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Bimodal hybrid zones and speciation.

Contact zones exemplify a series of stages in speciation. In unimodal hybrid zones intermediates predominate; in bimodal zones hybrids are rare and parental forms predominate; and finally, species might overlap, but never hybridize. Recent studies show bimodality to be associated strongly with assortative mating or fertilization, and only weakly with overall levels of genetic divergence or intrinsic genomic incompatibility. Ecological divergence across most bimodal hybrid zones suggests that ecology contributes more to speciation than genomic incompatibility. This continuum of stable contact zones provides empirical evidence for a route to speciation, which does not require allopatry.

Journal Article↗

Niche lability in the evolution of a Caribbean lizard community.

Niche conservatism--the tendency for closely related species to be ecologically similar--is widespread. However, most studies compare closely related taxa that occur in allopatry; in sympatry, the stabilizing forces that promote niche conservatism, and thus inhibit niche shifts, may be countered by natural selection favouring ecological divergence to minimize the intensity of interspecific interactions. Consequently, the relative importance of niche conservatism versus niche divergence in determining community structure has received little attention. Here, we examine a tropical lizard community in which species have a long evolutionary history of ecological interaction. We find that evolutionary divergence overcomes niche conservatism: closely related species are no more ecologically similar than expected by random divergence and some distantly related species are ecologically similar, leading to a community in which the relationship between ecological similarity and phylogenetic relatedness is very weak. Despite this lack of niche conservatism, the ecological structuring of the community has a phylogenetic component: niche complementarity only occurs among distantly related species, which suggests that the strength of ecological interactions among species may be related to phylogeny, but it is not necessarily the most closely related species that interact most strongly.

Animals↗

Genomic evidence for divergence with gene flow in host races of the larch budmoth.

Ecological divergence in the face of gene flow has recently become implicated as a potentially important cause of speciation and adaptive radiation. Here, we develop a genomic approach to test for divergent selection in sympatric host races of the larch budmoth Zeiraphera diniana (Lepidoptera: Tortricidae). We analysed hundreds of amplified fragment length polymorphism markers in 92 individuals in sympatric and allopatric populations, and in two backcross broods used to map the markers to individual chromosomes. The results directly confirm the existence of natural hybridization and demonstrate strong heterogeneity between chromosomes in terms of molecular divergence between host races (the average level of divergence was FST = 0.216). However, genomic heterogeneity was not found when we analysed divergence between geographically separated populations of the same host race. We conclude that the variance of the level of sympatric divergence among chromosomes is the footprint of divergent selection acting on a few linkage groups, combined with appreciable gene flow that homogenizes between-race variation at the remaining linkage groups. These results, coupled with other recent multilocus analyses of sister species pairs, demonstrate that selection-driven sympatric phase of genetic divergence in the presence of gene flow is a likely feature of speciation.

Analysis of Variance↗

Host-plant adaptation drives the parallel evolution of reproductive isolation.

Parallel evolution of similar traits in independent populations that inhabit ecologically similar environments strongly implicates natural selection as the cause of evolution. Parallel speciation is a special form of parallel evolution where traits that determine reproductive isolation evolve repeatedly, in closely related populations, as by-products of adaptation to ecological conditions. The outcome of such parallel evolution is that ecologically divergent pairs of populations exhibit greater levels of reproductive isolation than ecologically similar pairs of populations of a similar or younger age. The parallel evolution of reproductive isolation provides strong evidence for natural selection in the process of speciation, but only one conclusive example from nature is known. Populations of the walking-stick insect Timema cristinae that use different host-plant species have diverged in body size and shape, host preference, behaviour and the relative frequency of two highly cryptic colour-pattern morphs. Here we report that divergent selection for host adaptation, and not genetic drift, has promoted the parallel evolution of sexual isolation in this species. Our findings represent a clear demonstration that host-plant adaptation can play a crucial and repeatable role in the early stages of speciation.

Adaptation, Physiological↗

Variation in vasotocin immunoreactivity in the brain of recently isolated populations of a death valley pupfish, Cyprinodon nevadensis.

