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A test of alternative models of diversification in tropical rainforests: ecological gradients vs. rainforest refugia.

Comparison of mitochondrial and morphological divergence in eight populations of a widespread leaf-litter skink is used to determine the relative importance of geographic isolation and natural selection in generating phenotypic diversity in the Wet Tropics Rainforest region of Australia. The populations occur in two geographically isolated regions, and within each region, in two different habitats (closed rainforest and tall open forest) that span a well characterized ecological gradient. Morphological differences among ancient geographic isolates (separated for several million years, judging by their mitochondrial DNA sequence divergence) were slight, but morphological and life history differences among habitats were large and occurred despite moderate to high levels of mitochondrial gene flow. A field experiment identified avian predation as one potential agent of natural selection. These results indicate that natural selection operating across ecological gradients can be more important than geographic isolation in similar habitats in generating phenotypic diversity. In addition, our results indicate that selection is sufficiently strong to overcome the homogenizing effects of gene flow, a necessary first step toward speciation in continuously distributed populations. Because ecological gradients may be a source of evolutionary novelty, and perhaps new species, their conservation warrants greater attention. This is particularly true in tropical regions, where most reserves do not include ecological gradients and transitional habitats.

Animals↗

Ecological divergence promotes the evolution of cryptic reproductive isolation.

Speciation can involve the evolution of 'cryptic' reproductive isolation that occurs after copulation but before hybrid offspring are produced. Because such cryptic barriers to gene exchange involve post-mating sexual interactions, analyses of their evolution have focused on sexual conflict or traditional sexual selection. Here, we show that ecological divergence between populations of herbivorous walking sticks is integral to the evolution of cryptic reproductive isolation. Low female fitness following between-population mating can reduce gene exchange between populations, thus acting as a form of cryptic isolation. Female walking sticks show reduced oviposition rate and lower lifetime fecundity following between-population versus within-population mating, but only for mating between populations using different host-plant species. Our results indicate that even inherently sexual forms of reproductive isolation can evolve as a by-product of ecological divergence and that post-mating sexual interactions do not necessarily evolve independently of the ecological environment.

Animals↗

Genetic structure and evolutionary history of a diploid hybrid pine Pinus densata inferred from the nucleotide variation at seven gene loci.

Although homoploid hybridization is increasingly recognized as an important phenomenon in plant evolution, its evolutionary genetic mechanisms are poorly documented and understood. Pinus densata, a pine native to the Tibetan Plateau, represents a good example of a homoploid hybrid speciation facilitated by adaptation to extreme environment and ecological isolation from the parents. Its ecologically and reproductively stabilized nature offers excellent opportunity for studying genetic processes associated with hybrid speciation. In this study, we investigated the levels and patterns of nucleotide variation in P. densata and its putative parents. Haplotype composition, gene genealogies, and the levels and patterns of nucleotide variation gave further support to the hybrid nature of P. densata. Allelic history, as revealed by our data, suggests the ancient nature of the hybrid preceding elevation of the Tibetan Plateau. We detected more deviations from neutrality in P. densata than in the parental species. Thus, at least some of the evolutionary forces that have shaped the genetic variation in P. densata are likely to be different from those acting upon parental species. We speculate that when populations of P. densata invaded new territories, they had elevated rates of response to selection in order to develop traits that help them to survive and adapt in the new environments.

Diploidy↗

Parasite extinction and colonisation and the evolution of parasite communities: a simulation study.

We determined what evolutionary processes influence the likelihood of detecting an effect of host ecological characteristics on the richness of parasite communities in comparative analyses among related host species. We used a mathematical model to generate phylogenies of hosts in which parasite communities varied over evolutionary time as parasite species were either gained or lost during host speciation events. Gain or loss of parasites were stochastic and could either be strongly, moderately, weakly, or not, affected by host ecological characteristics. The model was evaluated over this range of effects of host ecology, and for various mean probabilities of parasite gain and loss and various rates of change in host ecological characteristics at speciation events. Our results suggest that phylogenetic effects (the passing of parasite species from mother to daughter host species) are likely to obscure ecological effects (the effect of host body size, diet, habitat, lifespan, etc.) except when the effects of host ecology are strong, and the probabilities of gain or loss of parasites are high, or host ecological characteristics change markedly at speciation events. This outcome was not influenced by the shape of the phylogenetic tree used in the simulations. Sensitivity analysis of our model also shows this result to be robust to a wide range of assumptions and parameter values. Thus, because the composition of parasite communities tends to reflect their ancestry, the effect of host ecology will often be very difficult to detect.

Animals↗

Conservatism of ecological niches in evolutionary time

Theory predicts low niche differentiation between species over evolutionary time scales, but little empirical evidence is available. Reciprocal geographic predictions based on ecological niche models of sister taxon pairs of birds, mammals, and butterflies in southern Mexico indicate niche conservatism over several million years of independent evolution (between putative sister taxon pairs) but little conservatism at the level of families. Niche conservatism over such time scales indicates that speciation takes place in geographic, not ecological, dimensions and that ecological differences evolve later.

Journal Article↗

Strong assortative mating between allopatric sticklebacks as a by-product of adaptation to different environments.

