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Homoploid Hybrid Speciation in a Marine Pelagic Fish.

Homoploid hybrid speciation (HHS) is an enigmatic evolutionary process where new species arise through hybridisation of divergent lineages without changes in chromosome number. Although increasingly documented in various taxa and ecosystems, convincing cases of HHS in marine fishes have been lacking. This study presents a possible case of HHS in a pelagic marine fish based on comprehensive genomic, morphological, and ecological analyses. Population genomics, species tree estimation, and tests of introgression and admixture identified three sympatric clusters in Megalaspis cordyla in the western Pacific and the admixed nature of one cluster between the others. Moreover, model-based demographic inference favoured a hybrid speciation scenario over introgression for the origin of the admixed cluster. While contemporary gene flow suggested partial reproductive isolation, examination of occurrence data and ecologically relevant morphological characters suggested ecological differences between the clusters, potentially contributing to the reproductive isolation and niche partitioning in sympatry. The clusters are also morphologically distinguishable and thus can be taxonomically recognised as separate species. The hybrid cluster is restricted to the coasts of Taiwan and Japan, where all three clusters coexist. The parental clusters are additionally found in lower latitudes, where they display non-overlapping distributions. Given the geographical distributions, estimated times of species formation, and patterns of historical demographic changes, we propose that the Pleistocene glacial cycles were the primary driver of HHS in this system. We also develop an ecogeographic model of HHS in marine coastal ecosystems, including a novel hypothesis to explain the initial stages of HHS.

Animals↗

Drosophila flies in "Evolution Canyon" as a model for incipient sympatric speciation.

The genetic basis of population divergence leading to adaptive radiation and speciation is a major unresolved problem of evolutionary biology. Molecular elucidation of "speciation genes" advanced recently, yet it remains without clear identification of the gene complexes participating in reproductive isolation between natural populations, particularly, in sympatry. Genetic divergence was discovered between Drosophila melanogaster populations inhabiting ecologically contrasting, opposite slopes in "Evolution Canyon" (EC), Mt. Carmel, Israel. Interslope migration of flies is easy and verified. Nevertheless, significant interslope D. melanogaster population divergence was established at EC involving habitat choice, mate choice, thermal and drought tolerances, adaptive genes, and mobile elements. Parallel patterns of stress tolerance, habitat choice, and mate choice were demonstrated in Drosophila simulans at EC, although on a smaller scale. However, some tests for interslope genetic differentiation in Drosophila, derived from the opposite EC slopes, gave somewhat controversial results. Here we present new empirical data on interslope genetic divergence of Drosophila at EC, and summarize previous supporting and controversial results. We suggest that Drosophila populations at EC represent a rare example, demonstrating how selection overrides migration, and propose an ad hoc ecological model of incipient sympatric divergence.

Adaptation, Biological↗

Evolutionary and ecological causes of the latitudinal diversity gradient in hylid frogs: treefrog trees unearth the roots of high tropical diversity.

Why are there more species in the tropics than in temperate regions? In recent years, this long-standing question has been addressed primarily by seeking environmental correlates of diversity. But to understand the ultimate causes of diversity patterns, we must also examine the evolutionary and biogeographic processes that directly change species numbers (i.e., speciation, extinction, and dispersal). With this perspective, we dissect the latitudinal diversity gradient in hylid frogs. We reconstruct a phylogeny for 124 hylid species, estimate divergence times and diversification rates for major clades, reconstruct biogeographic changes, and use ecological niche modeling to identify climatic variables that potentially limit dispersal. We find that hylids originated in tropical South America and spread to temperate regions only recently (leaving limited time for speciation). There is a strong relationship between the species richness of each region and when that region was colonized but not between the latitudinal positions of clades and their rates of diversification. Temperature seasonality seemingly limits dispersal of many tropical clades into temperate regions and shows significant phylogenetic conservatism. Overall, our study illustrates how two general principles (niche conservatism and the time-for-speciation effect) may help explain the latitudinal diversity gradient as well as many other diversity patterns across taxa and regions.

Animals↗

Modeling the effect of algal dynamics on arsenic speciation in Lake Biwa.