Pupfishes in the Death Valley region of California and Nevada comprise a monophyletic group of populations that became isolated in remote streams and springs over the past 20,000 years. These aquatic habitats show considerable ecological diversity, and allopatric populations have evolved differences in morphology and behavior. Here we investigated whether the divergence of pupfish populations in Death Valley might be associated with changes in arginine vasotocin (AVT). We used immunocytochemistry to compare the expression of AVT in the brain of Amargosa pupfish (Cyprinodon nevadensis) from two Death Valley populations: (1) the Amargosa River-a highly variable desert stream containing the Cyprinodon nevadensis amargosae subspecies, and (2) Big Spring-a comparatively stable springhead and outflow inhabited by Cyprinodon nevadensis mionectes. These particular populations have been isolated from each other for only 400-4000 years. In both populations AVT-immunoreactive somata localized to parvocellular and magnocellular neurons in the preoptic area, with AVT-immunoreactive fibers extending ventrolaterally to innervate the pituitary. We found that both parvocellular and magnocellular AVT-immunoreactive neurons were significantly larger in males and females from the Amargosa River population than in same sex pupfish from Big Spring. Our findings suggest that the divergent ecological conditions of these two habitats have brought about changes in AVT pathways in the brain.

Animals↗

Molecular and morphological evolution of the amphipod radiation of Lake Baikal.

Lake Baikal, in Siberia, Russia, contains the highest biodiversity of any extant lake, including an impressive radiation of gammaroidean amphipods that are often cited as a classic case of adaptive radiation. However, relationships among Baikal's amphipods remain poorly understood. The phylogenetic history of 32 Lake Baikal amphipod species, representing most major lineages of the endemic fauna, was examined using three genes (COI, 16S rRNA, and 18S rRNA), and 152 morphological characters. Results support monophyly of the largest and most diverse of the Baikalian families, the Acanthogammaridae. Analyses suggest that a second Baikalian family, the fossorial Micruropodidae, is paraphyletic and composed of two divergent clades, one of which includes Macrohectopus branickii, a morphologically specialized pelagic planktivore traditionally assigned its own family. The extreme morphological and ecological divergence of Macrohectopus from its close genetic relatives, and conversely, the large genetic distances among other morphologically similar micruropodids, suggest that morphological and molecular evolution have often been uncoupled during the radiation of Baikal's amphipods. This study suggests that the amphipod fauna of Lake Baikal is polyphyletic; originating from two independent invasions of the lake.

Animals↗

Hidden genomic structure and widespread structural polymorphism across environmental gradients in the spiny sea star Marthasterias glacialis.

Genomic regions of reduced recombination can preserve linkage among co-adapted alleles, facilitating local adaptation despite high connectivity. Such regions-often generated by chromosomal inversions-may be especially important in highly dispersive marine taxa yet remain poorly documented in echinoderms. Here, we combined a chromosome-level reference genome with genome-wide ddRAD-seq from 296 Marthasterias glacialis individuals across 19 Atlantic-Mediterranean locations to quantify population structure and scan for recombination-suppressed haploblocks. Genome-wide neutral markers showed significant population differentiation together with evidence of high connectivity, revealed by the presence of inter-ecoregion migrants. Additionally, we identified 16 polymorphic haploblocks with patterns consistent with putative chromosomal inversions spanning 18.6% of the genome. Haploblock haplotypes were strongly environmentally and geographically structured and contained genes with key functions in stress response, osmoregulation and thermal tolerance. Haplotype distributions also paralleled previously described mitochondrial lineages despite nuclear gene flow, consistent with a model of ancient divergence followed by secondary contact. Overall, our results suggest a role for widespread structural polymorphism in adaptive differentiation in Echinodermata, providing a framework for linking echinoderm genome rearrangements to ecological divergence. Marthasterias glacialis thus emerges as a promising system to explore how structural variation contributes to adaptation and genome evolution in highly dispersive organisms.

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↗

Phylogeography of the Calonectris shearwaters using molecular and morphometric data.

