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PubMed2006

Microgeographic evolution of snail shell shape and predator behavior.

Genetic divergence in geographically isolated populations is a prerequisite for allopatric speciation, one of the most common modes of speciation. In ecologically equivalent populations existing within a small, environmentally homogeneous area, an important role for environmentally neutral divergence is often found or inferred. We studied a species complex of conspicuously shaped Opisthostoma land snails on scattered limestone outcrops within a small area of lowland rainforest in Borneo. We used shell morphometrics, mitochondrial and nuclear DNA sequences, and marks of predation to study the factors involved in allopatric divergence. We found that a striking geographic divergence exists in shell morphology, which is partly associated with neutral genetic divergence. We also found geographic differentiation in the behavior of the snails' invertebrate predator and evidence of an evolutionary interaction between aspects of shell shape and predator behavior. Our study shows that adaptation to biotic aspects of the environment may play a more important role in allopatric speciation than previously suspected, even on a geographically very small scale.

Adaptation, Physiological↗
PubMed2006

Ancient trans-Atlantic flight explains locust biogeography: molecular phylogenetics of Schistocerca.

The desert locust (Schistocerca gregaria) has been an important agricultural pest at least since biblical times. Although the ecology, physiology and behaviour of this insect species have been well characterized, its biogeographical origins and evolutionary history are more obscure. Schistocerca gregaria occurs throughout Africa, the Middle East and Western Asia, but all other species in the genus Schistocerca are found in the New World. Because S. gregaria has the capacity for extreme long-distance movement associated with swarming behaviour, dispersal may have played an important role in determining current distribution patterns. Some authors have argued that S. gregaria is the product of an eastward trans-Atlantic dispersal from North America to Africa; others consider it more likely that the New World taxa are the product of westward dispersal from Africa. Here, we present a mitochondrial DNA phylogeny of Schistocerca species that supports the monophyly of New World species (including the Galapagos endemic Halmenus) relative to S. gregaria. In concert with observed patterns of molecular divergence, and in contrast to previous morphological studies, our analysis indicates a single trans-Atlantic flight from Africa to South America, followed by extensive speciation and ecological divergence in the New World.

Africa↗
PubMed1999

Molecular evolution of cytochrome b of subterranean mole rats, Spalax ehrenbergi superspecies, in Israel.

We describe the molecular evolution of cytochrome b of blind subterranean mole rats. We examined 12 individuals for nucleotide differences in the region of 402 base pairs of mitochondrial cytochrome b. Each individual represents a different population from the entire ecological and speciational range of the four chromosomal species in Israel (2n = 52, 54, 58, and 60) belonging to the Spalax ehrenbergi superspecies. Our results indicate the following. (i) There are seven first-position transitional differences, compared to 34 variable third positions, with no observed second-position substitutions. (ii) A maximum of four amino acids differences occurs across the range. (iii) Within-species diversity increases southward. Only 1 autoapomorphic substitution characterizes either 2n = 52 or 2n = 54, but 6-11 substitutions characterize 2n = 58, and 9-13 substitutions characterize 2n = 60. (iv) Both parsimony and maximum-likelihood trees suggest two monophyletic groups: (a) 2n = 52 and 54, and (b) 2n = 58 and 60, as identified earlier by other protein and DNA markers. (v) Mitochondrial cytochrome b heterogeneity is significantly correlated with climatic factors (rainfall) and biotic factors (body size and allozymes). We hypothesize that two selective regimes direct cytochrome b evolution in the S. ehrenbergi superspecies: (i) purifying selection in the flooded, mesic, hypoxic northern range of 2n = 52 and 54 and (ii) diversifying selection in the climatically spatiotemporal, xeric, and variable southern range of 2n = 58 and 60. Thus, the molecular evolution of mitochondrial cytochrome b in S. ehrenbergi is explicable by opposite selective stresses across the range of S. ehrenbergi in Israel, associated with the ecological adaptive radiation of the complex.

Amino Acids↗
PubMed2005

Comparative landscape genetics and the adaptive radiation of Darwin's finches: the role of peripheral isolation.

