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Pollinator-mediated assortative mating in mixed ploidy populations of Chamerion angustifolium (Onagraceae).

Establishment of polyploid individuals within diploid populations is theoretically unlikely unless polyploids are reproductively isolated, pre-zygotically, through assortative pollination. Here, we quantify the contribution of pollinator diversity and foraging behaviour to assortative pollen deposition in three mixed-ploidy populations of Chamerion angustifolium (Onagraceae). Diploids and tetraploids were not differentiated with respect to composition of insect visitors. However, foraging patterns of the three most common insect visitors (all bees) reinforced assortative pollination. Bees visited tetraploids disproportionately often and exhibited higher constancy on tetraploids in all three populations. In total, 73% of all bee flights were between flowers of the same ploidy (2x-2x, 4x-4x); 58% of all flights to diploids and 83% to tetraploids originated from diploid and tetraploid plants, respectively. Patterns of pollen deposition on stigmas mirrored pollinator foraging behaviour; 73% of all pollen on stigmas (70 and 75% of pollen on diploid and tetraploid stigmas, respectively) came from within-ploidy pollinations. These results indicate that pollinators contribute to high rates of pre-zygotic reproductive isolation. If patterns of fertilization track pollen deposition, pollinator-plant interactions may help explain the persistence and spread of tetraploids in mixed-ploidy populations.

Alberta↗

The evolutionary forces maintaining a wild polymorphism of Littorina saxatilis: model selection by computer simulations.

Two rocky shore ecotypes of Littorina saxatilis from north-west Spain live at different shore levels and habitats and have developed an incomplete reproductive isolation through size assortative mating. The system is regarded as an example of sympatric ecological speciation. Several experiments have indicated that different evolutionary forces (migration, assortative mating and habitat-dependent selection) play a role in maintaining the polymorphism. However, an assessment of the combined contributions of these forces supporting the observed pattern in the wild is absent. A model selection procedure using computer simulations was used to investigate the contribution of the different evolutionary forces towards the maintenance of the polymorphism. The agreement between alternative models and experimental estimates for a number of parameters was quantified by a least square method. The results of the analysis show that the fittest evolutionary model for the observed polymorphism is characterized by a high gene flow, intermediate-high reproductive isolation between ecotypes, and a moderate to strong selection against the nonresident ecotypes on each shore level. In addition, a substantial number of additive loci contributing to the selected trait and a narrow hybrid definition with respect to the phenotype are scenarios that better explain the polymorphism, whereas the ecotype fitnesses at the mid-shore, the level of phenotypic plasticity, and environmental effects are not key parameters.

Animals↗

Evolutionary animation: how do molecular phylogenies compare to Mayr's reconstruction of speciation patterns in the sea?

Ernst Mayr used the geography of closely related species in various stages of increasing divergence to "animate" the process of geographic, or allopatric, speciation. This approach was applied to a wide set of taxa, and a seminal paper by Mayr used it to explore speciation patterns in tropical sea urchins. Since then, taxonomic information in several of these genera has been augmented by detailed molecular phylogenies. We compare Mayr's animation with the phylogenies of eight sea urchin genera placed by Mayr into four speciation groups. True to Mayr's predictions, early-stage genera have on average lower species divergence and more polytypic species than genera in later stages. For six of these genera, we also have information about the evolution of the gamete recognition protein bindin, which is critical to reproductive isolation. These comparisons show that later-stage genera with many sympatric species tend to be those with rapid bindin evolution. By contrast, early-stage genera with few sympatric species are not necessarily earlier in the divergence process; they happen to be those with slow rates of bindin evolution. These results show that the rate of speciation in sea urchins does not only depend on the steady accumulation of genome divergence over time, but also on the rate of evolution of gamete recognition proteins. The animation method used by Mayr is generally supported by molecular phylogenies. However, the existence of multiple rates in the acquisition of reproductive isolation complicates placement of different genera in an evolutionary series.

Animals↗

Genetic variation in sympatric Ascaris populations from humans and pigs in China.

