variation, within and among 31 populations from river systems across northern Australia, was analysed to test this hypothesis. To allow comparisons within and between river systems, trees were sampled from up to three populations per river system. Allele frequencies were correlated with longitude for more than half the 33 RFLP loci surveyed. Genetic identity was greatest between populations in closest geographic proximity, irrespective of river system, suggesting that sampling strategies for breeding programmes should be based on geographic distance rather than river system. The level of genetic variation was similar throughout the geographic range examined (mean H(E) = 0.49). However, there was evidence of a barrier to gene flow between populations in the east and west of the species range. The RFLP data support morphological evidence of hybridisation between E. camaldulensis and E. tereticornis Sm. in several populations in northeast Queensland and the genetic divergence of E. camaldulensis subsp. simulata Brooker and Kleinig.