Algae reduce and methylate arsenate and the end product of the reaction is correlated to their growth rate. At slow growth rates, dimethylarsinate (DMA) is produced, and at fast growth rates arsenite (As(III)) is produced. Previous work has linked this phenomenon to the phosphorus luxury uptake mechanism of algae, and a model was developed for the process (Hellweger et al. Limnol. Oceanogr. 2003, 48, 2275). This paper presents the integration of that process model for arsenic transformation by algae into a full ecological model and application to Lake Biwa, Japan. The model application allows for a quantitative analysis of the field data, consistent with the process model and the ecological dynamics of the lake. The newly developed ecological model includes a variable phytoplankton composition, which is needed to simulate luxury uptake. This is in contrast to most existing ecological models, which typically assume a fixed "Redfield" composition. The model adequately reproduces the observed ecology of Lake Biwa, including the rapid uptake of phosphate by phytoplankton without immediate growth (luxury uptake) following lake overturn. The model also reproduces the observed arsenic speciation, including the gradual appearance of DMA during the summer and peaks in As(III) concentration at the onset of spring and fall algal blooms.

Arsenic↗

The diversity of parasites.

Parasitism is one of the most successful modes of life displayed by living organisms, as measured by how often it evolved and how many parasitic species are presently in existence. Studying the diversity of parasites is particularly relevant because sympatric diversification may be important in some parasite taxa, and because of the opportunity for independent tests of evolutionary hypotheses in the many separate lineages in which parasitism evolved. Our incomplete knowledge of existing parasite species--the result of a range of phenomena that includes inadequate sampling effort or the lumping of different cryptic species under one name--is not always a major obstacle for the study of parasite diversity. Patterns in the diversity of parasites may be associated with either host or parasite characteristics. The distribution of parasite diversity among host taxa does not simply reflect the species diversity of the host taxa themselves; life history and ecological traits of hosts appear to play important roles. These may determine the likelihood that hosts are colonized by parasite species over evolutionary time. It is not yet clear whether some host traits also favor intrahost speciation and diversification of parasites, and the formation of new parasite species. Certain features of parasites may also be associated with speciation and diversification. Only parasite body size has received much attention; the patterns observed are not greatly different from those of free-living species, with small-bodied parasite taxa being more speciose than related large-bodied taxa. Epidemiological parameters such as the basic reproductive rate of parasites, or R0, can also generate predictions regarding the distribution or evolution of parasite diversity. For instance, parasite taxa characterized by high R0 values may be more speciose than related taxa with lower values of R0; such predictions remain untested. Large-scale biogeographical patterns of diversity have only been well studied for metazoan parasites of marine fish; for these parasites, latitudinal patterns can be explained by effects of temperature on speciation rates and epidemiological variables, though other causes are possible. The emphasis for future research must shift from pattern description to the elucidation of the processes responsible for the structure and diversity of parasite faunas. A better integration of ecological and historical (or phylogenetic) approaches to the study of parasite diversity should make this objective possible.

Animals↗

Socially mediated speciation.

We employ a simple model to show that social selection can lead to prezygotic reproductive isolation. The evolution of social discrimination causes the congealing of phenotypically similar individuals into different, spatially distinct tribes. However, tribal formation is only obtained for certain types of social behavior: altruistic and selfish acts can produce tribes, whereas spiteful and mutualistic behaviors never do. Moreover, reduced hybrid fitness at tribal borders leads to the selection of mating preferences, which then spread to the core areas of the respective tribes. Unlike models of resource competition, our model generates reproductive isolation in an ecologically homogeneous environment. We elaborate on how altruistic acts can lead to reproductive isolation, but also predict that certain types of competition can lead to the speciation effect. Our theory provides a framework for how individual-level interactions mold lineage diversification, with parapatric speciation as a possible end product.

Social Behavior↗

What initiates speciation in passion-vine butterflies?

Studies of the continuum between geographic races and species provide the clearest insights into the causes of speciation. Here we report on mate choice and hybrid viability experiments in a pair of warningly colored butterflies, Heliconius erato and Heliconius himera, that maintain their genetic integrity in the face of hybridization. Hybrid sterility and inviability have been unimportant in the early stages of speciation of these two Heliconius. We find no evidence of reduced fecundity, egg hatch, or larval survival nor increases in developmental time in three generations of hybrid crosses. Instead, speciation in this pair appears to have been catalyzed by the association of strong mating preferences with divergence in warning coloration and ecology. In mate choice experiments, matings between the two species are a tenth as likely as matings within species. F1 hybrids of both sexes mate frequently with both pure forms. However, male F1 progeny from crosses between H. himera mothers and H. erato fathers have somewhat reduced mating success. The strong barrier to gene flow provided by divergence in mate preference is probably enhanced by frequency-dependent predation against hybrids similar to the type known to occur across interracial hybrid zones of H. erato. In addition, the transition between this pair falls at the boundary between wet and dry forest, and rare hybrids may also be selected against because they are poorly adapted to either biotope. These results add to a growing body of evidence that challenge the importance of genomic incompatibilities in the earliest stages of speciation.