We use genetic divergence at 16 microsatellite loci to investigate how geographical features of the Galápagos landscape structure island populations of Darwin's finches. We compare the three most genetically divergent groups of Darwin's finches comprising morphologically and ecologically similar allopatric populations: the cactus finches (Geospiza scandens and Geospiza conirostris), the sharp-beaked ground finches (Geospiza difficilis) and the warbler finches (Certhidea olivacea and Certhidea fusca). Evidence of reduced genetic diversity due to drift was limited to warbler finches on small, peripheral islands. Evidence of low levels of recent interisland migration was widespread throughout all three groups. The hypothesis of distance-limited dispersal received the strongest support in cactus and sharp-beaked ground finches as evidenced by patterns of isolation by distance, while warbler finches showed a weaker relationship. Support for the hypothesis that gene flow constrains morphological divergence was only found in one of eight comparisons within these groups. Among warbler finches, genetic divergence was relatively high while phenotypic divergence was low, implicating stabilizing selection rather than constraint due to gene flow. We conclude that the adaptive radiation of Darwin's finches has occurred in the presence of ongoing but low levels of gene flow caused by distance-dependent interisland dispersal. Gene flow does not constrain phenotypic divergence, but may augment genetic variation and facilitate evolution due to natural selection. Both microsatellites and mtDNA agree in that subsets of peripheral populations of two older groups are genetically more similar to other species that underwent dramatic morphological change. The apparent decoupling of morphological and molecular evolution may be accounted for by a modification of Lack's two-stage model of speciation: relative ecological stasis in allopatry followed by secondary contact, ecological interactions and asymmetric phenotypic divergence.

Adaptation, Biological↗
PubMed2007

Genetic transformation of Arabidopsis lyrata: specific expression of the green fluorescent protein (GFP) in pistil tissues.

Arabidopsis thaliana has become widely used as a model system for plant biology. Recent phylogenetic studies led to a severe revision of the systematic relationships across species of the Brassicaceae family. This provided an opportunity to examine close relatives of A. thaliana and to study the function and molecular evolution of genes that play roles in ecology and speciation. In this context, developing tools to genetically transform "non-model plants" appears as a major issue to ascertain gene function. Here, we report a method to transform A. lyrata, one of the closest relatives of A. thaliana.

Arabidopsis↗
PubMed2003

Methodologies for assessing exposure to metals: speciation, bioavailability of metals, and ecological host factors.

Host factors play a role in the bioavailability of metals, making it critical to understand their nature and how to measure them, as well as how to measure bioavailability with respect to host factors. The host factors that are critical to consider during all phases of bioavailability studies are age, gender, size, genetic characteristics, behavior (food chain considerations), and interactions between all of them. Some of these vulnerabilities are unique to individuals, populations, species, or communities. There are many interactions between and among metals, the species of metals, and the physical environment (pH, salinity). Some factors enhance uptake and absorption, whereas others moderate it. Moreover, some metals have greater effects on invertebrate organisms, whereas other metals (or species thereof) affect vertebrates more strongly. Fish and wildlife are useful as sentinel species and bioindicators because they can help us understand the risk to the organisms themselves, to the ecosystem, and to humans.

Animals↗
PubMed2005

Genetic differentiation and estimation of effective population size and migration rates in two sympatric ecotypes of the marine snail Littorina saxatilis.

On exposed rocky shores in Galicia (northwest Spain), a striking polymorphism exists between two ecotypes (RB and SU) of Littorina saxatilis that occupy different levels of the intertidal zone and exhibit an incomplete reproductive isolation. The setting has been suggested to represent ongoing sympatric speciation by ecological adaptation of the two ecotypes to their respective habitats. In this article we address whether or not the ecotypes have developed their own population structures in response to the rigors of their corresponding environments and life histories. We analyzed four to five allozymic loci from three surveys of the same sites, spanning a 14-year period. An experimental design including three localities with two transects per locality and three shore levels allowed studying temporal and spatial population structure and estimation of effective population sizes (N(e)), neighborhood sizes (N(n)), and migration rates (m). Genetic differentiation was significantly lower in RB populations (theta(ST) = 0.067) than in SU ones (theta(ST) = 0.124). Mean estimates of N(e), N(n), and m did not differ significantly between ecotypes, but local ecotype differences in migration between the two closest localities (larger migration rates in RB than in SU populations) could explain the pattern in population differentiation.

Animals↗
PubMed2004

Evolutionary dynamics of biological games.

Darwinian dynamics based on mutation and selection form the core of mathematical models for adaptation and coevolution of biological populations. The evolutionary outcome is often not a fitness-maximizing equilibrium but can include oscillations and chaos. For studying frequency-dependent selection, game-theoretic arguments are more appropriate than optimization algorithms. Replicator and adaptive dynamics describe short- and long-term evolution in phenotype space and have found applications ranging from animal behavior and ecology to speciation, macroevolution, and human language. Evolutionary game theory is an essential component of a mathematical and computational approach to biology.

Adaptation, Biological↗
PubMed1997

Mass Spawning by Green Algae on Coral Reefs

Predawn episodes of mass spawning by green algae (up to nine species in five genera on a single morning) intermittently cloud Caribbean waters. Species- and sex-specific bouts of anisogamous gamete release occurred synchronously and predictably on a given morning, with closely related species spawning at different times. Algal sexual reproduction was seasonal, but, unlike the mass-spawning behavior of other sessile marine organisms, showed no lunar or tidal cycling. The discovery of mass-spawning behavior by these algae has important implications for future studies of the reproductive ecology and speciation of a vital, yet poorly understood, component of the coral reef community.