It has recently been shown using genetic markers that Ascaris in humans and pigs in Central America comprise reproductively isolated populations. We present a similar analysis for a region of China in which close association between pigs and humans has been the norm for thousands of years, and agricultural practices will result in frequent exposure to eggs from both sources. DNA fragments from selected regions of mitochondrial and ribosomal DNA were amplified by PCR and allelic forms identified following digestion with a panel of restriction enzymes, using DNA from a total of 115 individual worms from both people and pigs from 2 neighbouring villages. Significant frequency differences in both mtDNA haplotypes and the rDNA spacer were found between the 2 host-associated populations, indicating that they represented reproductively isolated populations. Mitochondrial haplotype frequencies were different from those observed in Guatemala and also from other Asian Ascaris populations, suggesting low levels of gene flow between populations. However, we found no evidence for significant heterogeneity in the genetic composition of Ascaris infrapopulations in either humans or pigs, possibly indicative of agricultural practices in China which have resulted in a random distribution of alleles within the parasite populations.

Animals↗

Penetration of zona-free hamster eggs in vitro by goat spermatozoa preincubated in the reproductive tract isolated from a maturing gilt.

When goat spermatozoa were preincubated for 5-5.5 hr in the isolated uterus and oviduct from a maturing gilt, they developed the ability to penetrate zona-free hamster eggs in vitro. Overall penetration rates were 17% and 0%, respectively, 1 hr after insemination, and these increased to 59% and 54% 2 hr later. The penetration rates were not improved by incubation longer than 2 hr after insemination up to 5 hr. Goat spermatozoa preincubated for 5 hr in a m-KRB solution with or without goat serum penetrated zona-free hamster eggs with difficulty. The results suggest that a functional change in the ability of goat spermatozoa to penetrate zona-free hamster eggs may occur in the reproductive tract of different species.

Animals↗

Early events in speciation: polymorphism for hybrid male sterility in Drosophila.

Capturing the process of speciation early enough to determine the initial genetic causes of reproductive isolation remains a major challenge in evolutionary biology. We have found, to our knowledge, the first example of substantial intraspecific polymorphism for genetic factors contributing to hybrid male sterility. Specifically, we show that the occurrence of hybrid male sterility in crosses between Drosophila mojavensis and its sister species, Drosophila arizonae, is controlled by factors present at different frequencies in different populations of D. mojavensis. In addition, we show that hybrid male sterility is a complex phenotype; some hybrid males with motile sperm still cannot sire offspring. Because male sterility factors in hybrids between these species are not yet fixed within D. mojavensis, this system provides an invaluable opportunity to characterize the genetics of reproductive isolation at an early stage.

Age Factors↗

Mitochondrial and ribosomal internal transcribed spacer (ITS2) diversity of the African malaria vector Anopheles funestus.

The pattern of sequence variation in the mitochondrial DNA cytochrome b gene (cyt-b) and ribosomal DNA internal transcribed spacer 2 (ITS2) was examined in Anopheles funestus from Senegal and Burkina Faso in West Africa and Kenya in East Africa. From both West African countries, samples included individuals hypothesized to represent reproductively isolated taxa based upon different karyotypes and behaviours. Analysis of the cyt-b data revealed high haplotypic diversity (86%) and an average pairwise difference per site of 0.42%. Sequence variation was not partitioned by geographical origin or karyotype class. The most common haplotype was sampled across Africa (approximately 6000 km). Analysis of the ITS2 data revealed one of the longest spacers yet found in anophelines (approximately 704 bp). In common with other anopheline ITS2 sequences, this one had microsatellites and frequent runs of individual nucleotides. Also in common with data from other anopheline ITS2 studies, the An. funestus sequences were almost monomorphic, with only two rare polymorphisms detected. The results from both markers are congruent and do not support the hypothesis of reproductively isolated chromosomal taxa within An. funestus. Whether the lack of support by mitochondrial DNA (mtDNA) and ribosomal DNA (rDNA) sequences is a result of the recent origin of the presumptive taxa, or of the absence of barriers to gene flow, remains to be elucidated, using more rapidly evolving markers such as microsatellites.