Animals↗

Molecular support for vicariance as a source of diversity in rainforest.

The origin of high diversity in tropical rainforests is attributed to divergence amongst relatively mesic, late Pleistocene refuges. To test this hypothesis we analysed sequence variation within the mitochondrial DNA of populations of six rainforest-dwelling vertebrate species (one lizard and five birds) in the Wet Tropical rainforests of northeastern Australia. Vicariance among historical refuges was indicated by geographically congruent variation across a historical climatic barrier previously predicted by modelling. Sequence divergence across the barrier varied widely, being highest in species with lowest vagility and greatest restriction to rainforest. A high altitude, rainforest-restricted species was exceptional in lacking mtDNA variation. These data support the suggested role of vicariance in promoting evolutionary diversity in rainforests but also indicate variation in the timing of vicariance events, with most speciation or divergence events occurring well before the late Pleistocene. The relation between the amount of molecular divergence and current ecology suggests a sequence of isolation events, perhaps of increasing severity, occurring between the late Tertiary and the late Pleistocene.

Animals↗

[Intraspecific variability and sympatry in closely related chigger mite species of the genus Hirsutiella (Acari: Trombiculidae)].

Intraspecific variability in three closely related chigger mite species of the genus Hirsutiella Schluger et Vysotzkaya, 1970, H. steineri (Kepka, 1966), H. llogorensis (Daniel, 1960), and H. alpina Stekolnikov, 2001 has been studied based on materials collected in Caucasus and Turkey. It is established that both H. steineri and H. llogorensis include Western Caucasian and Asian forms, the first one being larger than the second one. Western Caucasian samples of H. steineri are also split into large and small forms. The large form inhabits screes with larvae occurring mainly on snow voles, and the small form inhabits meadows and forests with larvae parasitizing snow voles as well as mice of the genus Apodemus and pine voles. Asian population of H. steineri include small low-mountain form which is hard to distinguish from H. llogorensis. In the light of the new data on variability, morphological border between these two species are specified. Correlations between some characters and altitude above sea level are shown within the Western Caucasian and Asian forms of both species, and also in H. alpina. All the three species can occur together on the same individual hosts, but they have different sets of main hosts and different distribution among biotopes in the area of sympatria. The border between H. steineri and H. llogorensis in the places of sympatria can be indistinct owing to the presence of small ecological forms of H. steineri. Our results give a basis for the construction of alternative hypotheses concerning the processes of speciation in trombiculids. Chiggers species could be formed in allopatric way, on the base of such geographical forms, as Western and Asian forms of the Caucasian Hirsutiella species, but they could also be formed in sympatric way, distributing among neighbouring biotopes (e.g., screes and meadows, or screes and forests), as large and small forms of Western Caucasian H. steineri do.

Animals↗

[Probable origin of the Robertsonian phenomena in domestic mice in Tunisia].

The Robertsonian phenomenon in house mice (Mus musculus domesticus) from Tunisia consists in the presence of only one 22-chromosome Robertsonian race (22Rb) carrying the maximum number of fusions observed until now. The 22Rb populations exclusively occupy urban centers in the Eastern-Central region of Tunisia where standard population with 40-all acrocentric chromosomes (40Std) occur in surrounding neighborhoods and rural environments. In addition to the habitat partition, allozyme and mitochondrial DNA analyses showed that the 22Rb populations were genetically differentiated from the 40Std ones. This differentiation mostly stemmed from an important decrease in genetic variability in the 22Rb populations from the Sahel towns. The extent of morphological, ecological and genetical divergence observed between these chromosomal races in Tunisia is in agreement with the predictions of the chromosomal speciation model of White which advocates that karyotypic differentiation between taxa can lead to their reproductive isolation and independent evolution. Such a process is verified if the Rb process in Tunisia results from local differentiation which is supported by both the genetic and morphological data. However, the hypothesis of an origin by introduction of these mice from another region of Tunisia or from another country cannot be totally dismissed. In this study, an allozymic analysis of mice (22Rb and 40Std) from the geographically distant city of Kairouan was performed. Results showed that 22Rb and 40Std mice from Kairouan shared the same high degree of variability, and were not genetically differentiated. This contrasts with the results registered in the two chromosomal races in the Sahel towns. Such data argue in favor of a local differentiation of the Robertsonian process in Tunisia and suggest that the decrease in variability of the structural nuclear genes in the Sahel 22Rb populations can be related to an introduction from Kairouan into a Sahel locality resulting in a founder effect or followed by a severe bottleneck prior to its dispersion throughout the Sahel region.