Journal Article↗
PubMed2002

Molecular systematics of Anopheles: from subgenera to subpopulations.

The century-old discovery of the role of Anopheles in human malaria transmission precipitated intense study of this genus at the alpha taxonomy level, but until recently little attention was focused on the systematics of this group. The application of molecular approaches to systematic problems ranging from subgeneric relationships to relationships at and below the species level is helping to address questions such as anopheline phylogenetics and biogeography, the nature of species boundaries, and the forces that have structured genetic variation within species. Current knowledge in these areas is reviewed, with an emphasis on the Anopheles gambiae model. The recent publication of the genome of this anopheline mosquito will have a profound impact on inquiries at all taxonomic levels, supplying better tools for estimating phylogeny and population structure in the short term, and ultimately allowing the identification of genes and/or regulatory networks underlying ecological differentiation, speciation, and vectorial capacity.

Animals↗
PubMed2003

Evaluation of the courtship and of the hybrid male sterility among Drosophila buzzatii cluster species (Diptera, Drosophilidae).

In the Drosophila repleta group the establishment of subgroups and complexes made on the basis of morphological and cytological evidences is supported by tests of reproductive isolation. Among species in the repleta group, the buzzatii cluster, due to its polymorphism and polytipism, is an excellent material for ecological and speciation studies. Some interspecific crosses involving Drosophila seriema, Drosophila sp. B, D. koepferae and D. buzzatii strains were completely sterile while others involving strains from these species produced F1 hybrids that did not yield F2. In the present work, data on courtship duration and copula occurrence obtained in the analysis of flies from parental sterile crosses and on spermatozoon mobility observed in F1 hybrids that did not yield F2 are presented. Copula did not occur during one hour of observation and the spermatozoon also did not show mobility at any of the analyzed stages (3, 7, 9 and 10 days old). There was a high variation in courtship average duration and in the percentage of males that courted the females. The reproductive isolation mechanisms indicated by these observations were pre and post-zygotic, as supported by the absence of copula and male sterility. Data obtained also showed the occurrence of different degrees of reproductive compatibility among the strains classified as the same species but from distinct geographic localities.

Animals↗
PubMed2006

Comment on "Neutral ecological theory reveals isolation and rapid speciation in a biodiversity hot spot".

Latimer et al. (Reports, 9 September 2005, p. 1722) used an approximate likelihood function to estimate parameters of Hubbell's neutral model of biodiversity. Reanalysis with the exact likelihood not only yields different estimates but also shows that two similar likelihood maxima for very different parameter combinations can occur. This reveals a limitation of using species abundance data to gain insight into speciation and dispersal.

Animals↗
PubMed2005

Neutral ecological theory reveals isolation and rapid speciation in a biodiversity hot spot.

South Africa's Mediterranean-climate fynbos shrubland is a hot spot of species diversity, but its diversity patterns contrast strongly with other high-diversity areas, including the Amazon rain forest. With its extremely high levels of endemism and species turnover, fynbos is made up of dissimilar local communities that are species-rich but relatively poor in rare species. Using neutral ecological theory, we show that the relative species-abundance distributions in fynbos can be explained by migration rates that are two orders of magnitude lower than they are in tropical rain forests. Speciation rates, which are indexed by the "biodiversity parameter" Theta, are estimated to be higher than they are in any previously examined plant system.

Bayes Theorem↗
PubMed2001

Linking ecological impact to metal concentrations and speciation: a microcosm experiment using a salt marsh meiofaunal community.

Microcosm experiments addressed the impact of a mixture of Cu, Cr, Cd, Pb, and Hg at three concentrations after 36 h, 12 d, and 30 d on a meiofauna-dominated salt marsh community. In addition to analyzing effects on meiofaunal abundances, the study quantified the sediment metal concentrations of all five metals and pore-water concentrations, speciation, and ligand complexation of Cu. Abundances of deposit feeders such as the polychaete Streblospio benedicti, gastropods, and bivalves were impacted at lower metal concentrations than the mainly algal-feeding copepods, ostracods, and nematodes. We suggest that this might be due to bulk ingestion of metal-contaminated sediments resulting in relatively higher metal exposure in the deposit feeders than in the other, nondeposit feeding taxa. Copepod and ostracod abundances decreased only in the highest metal treatment, where levels of inorganic Cu ([Cu']) in pore waters were similar to levels associated with both acute and subacute toxicity in published in vivo toxicity studies of marine copepods. The higher metal treatments yielded disproportionately higher pore-water [Cu] compared with sediment [Cu], suggesting saturation of sediment-associated ligands with increased additions of Cu. Similarly, the higher metal treatments appeared to reach saturation of the organic Cu ligands, with the excess pore-water [Cu] present in the more toxic, inorganic species of Cu. Acid-volatile sulfide (AVS) concentrations at sediment horizons inhabited by meiofauna were low and AVS was not considered a significant metal ligand at these depths. Since meiofauna are predominantly associated with oxic surface sediments, it is doubtful that AVS is a major factor controlling availability of free metal for exposure to these taxa.