We investigated phylogenetic relationships and the biogeographic history of the Calonectris species complex, using both molecular and biometric data from one population of the Cape Verde shearwater Calonectris edwardsii (Cape Verde Islands), one from the streaked shearwater C. leucomelas (western Pacific Ocean) and 26 from Cory's shearwater populations distributed across the Atlantic (C. d. borealis) and the Mediterranean (C. d. diomedea). The streaked shearwater appeared as the most basal and distant clades, whereas the genetic divergences among the three main clades within the Palearctic were similar. Clock calibrations match the first speciation event within Calonectris to the Panama Isthmus formation, suggesting a vicariant scenario for the divergence of the Pacific and the Palearctic clades. The separation between the Atlantic and Mediterranean clades would have occurred in allopatry by range contraction followed by local adaptation during the major biogeographic events of the Pleistocene. The endemic form from Cape Verde probably evolved as a result of ecological divergence from the Mediterranean subspecies. Finally, one Mediterranean population (Almeria) was unexpectedly grouped into the Atlantic subspecies clade, both by genetic and by morphometric analyses, pointing out the Almeria-Oran oceanographic front (AOOF) as the actual divide between the two Cory's shearwater subspecies. Our results highlight the importance of oceanographic boundaries as potentially effective barriers shaping population and species phylogeographical structure in pelagic seabirds.

Animals↗

Genomic signatures of host-range divergence in the generalist Beauveria bassiana and the specialist Beauveria brongniartii.

Entomopathogenic fungi of the genus Beauveria are widely used biological control agents that infect diverse insect hosts and can also associate with plants as rhizosphere colonizers and endophytes. Within this genus, Beauveria bassiana is a cosmopolitan generalist, whereas Beauveria brongniartii exhibits a narrower host range, primarily targeting soil-dwelling coleopteran larvae with limited evidence of plant colonization. To explore genomic differentiation associated with this ecological divergence, the commercially exploited B. brongniartii strain BIPESCO2 and B. bassiana ATHUM 4946 were sequenced using Oxford Nanopore technology, followed by comparative genomic analyses across multiple strains. Orthology identified species-specific gene families, although overall genome architecture and core gene content were highly conserved. The CAZyme repertoires were nearly identical, indicating retention of a versatile enzymatic toolkit supporting plant association, saprotrophy, and insect pathogenicity. In contrast, biosynthetic gene clusters displayed substantial variation, including structural remodeling of Beauveria-specific virulence-associated clusters and expansion of type I polyketide synthase clusters in B. brongniartii. Effector prediction revealed a conserved core of largely uncharacterized proteins alongside species-specific orthogroups enriched in adhesion-, immunity-, and cuticle-interaction domains. Together, these findings indicate that host-range divergence in Beauveria is associated with compartmentalized genomic differentiation, particularly in secondary metabolism and a limited subset of lineage-specific virulence factors, rather than in the conserved core infection machinery.

Beauveria↗

On the origin of species by sympatric speciation.

Understanding speciation is a fundamental biological problem. It is believed that many species originated through allopatric divergence, where new species arise from geographically isolated populations of the same ancestral species. In contrast, the possibility of sympatric speciation (in which new species arise without geographical isolation) has often been dismissed, partly because of theoretical difficulties. Most previous models analysing sympatric speciation concentrated on particular aspects of the problem while neglecting others. Here we present a model that integrates a novel combination of different features and show that sympatric speciation is a likely outcome of competition for resources. We use multilocus genetics to describe sexual reproduction in an individual-based model, and we consider the evolution of assortative mating (where individuals mate preferentially with like individuals) depending either on an ecological character affecting resource use or on a selectively neutral marker trait. In both cases, evolution of assortative mating often leads to reproductive isolation between ecologically diverging subpopulations. When assortative mating depends on a marker trait, and is therefore not directly linked to resource competition, speciation occurs when genetic drift breaks the linkage equilibrium between the marker and the ecological trait. Our theory conforms well with mounting empirical evidence for the sympatric origin of many species.

Animals↗

Molecular insight into patterns of colony composition and paternity in the common mole-rat Cryptomys hottentotus hottentotus.