Animals↗

Molecular evolutionary genetics of isozymes: pattern, theory, and application.

Isozyme studies at the population genetics-ecology interface conducted at the Institute of Evolution, University of Haifa, during 15 years, 1974-1989, are reviewed in terms of the evidence, theoretical, and practical implications. These studies involve numerous individuals, populations, species, and higher taxa in nature of plants, animals, and humans tested for variation at 15 to 50 primary isozyme loci. The isozyme studies have been conducted mainly in individuals sampled in natural populations at the local, regional, and global levels. Two of the species studied were wild cereals, the progenitors of wheat and barley in the Near East Fertile Crescent. These studies have been complemented by laboratory controlled a priori experimentation of inorganic and organic pollution biology. The human genetics laboratory compared isozyme structure of Jewish and non-Jewish populations. Our results indicate that: (i) isozyme diversity in nature in abundant, at least partly adaptive, and is oriented and maintained primarily by ecological factors. (ii) Natural selection in action is highlighted by stresses involving among others thermal, chemical, and climatic factors. (iii) Speciation can occur with little change in isozyme diversity. (iv) Jews from diverse countries, and in spite of 2,000 years of Diaspora, retain in the frequencies of some isozymes their Near Eastern origins. (v) Wild cereals harbor rich genetic resources exploitable in breeding either directly as adaptive structures, or indirectly as genetic markers for genotypic production of elite agronomic traits. (vi) Isozymes have been utilized as genetic monitors of marine pollution thereby contributing to environmental quality and conservation. (vii) Isozymes can substantially contribute to conservation biology. (viii) Isozymes have been successfully utilized in constructing molecular phylogenies and in revealing new sibling species. (ix) Future theoretical and practical directions of isozyme studies at the protein and DNA levels are outlined.

Animals↗

Natural selection and divergence in mate preference during speciation.

Sexual isolation can evolve due to natural selection against hybrids (reinforcement). However, many different forms of hybrid dysfunction, and selective processes that do not involve hybrids, can contribute to the evolution of sexual isolation. Here we review how different selective processes affect the evolution of sexual isolation, describe approaches for distinguishing among them, and assess how they contribute to variation in sexual isolation among populations of Timema cristinae stick-insects. Pairs of allopatric populations of T. cristinae living on different host-plant species exhibit greater sexual isolation than those on the same host, indicating that some sexual isolation has evolved due to host adaptation. Sexual isolation is strongest in regions where populations on different hosts are in geographic contact, a pattern of reproductive character displacement that is indicative of reinforcement. Ecological costs to hybridization do occur but traits under ecological selection (predation) do not co-vary strongly with the probability of between-population mating such that selection on ecological traits is not predicted to produce a strong correlated evolutionary response in mate preference. Moreover, F1 hybrid egg inviability is lacking and the factors contributing to reproductive character displacement require further study. Finally, we show that sexual isolation involves, at least in part, olfactory communication. Our results illustrate how understanding of the evolution of sexual isolation can be enhanced by isolating the roles of diverse ecological and evolutionary processes.

Adaptation, Biological↗

Inheritance of female mating preference in a sympatric sibling species pair of Lake Victoria cichlids: implications for speciation.

Female mate choice has often been proposed to play an important role in cases of rapid speciation, in particular in the explosively evolved haplochromine cichlid species flocks of the Great Lakes of East Africa. Little, if anything, is known in cichlid radiations about the heritability of female mating preferences. Entirely sympatric distribution, large ecological overlap and conspicuous differences in male nuptial coloration, and female preferences for these, make the sister species Pundamilia pundamilia and P. nyererei from Lake Victoria an ideally suited species pair to test assumptions on the genetics of mating preferences made in models of sympatric speciation. Female mate choice is necessary and sufficient to maintain reproductive isolation between these species, and it is perhaps not unlikely therefore, that female mate choice has been important during speciation. A prerequisite for this, which had remained untested in African cichlid fish, is that variation in female mating preferences is heritable. We investigated mating preferences of females of these sister species and their hybrids to test this assumption of most sympatric speciation models, and to further test the assumption of some models of sympatric speciation by sexual selection that female preference is a single-gene trait. We find that the differences in female mating preferences between the sister species are heritable, possibly with quite high heritabilities, and that few but probably more than one genetic loci contribute to this behavioural speciation trait with no apparent dominance. We discuss these results in the light of speciation models and the debate about the explosive radiation of cichlid fishes in Lake Victoria.