Africa↗

Multilocus models of sympatric speciation: Bush versus Rice versus Felsenstein.

In populations of phytophagous insects that use the host plant as a rendezvous for mating, divergence in host preference could lead to sympatric speciation. Speciation requires the elimination of "generalist" genotypes, that is, those with intermediate host preference. This could occur because such genotypes have an inherent fitness disadvantage, or because preference alleles become associated with alleles that are oppositely selected on the two hosts. Although the former mechanism has been shown to be plausible, the latter mechanism has not been studied in detail. I consider a multilocus model (the "Bush model") in which one set of biallelic loci affects host preference, and a second set affects viability on the hosts once chosen. Alleles that increase viability on one host decrease viability on the other, and all loci are assumed to be unlinked. With moderately strong selection on the viability loci, preference alleles rapidly become associated with viability alleles, and the population splits into two reproductively isolated host specialist populations. The conditions for speciation to occur in this model, as measured by the strength of selection required, are somewhat more stringent than in a model in which preference and viability are controlled by the same loci (one-trait model). In contrast, the conditions are much less stringent than in a model in which speciation requires buildup of associations between viability loci and loci controlling a host-independent assortative mating trait (canonical two-trait model). Moreover, in the one-trait model, and to a lesser extent the Bush model, the strength of selection needed to initiate speciation is only slightly greater than that needed to complete it. This indicates that documenting instances of sympatric species that are reproductively isolated only by host or habitat preference would provide evidence for the plausibility of sympatric speciation in nature.

Alleles↗

Distribution and evolution of genetic caste determination in Pogonomyrmex seed-harvester ants.

We examined the distribution and ancestral relationships of genetic caste determination (GCD) in 46 populations of the seed-harvester ants Pogonomyrmex barbatus and P. rugosus across the east-to-west range of their distributions. Using a mtDNA sequence and two nuclear markers diagnostic for GCD, we distinguished three classes of population phenotypes: those with GCD, no evidence of GCD, and mixed (both GCD and non-GCD colonies present). The GCD phenotype was geographically widespread across the range of both morphospecies, occurring in 20 of 46 sampled populations. Molecular data suggest three reproductively isolated and cryptic lineages within each morphospecies, and no present hybridization. Mapping the GCD phenotype onto a mtDNA phylogeny indicates that GCD in P. rugosus was acquired from P. barbatus, suggesting that interspecific hybridization may not be the causal agent of GCD, but may simply provide an avenue for GCD to spread from one species (or subspecies) to another. We hypothesize that the origin of GCD involved a genetic mutation with a major effect on caste determination. This mutation generates genetic conflict and results in the partitioning and maintenance of distinct allele (or gene set) combinations that confer differences in developmental caste fate. The outcome is two dependent lineages within each population; inter-lineage matings produce workers, while intra-lineage matings produce reproductives. Both lineages are needed to produce a caste-functional colony, resulting in two reproductively isolated yet interdependent lineages. Pogonomyrmex populations composed of dependent lineages provide a unique opportunity to investigate genetic variation underlying phenotypic plasticity and its impact on the evolution of social structure.

Animals↗

Genetics, experience, and host-plant preference in Eurosta solidaginis: implications for host shifts and speciation.