Animals↗
PubMed2000

Incipient speciation in sympatric Nicaraguan crater lake cichlid fishes: sexual selection versus ecological diversification.

The growing body of empirical evidence for sympatric speciation has been complemented by recent theoretical treatments that have identified evolutionary conditions conducive to speciation in sympatry. The Neotropical Midas cichlid (Amphilophus citrinellum) fits both of the key characteristics of these models, with strong assortative mating on the basis of a colour polymorphism coupled with trophic and ecological differentiation derived from a polymorphism in their pharyngeal jaws. We used microsatellite markers and a 480 bp fragment of the mitochondrial DNA control region to study four polymorphic populations of the Midas cichlid from three crater lakes and one large lake in Nicaragua in an investigation of incipient sympatric speciation. All populations were strongly genetically differentiated on the basis of geography. We identified strong genetic separation based on colour polymorphism for populations from Lake Nicaragua and one crater lake (Lake Apoyo), but failed to find significant genetic structuring based on trophic differences and ecological niche separation in any of the four populations studied. These data support the idea that sexual selection through assortative mating contributes more strongly or earlier during speciation in sympatry than ecological separation in these cichlids. The long-term persistence of divergent cichlid ecotypes (as measured by the percentage sequence divergence between populations) in Central American crater lakes, despite a lack of fixed genetic differentiation, differs strikingly from the patterns of extremely rapid speciation in the cichlids in Africa, including its crater lakes. It is unclear whether extrinsic environmental factors or intrinsic biological differences, e.g. in the degree of phenotypic plasticity, promote different mechanisms and thereby rates of speciation of cichlid fishes from the Old and New Worlds.

Animals↗
PubMed1994

Chromosomal speciation and adaptive radiation of mole rats in Asia Minor correlated with increased ecological stress.

The evolutionary forces causing chromosomal speciation and adaptation are still enigmatic. Here we tested the Israel evolutionary model of positive association of diploid chromosome number (2n) and genetic diversity with aridity stress in subterranean mole rats, on a 30-times-larger scale in Asia Minor. We analyzed both karyotype and allozyme diversity across Turkey, based on 37 allozymic loci in 20 localities of the Spalax leucodon and 4 localities of the Spalax ehrenbergi superspecies. We found extensive chromosomal speciation in S. leucodon (2n = 38, 40, 50, 54, 60, and 62) and in S. ehrenbergi (2n = 52, 56, and 58), presumably representing from 14 to > 20 additional biological species. Genetic diversity indices were low, but, like the chromosome number (2n), positively correlated with aridity stress, increasing centripetally from the periphery toward geologically young, arid, and climatically unpredictable central Anatolia. Nei's genetic distance D across all populations averaged 0.174 (range 0.002-0.422), supporting, combined with 2n and ecogeography, the biological species status of most tested populations. Chromosome evolution is the basis of speciation and adaptation in Spalax; it provides both postmating reproductive isolation, as well as higher levels of recombination with increased 2n. A mathematical model shows that a Robertsonian fission of a single metacentric considerably increases haplotype diversity. This haplotype diversity may contribute to population adaptation to climatic stress and ecological unpredictability in space and time. The increase in diversity corroborates the nichewidth genetic-variation hypothesis.

Animals↗
PubMed2003

Speciation along environmental gradients.

Traditional discussions of speciation are based on geographical patterns of species ranges. In allopatric speciation, long-term geographical isolation generates reproductively isolated and spatially segregated descendant species. In the absence of geographical barriers, diversification is hindered by gene flow. Yet a growing body of phylogenetic and experimental data suggests that closely related species often occur in sympatry or have adjacent ranges in regions over which environmental changes are gradual and do not prevent gene flow. Theory has identified a variety of evolutionary processes that can result in speciation under sympatric conditions, with some recent advances concentrating on the phenomenon of evolutionary branching. Here we establish a link between geographical patterns and ecological processes of speciation by studying evolutionary branching in spatially structured populations. We show that along an environmental gradient, evolutionary branching can occur much more easily than in non-spatial models. This facilitation is most pronounced for gradients of intermediate slope. Moreover, spatial evolutionary branching readily generates patterns of spatial segregation and abutment between the emerging species. Our results highlight the importance of local processes of adaptive divergence for geographical patterns of speciation, and caution against pitfalls of inferring past speciation processes from present biogeographical patterns.

Animals↗