Animals↗

Adverse effects of ecologically relevant dietary mercury exposure in southern leopard frog (Rana sphenocephala) larvae.

Southern leopard frog (Rana sphenocephala) larvae were exposed to experimental diets supplemented with aufwuchs from control and mercury-enriched mesocosms combined in proportions intended to mimic mercury concentrations and speciation in aufwuchs observed from aquatic systems contaminated by atmospheric deposition. Observations on rates of mortality, malformation, and larval growth and development were made for 254 d. Increased incidence of mortality, malformation, and changes in growth and development were observed at concentrations that reflect the highest concentrations expected in the amphibian diet from atmospheric deposition (1,500-3,300 ng Hg/g dry wt). The results of this study are probably more ecologically realistic than results obtained from previous studies of aqueous mercury toxicity and suggest that dietary mercury exposure in habitats contaminated primarily by atmospheric deposition has the potential to cause adverse effects in amphibian larvae.

Animals↗

Molecular phylogenetics of the holly leafminers (Diptera: Agromyzidae: Phytomyza): species limits, speciation, and dietary specialization.

A molecular phylogenetic analysis was conducted to determine relationships and to investigate character evolution in the Phytomyza ilicis group of leafmining flies on hollies (Aquifoliaceae: Ilex). A total of 2207 bp of the mitochondrial cytochrome oxidase I and II genes were sequenced for all known holly leafminers, as well as for several undescribed members of this group. Maximum-parsimony analysis of the sequence data indicates that these leafminers form a monophyletic group with the inclusion of an undescribed leafminer that feeds on the distantly related plant Gelsemium sempevirens (Loganiaceae). Species boundaries of previously known and of undescribed holly leafmining species were confirmed with the molecular data, with one exception. Optimization of variable ecological and morphological characters onto the most parsimonious phylogeny suggests that these traits are evolutionarily labile, requiring multiple instances of convergence and/or reversal to explain their evolutionary history. Speciation in holly leafminers is associated with host shifts and appears to involve colonization of new hosts more often than cospeciation as the hosts diverge. Monophagy is the most common feeding pattern in holly leafminers, and more generalized feeding is inferred to have evolved at least two separate times, possibly as a prelude to speciation.

Animals↗

Habitat avoidance: overlooking an important aspect of host-specific mating and sympatric speciation?

Understanding speciation requires discerning how reproductive barriers to gene flow evolve between previously interbreeding populations. Models of sympatric speciation for phytophagous insects posit that reproductive isolation can evolve in the absence of geographic isolation as a consequence of an insect shifting and ecologically adapting to a new host plant. One important adaptation contributing to sympatric differentiation is host-specific mating. When organisms mate in preferred habitats, a system of positive assortative mating is established that facilitates sympatric divergence. Models of host fidelity generally assume that host choice is determined by the aggregate effect of alleles imparting positive preferences for different plant species. But negative effect genes for avoiding nonnatal plants may also influence host use. Previous studies have shown that apple and hawthorn-infesting races of Rhagoletis pomonella flies use volatile compounds emitted from the surface of fruit as key chemosensory cues to recognize and distinguish between their host plants. Here, we report results from field trials indicating that in addition to preferring the odor of their natal fruit, apple and hawthorn flies, and their undescribed sister species infesting flowering dogwood (Cornus florida), also avoid the odors of nonnatal fruit. We discuss the implications of nonnatal fruit avoidance for the evolutionary dynamics and genetics of sympatric speciation. Our findings reveal an underappreciated role for habitat avoidance as a potential postmating, as well as prezygotic, barrier to gene flow.

Animals↗

Simple, rapid method for direct isolation of nucleic acids from aquatic environments.

Direct isolation of nucleic acids from the environment may be useful in several respects, including the estimation of total biomass, detection of specific organisms and genes, estimations of species diversity, and cloning applications. We have developed a method that facilitates the concentration of microorganisms from aquatic samples and the extraction of their nucleic acids. Natural water samples of 350 to greater than 1,000 ml are concentrated on a single cylindrical filter membrane (type SVGS01015; Millipore Corp., Bedford, Mass.), and cell lysis and proteolysis are carried out within the filter housing. Crude, high-molecular-weight nucleic acid solutions are then drawn off the filter. These solutions can be immediately analyzed, concentrated, or purified, depending on the intended application. The method is simple, rapid, and economical and provides high-molecular-weight chromosomal DNA, plasmid DNA, and speciated RNAs which comigrate with 5S, 16S, and 23S rRNAs. The methods presented here should prove useful in studying both the ecology and the phylogeny of microbes that resist classical culture methods.

Acetates↗