Host-associated mating is crucial in maintaining the partial reproductive isolation between the host races of Eurosta solidaginis (Diptera: Tephritidae), a fly that forms galls on Solidago altissima and S. gigantea. (We refer to flies reared from S. gigantea as gigantea flies and those reared from S. altissima as altissima flies.) We measured the host preference of males and females of both host races, F1 hybrids between the host races, F2, and backcrosses to both host races. Male and female altissima flies and female gigantea flies had high host fidelity, whereas male gigantea flies had low host fidelity. This result suggests that there may be gene flow between the host races due to nonassortative mating that occurs when male gigantea mate with altissima females on S. altissima. This indicates assortative-mating mechanisms in addition to host-associated mating are required to produce the partial reproductive isolation between the host races that has been observed. Nongenetic factors had no influence on host preference. Larval conditioning did not influence host preference: reciprocal F1 hybrids reared in S. altissima and S. gigantea both preferred S. gigantea. Adult experience had no impact on host preference: females preferred their natal host plant regardless of which host they encountered first as an adult. The hypothesis that maternal effects influence preferences was rejected because male and female flies did not show a consistent preference for the host plant of their mother. We also found no evidence that preference was a sex-linked trait because F1 and backcrosses to the host races with different combinations of X chromosomes from the two host races preferred S. gigantea. Our results indicate that host preference is not determined by a large number of genes because preference of hybrids did not correspond to the proportion of the genome derived from each host race. The strength of the ovipuncture preference for S. gigantea by gigantea females, the females of both reciprocal F1 hybrids, the backcross to gigantea, and F2s indicates that preference is inherited nonadditively at a limited number of loci. The F1 female hybrids, however, had a weaker host preference for S. gigantea than the pure gigantea host race, indicating that there may be incomplete dominance or modifier loci. Males had different host preference patterns than females, with individual male gigantea and male F1 hybrids usually exhibiting preference exclusively for S. gigantea or S. altissima. One hypothesis explaining the difference in host preference between males and females is that the same gene influences both female and male host preference, but it is a sex-influenced gene. Thus, males carrying the gene for S. gigantea preference have an intermediate host preference, whereas females have a strong host preference to S. gigantea. In summary, we found that the host preference that produces host-associated mating is inherited nonadditively at a relatively small number of loci on autosomal genes. This mode of inheritance meets the assumptions of models of sympatric speciation, indicating that the host races could have evolved in sympatry.

Animals↗

Genetic variation in wild and hatchery stocks of Suminoe Oyster (Crassostrea ariakensis) assessed by PCR-RFLP and microsatellite markers.

Genetic variation in wild Asian populations and U.S. hatchery stocks of Crassostrea ariakensis was examined using polymerase chain reactions with restriction fragment length polymorphism (PCR-RFLP) analysis of both the mitochondrial COI gene and the nuclear internal transcribed spacer (ITS) 1 region and using 3 microsatellite markers. Hierarchical analysis of molecular variance and pairwise comparisons revealed significant differentiation (P < 0.05) between samples from the northern region, represented by collections from China and Japan, and 2 of 3 samples from southern China. PCR-RFLP patterns were identified that were diagnostic for the northern (N-type) and southern (S-type) groups. Microsatellite marker profiles were used to assign each oyster to one of the two northern or two southern populations. Results for more than 97% of the oysters were consistent with the PCR-RFLP patterns observed for each individual in that oysters with N-type patterns were assigned to one of the northern populations and those with S-type patterns to one of the southern populations. At one site of the Beihai (B) region in southern China a mix of individuals with either the N-type or S-type PCR-RFLP genotypes was found. No heterozygotes at the nuclear ITS-1 locus were found in the sample, possibly indicating reproductive isolation in sympatry. Microsatellite assignment test results of the B individuals were also consistent with identifications as either the N-type or S-type based on PCR-RFLP patterns. The parental population for one hatchery stock was this B sample, which initially was composed of almost equal numbers of northern and southern genetic types. After hatchery spawns, however, more than 97% of the progeny fell into the northern genetic group by PCR-RFLP and microsatellite assignment test analyses, indicating that the individuals with the southern genotype contributed little to the spawn, owing to gametic incompatibility, differential larval survival, or a difference in timing of sexual maturity. Overall, results suggested that oysters collected as C. ariakensis in this study, and likely in other studies as well, include two different sympatric species with some degree of reproductive isolation.

Analysis of Variance↗

Marriage behaviour in the Alpine Non Valley from 1825 to 1923.