We report the discovery of intraspecific variation in both colony composition and patterns of paternity in two populations of the social common mole-rat Cryptomys hottentotus hottentotus. These two populations represent the mesic and arid habitat extremes of the species' broad ecological range in South Africa. Until recently colonies of the common mole-rat were thought to consist of familial groups whereby all colony members were the offspring of a monogamous reproductive pair. The remaining colony members were thought to forego reproduction until both social and ecological conditions favoured dispersal and opportunities for independent outbreeding. Results from genetic assignment tests using microsatellite markers indicate that while colony composition is dominated by familial groups, colonies within both populations included both adult and subadult foreign conspecifics. Analysis of parentage reveals that the social organization of C. h. hottentotus is not that of strict monogamy; paternity of offspring was not assigned consistently to the largest, most dominant male within the colony. Moreover, a number of significantly smaller males were found to sire offspring, suggesting a sneak-mating strategy by subordinate within-colony males. Extra-colony extra-pair paternity (ECP) was also found to characterize C. h. hottentotus colonies, occurring with similar frequencies in both habitats. Both dominant established breeding males and subordinate males were identified as siring young in nonsource colonies. Furthermore, established breeding males were found to sire extra-colony young in the same season as siring young within their source colonies. We discuss the significance of these results within the context of the divergent ecological regimes characterizing the two sites and observe that our results revisit the accuracy of using behavioural and morphological characters, which have structured the basis of our understanding of the behavioural ecology of this species, as indicators of breeding status in mark-recapture studies.

Animals↗

Capture techniques and morphological measurements of the mona monkey (Cercopithecus mona) on the island of Grenada, West Indies.

Morphological measurements were collected from 12 wild and 12 captive mona monkeys (Cercopithecus mona) on the Caribbean island of Grenada. Mona monkeys were introduced to Grenada from Africa approximately 200 to 300 years ago during the slave trade era. Wild monkeys were captured using either 1) a baited treadle-door trap and anesthetic-filled darts fired from a blowpipe, or 2) rifle-fired anesthetic-filled darts. All wild monkeys were released back into the forest after capture and were seen with their original groups within 24 hours of release. Captive monkeys were anesthetized using blowpipe-fired darts. A Ketaset/Rompun mixture was the most effective anesthetic for wild monkeys while Ketaset alone was suitable for captive monkeys. Responses to and recovery times from both drugs varied among individuals. Data on eight linear body measurements, canine length, testicle size, and weight were collected from all monkeys. Adult monkeys were significantly sexually dimorphic across all measurements. Mean adult male weight (mean = 4.7, SD = 0.9, n = 13) was almost twice that of adult females (mean = 2.8, SD = 0.8, n = 7). No significant differences in weight or measurements were found between adult wild and captive males. Preliminary comparisons with morphometrics for African C. mona from the literature showed the upper limit of Grenada mona body length and weight to be smaller than that of African monas for both sexes. These differences may be due to genetic divergence, ecological adaptation, inter-African geographic variation, and/or small sample sizes.

Africa↗

Sexual isolation among widely distributed populations of Drosophila immigrans.

Six strains of the cosmopolitan species D. immigrans from the Australian life zone plus one from the USA, show weak sexual isolation and more rarely sexual selection. Levels of sexual isolation cannot be related to geographic distances. Assortative mating may have evolved as a byproduct of ecological divergence.

Animals↗

Germination characteristics of native Japanese dandelion autopolyploids and their putative diploid parent species.

Germination characteristics of native Japanese Taraxacum lineages of Taraxacum platycarpum (diploid), T. venustum (triploid and tetraploid), and T. albidum (pentaploid) have been studied at different temperatures. Taraxacum platycarpum ssp. hondoense is the putative diploid parent of T. venustum. Diploid T. platycarpum ssp. hondoense and the polyploids T. venustum and T. albidum are found in different areas of Japan, and distribution differences may reflect divergent ecological and physiological traits among ploidy levels. In this study, to prevent mixing of seeds of different polyploidy we used flow cytometry to examine the ploidy level of the plants from which seeds were collected. Results from seed-germination experiments showed that dependence on temperature of final percentage germination was qualitatively similar for both autopolyploids T. venustum and diploids T. platycarpum--germination was suppressed at high and low temperatures. It was also shown that seed germination of autopolyploids was suppressed more than that of the ancestral diploid at low temperatures and that seed germination for polyploids was higher than for the diploid. Threshold variations at low temperatures might affect the distribution of native dandelions. Taraxacum venustum, which occurs in cool climates, might have developed a distinctly lower germination threshold at low temperatures whereas T. albidum, which is native to warm climates, might have developed an adaptive threshold at high temperatures.