BACKGROUND: The study is part of a research project on the marital structure of mountain populations from the Eastern Italian Alps. Little is known about marriage patterns in this Alpine area. AIM: The aim of the study is to evaluate the extent of reproductive isolation in some communities of the Non Valley (Trentino, Italy) and to investigate its microgeographic and temporal changes over the period 1825-1923. SUBJECTS AND METHODS: 4518 microfilmed marriage records from registers of seven parishes of the Non Valley were used to analyse the following: endogamy rate, inbreeding calculated both from dispensations and from isonymy, repeating pairs of surnames in marriages, isonymic relationships. RESULTS: The results show notable variability among parishes in the levels of endogamy (40-73%), inbreeding (alpha: 1.9-4.57; Ft: 0.0073-0.019) and subdivision (RPr/RP: 0.5-1.3). The values are relatively stable over the course of a century, apart from a rise in inbreeding indicated by dispensations and a slight decrease of endogamy at the beginning of the 20th century. Isonymic relationships reflect geographic proximity between populations, with minimum changes through time. CONCLUSIONS: Variations in the level of reproductive isolation within the Non Valley are consistent with the different geographic characteristics and population sizes of the settlements. Comparison with data obtained from previous studies in the Eastern Italian Alps shows that the values of the investigated biodemographic indicators are in line with the geography and altitude of the area. The slight differences in temporal trend of endogamy and inbreeding can be correlated with different migration patterns.

Consanguinity↗

A study of fluctuating asymmetry in hybrids of dwarf and normal lake whitefish ecotypes (Coregonus clupeaformis) from different glacial races.

Fish ecotypes found in north temperate lakes are increasingly used as model organisms to explore patterns and processes of population divergence that may ultimately cause speciation. Processes involved in their reproductive isolation are, however, still poorly understood. Recent experimental studies on whitefish ecotypes from different glacial races revealed that embryonic mortality of hybrids was 2.4-4.7 times higher than for parental forms. In this study, we compared fluctuating asymmetry (FA) in morphological traits of these same hybrid and pure crosses to test the hypothesis that genetic stress observed in hybrids at embryonic stages is also manifested at later developmental stages. Twelve morphological traits were used to measure asymmetry. Variable degrees of asymmetry were observed depending on traits and crosses, however there was no significant difference in FA among crosses. These results thus provided no evidence in support of the working hypothesis and indicated that genetic stress may differ among life stages. It is more likely that high hybrid embryonic mortality acts together with ecological factors at later stages to increase further the extent of reproductive isolation between sympatric whitefish ecotypes.

Animals↗

Genetic, ecological, and behavioural divergence between two sibling snapping shrimp species (Crustacea: Decapoda: Alpheus).

Examination of genetic and ecological relationships within sibling species complexes can provide insights into species diversity and speciation processes. Alpheus angulatus and A. armillatus, two snapping shrimp species with overlapping ranges in the north-western Atlantic, are similar in morphology, exploit similar ecological niches and appear to represent recently diverged sibling species. We examined phylogenetic and ecological relationships between these two species with: (i) sequence data from two mitochondrial genes (16S rRNA and COI); (ii) data on potential differences in microhabitat distribution for A. armillatus and A. angulatus; and (iii) data from laboratory experiments on the level of reproductive isolation between the two species. DNA sequence data suggest A. armillatus and A. angulatus are sister species that diverged subsequent to the close of the Isthmus of Panama, and that haplotype diversity is lower in A. armillatus than in A. angulatus. Both species are distantly related to A. heterochaelis and A. estuariensis, two species with which A. angulatus shares some similarities in coloration. Ecological data on the distribution of A. angulatus and A. armillatus from two locations revealed differences in distribution of the two species between habitat patches, with each patch dominated by one or the other species. However, there was no apparent difference in distribution of the two species within habitat patches with respect to microhabitat location. Ecological data also revealed that heterospecific individuals often occur in close proximity (i.e. within metres or centimetres) where sympatric. Behavioural data indicated that these species are reproductively isolated, which is consistent with speciation in transient allopatry followed by post-divergence secondary contact. Our data further resolve taxonomic confusion between the sibling species, A. armillatus and A. angulatus, and suggest that sympatry in areas of range overlap and exploitation of similar ecological niches by these two recently diverged species have selected for high levels of behavioural incompatibility.

Animals↗

Quantitative trait loci affecting differences in floral morphology between two species of monkeyflower (Mimulus).