Adaptation, Physiological↗

Hybridization and the colonization of novel habitats by annual sunflowers.

Although invasive plant species often have a hybrid ancestry, unambiguous evidence that hybridization has stimulated the evolution of invasive behaviors has been difficult to come by. Here, we briefly review how hybridization might contribute to the colonization of novel habitats, range expansions, and invasiveness and then describe work on hybrid sunflowers that forges a direct link between hybridization and ecological divergence. We first discuss the invasion of Texas by the common sunflower and show that the introgression of chromosomal segments from a locally adapted species may have facilitated range expansion. We then present evidence that the colonization of sand dune, desert floor, and salt marsh habitats by three hybrid sunflower species was made possible by selection on extreme or "transgressive" phenotypes generated by hybridization. This body of work corroborates earlier claims regarding the role of hybridization in adaptive evolution and provides an experimental and conceptual framework for ongoing studies in this area.

Biological Evolution↗

Discordant temporal and geographic patterns in maternal lineages of eastern North American frogs, Rana catesbeiana (Ranidae) and Pseudacris crucifer (Hylidae).

Whether the Pleistocene has had a disproportionate impact on the recent diversification of temperate species, or played a lesser role in a more protracted process, has been a prominent evolutionary debate for the past decade. We used cytochrome b sequences to reconstruct the evolutionary histories of two widely co-distributed, and ecologically divergent frogs (Rana catesbeiana and Pseudacris crucifer) to examine the role of the Pleistocene in structuring these species. Results for R. catesbeiana reflect a pattern of allopatric fragmentation, likely in Coastal Plain refugia on either side of the Mississippi River dating to the mid to early Pleistocene. In contrast, P. crucifer contains numerous divergent lineages, including one west of the Mississippi River in the Interior Highlands, and in the east, multiple lineages that likely expanded from a number of southern Appalachian refugia with lineage sundering originating in the late Pliocene. Large-scale phylogeographic comparisons between these and other eastern North American species reflect both congruent and independent patterns of diversification, possibly reflecting the relative importance of dispersal ability and habitat associations. Although intra-lineage diversification has been structured by repeated Pleistocene glaciations, lineage sundering likely dates at least to the Pliocene in most (but not all) northern temperate amphibian and reptile species studied to date.

Animals↗

Recovery of fish stocks in the Seto Inland Sea.

The total amount of fisheries' yield in the Seto Inland Sea in 1999 was 571,000 MT, consisting of 256,000 MT by fishing and 315,000 MT from aquaculture. About 40,000 people engaged in the fishing and aquaculture industries, earned 129 billion yen (1100 million US$) by fishing and 89 billion yen (770 million $US) by aquaculture. The averaged annual catch for the Seto Inland Sea by fishing was 13 MT/km(2). Division into time periods in terms of eutrophication levels can be made: before 1960 when red sea bream were abundant with ecological divergence (before eutrophication), from 1960 to 1990 when the biomass of anchovy was large (during eutrophication), and after 1990 when the jellyfishes were abundant (excessive eutrophication or high N:P ratio). The fish production will decrease in the sea of jellyfishes. Actually, the amount of catch was 462,000 MT in 1982 which decreased 265,000 MT in 1993, corresponding to 43% in twelve years, then keeping the same level. A big reduction was seen in the catches of the spotlined sardine, anchovy, Spanish mackerel, tiger puffer, short-necked clam, sea cucumber and others. The tiger puffer and Spanish mackerel were abundant as predators in the sea of anchovy. The biomass of anchovy was at its maximum in 1986 and decreased to less than one third in 1996. The stocks of tiger puffer and Spanish mackerel greatly decreased because of the higher fishing pressure compared to the anchovy stock. The fishing power of individual fisheries targeting on the tiger puffer and Spanish mackerel increased substantially when fishing vessel and fishing gear improved, resulting in an excessive fishing effort. A large quantity of small immature fishes is usually caught in the Seto Inland Sea, resulting in growth and/or recruitment overfishing for many species. Hence, it is necessary to promote management of the fisheries so as not to reduce the fish stocks, and to allow the Seto Inland Sea to return from being a sea of jellyfishes to a sea of anchovy, with decreased eutrophication levels.

Animals↗