Conspicuous differences in floral morphology are partly responsible for reproductive isolation between two sympatric species of monkeyflower because of their effect on visitation of the flowers by different pollinators. Mimulus lewisii flowers are visited primarily by bumblebees, whereas M. cardinalis flowers are visited mostly by hummingbirds. The genetic control of 12 morphological differences between the flowers of M. lewisii and M. cardinalis was explored in a large linkage mapping population of F2 plants n = 465 to provide an accurate estimate of the number and magnitude of effect of quantitative trait loci (QTLs) governing each character. Between one and six QTLs were identified for each trait. Most (9/12) traits appear to be controlled in part by at least one major QTL explaining >/=25% of the total phenotypic variance. This implies that either single genes of individually large effect or linked clusters of genes with a large cumulative effect can play a role in the evolution of reproductive isolation and speciation.

Chromosome Mapping↗

Ring species as bridges between microevolution and speciation.

A demonstration of how small changes can lead to species-level differences is provided by ring species, in which two reproductively isolated forms are connected by a chain of intermediate populations. We review proposed cases of ring species and the insights they provide into speciation. Ring species have been viewed both as illustrations of the history of divergence of two species from their common ancestor and as demonstrations that speciation can occur in spite of gene flow between the diverging forms. Theoretical models predict that speciation with gene flow can occur when there is divergent ecological selection, and geographical differentiation increases the likelihood of speciation. Thus ring species are ideal systems for research into the role of both ecological and geographical differentiation in speciation, but few examples have been studied in detail. The Greenish warbler is a ring species in which two northward expansions around the Tibetan plateau have been accompanied by parallel evolution in morphology, ecology, and song length and complexity. However, songs have diverged in structure, resulting in a lack of recognition where the reproductively isolated forms come into contact in Siberia. Our analysis suggests that these differences could have arisen even with gene flow, and that parallel rather than divergent ecological changes have led to divergence in sexually selected traits and subsequent speciation.

Animals↗

Evidence that female preferences have shaped male signal evolution in a clade of specialized plant-feeding insects.

Mate choice is considered an important influence in the evolution of mating signals and other sexual traits, and--since divergence in sexual traits causes reproductive isolation--it can be an agent of population divergence. The importance of mate choice in signal evolution can be evaluated by comparing male signal traits with female preference functions, taking into account the shape and strength of preferences. Specifically, when preferences are closed (favouring intermediate values), there should be a correlation between the preferred values and the trait means, and stronger preferences should be associated with greater preference-signal correspondence and lower signal variability. When preferences are open (favouring extreme values), signal traits are not only expected to be more variable, but should also be shifted towards the preferred values. We tested the role of female preferences in signal evolution in the Enchenopa binotata species complex of treehoppers, a clade of plant-feeding insects hypothesized to have speciated in sympatry. We found the expected relationship between signals and preferences, implicating mate choice as an agent of signal evolution. Because differences in sexual communication systems lead to reproductive isolation, the factors that promote divergence in female preferences--and, consequently, in male signals--may have an important role in the process of speciation.

Acoustic Stimulation↗

Experimental manipulation of sexual selection and the evolution of courtship song in Drosophila pseudoobscura.

Courtship song serves as a sexual signal and may contribute to reproductive isolation between closely related species. Using lines of Drosophila pseudoobscura experimentally selected under different sexual selection regimes, we tested whether increased promiscuity and enforced monogamy led to evolutionary changes in courtship song elements. In D. pseudoobscura, males produce both a low and high rate repetition song. We found that both song types diverged after selection and that the direction of changes was consistent with ordered hypotheses of predicted directions of change under the different mating system structures. In particular, latency to the initiation of song and duration of the interpulse interval (IPI) decreased in highly promiscuous lines and increased in monogamous lines. These results suggest that courtship song may rapidly evolve under different mating system structures, representing either functional evolution of more stimulatory song under conditions of strong sexual selection, or might result from increased courtship vigor, or represent correlated evolution. Some speciation theory predicts that increased sexual selection should result in increased variance of traits between allopatric populations, facilitating reproductive isolation. We also found that courtship song elements were not equally variable between replicate lines.

Animals↗