Speciation involves the evolution of reproductive isolation between populations. One potentially important mechanism is the evolution of pre- or postzygotic isolation between populations as a by-product of adaptation to different environments. In this paper, we tested for assortative mating between allopatric stickleback populations adapted to different ecological niches. Our experimental design controlled for interpopulation interactions and non-adaptive explanations for assortative mating. We found that prezygotic isolation was surprisingly strong: when given a choice, the majority of matings occurred between individuals from similar environments. Our results indicate that the by-product mechanism is a potent source of reproductive isolation, and likely contributed to the origin of sympatric species of sticklebacks.

Adaptation, Physiological↗

Quantitative analysis of dental microwear in threespine stickleback: a new approach to analysis of trophic ecology in aquatic vertebrates.

1. The threespine stickleback Gasterosteus aculeatus is an important model organism in studies of genomic and phenotypic evolution, adaptation and speciation. Fossil Gasterosteus offer the potential to test models derived from studies of extant fishes over true evolutionary time-scales. Competition for food resources, for example, plays an important part in stickleback speciation, causing divergence in food gathering traits and ecological character displacement, but it is not possible to test this model in fossils because evidence of diet is almost never preserved. 2. We demonstrate here that quantitative analysis of dental microwear, a technique previously applied only to mammals, provides a reliable guide to the dietary preferences of stickleback. Teeth from stickleback raised under laboratory conditions exhibit microwear patterns that vary systematically according to substrate coarseness and whether fishes feed on Daphnia within the water column, or on chironomid larvae from the bottom. Furthermore, microwear data exhibit a progressive shift in their distribution that tracks differences in experimental feeding treatments. 3. Microwear in wild populations also exhibits a relationship with feeding. In blind assessments of trophic niche based on microwear patterns we were able to correctly assign all but one equivocal population to trophic group. Microwear data from wild stickleback exhibit a shift in distribution comparable with that observed across the range of treatments in the laboratory and these allow populations to be ranked according to the degree to which they approach fully benthic or fully limnetic feeding. 4. Our results demonstrate that microwear has the potential to be a powerful tool in the analysis of fish trophic ecology, particularly in the analysis of species pairs and niche differentiation. It has advantages over the trophic snapshot provided by analysis of stomach contents in that microwear reflects feeding and food preferences over a longer period of time, and can be applied where these data are unavailable. Furthermore, it is applicable to extinct organisms and fossils, allowing the role of trophic ecology, niche partitioning and competition over evolutionary time-scales to be investigated for the first time.

Animals↗

Axes of differentiation in the bower-building cichlids of Lake Malawi.

The 500-1000 cichlid species endemic to Lake Malawi constitute one of the most rapid and extensive radiations of vertebrates known. There is a growing debate over the role natural and sexual selection have played in creating this remarkable assemblage of species. Phylogenetic analysis of the Lake Malawi species flock has been confounded by the lack of appropriate morphological characters and an exceptional rate of speciation, which has allowed ancestral molecular polymorphisms to persist within species. To overcome this problem we used amplified fragment length polymorphism (AFLP) to reconstruct the evolution of species within three genera of Lake Malawi sand-dwelling cichlids that construct elaborate male display platforms, or bowers. Sister taxa with distinct bower morphologies, and that exist in discrete leks separated by only 1-2 m of depth, are divergent in both sexually selected and ecological traits. Our phylogeny suggests that the forces of sexual and ecological selection are intertwined during the speciation of this group and that specific bower characteristics and trophic morphologies have evolved repeatedly. These results suggest that trophic morphology and bower form may be inappropriate characters for delineating taxonomic lineages. Specifically the morphological characters used to describe the genera Lethrinops and Tramitichromis do not define monophyletic clades. Using a combination of behavioural and genetic characters, we were able to identify several cryptic cichlid species on a single beach, which suggests that sand dweller species richness has been severely underestimated.

Animals↗

Allopatric origins of microbial species.

Although allopatric divergence is a well-accepted mechanism of speciation for eukaryotic macro-organisms, the importance of geographical barriers to divergence in microbial populations is a subject of great debate. Do geographically separated populations of micro-organisms diverge independently, or does their structure fit the often quoted Bass-Becking description 'everything is everywhere; the environment selects'? Aided by high-resolution genetic and genomic tools, the search for 'microbial marsupials' has revealed that in fact both are true; some species of micro-organisms demonstrate allopatric divergence, while others do not. This discovery opens the door for comparative analyses, where questions about the differences in evolutionary and ecological mechanisms that drive divergence and speciation in different microbial species can begin to be explored. Investigating these differences in evolutionary mechanisms will greatly enhance interest in, and understanding of, the dynamic processes that create and maintain the vast diversity of the microbial world.

Bacteria↗

Modeling the ecological impact of heavy metals on aquatic ecosystems: a framework for the development of an ecological model.

In this paper, an ecological model is proposed to predict the effects of heavy metals on aquatic ecosystems. The bioavailable concentration of metals and a concept of toxicity strength (TS) are combined. The integrated ecological model relates the transport, distribution and speciation of heavy metals and their toxicity, and the effect of environmental variability on metal toxicity. It also emphasizes the link between physical and chemical processes of heavy metals in rivers and ecological effects. Based on the data obtained from research in the CERP project (Co-operative Ecological Research Project), the ecological impact of heavy metals on the aquatic ecosystem of the Le An River (polluted by heavy metals from a copper mine) was predicted. The results show that the estimated values of toxicity strength for surface water are in agreement with the percentage inhibition for the test organism (P. phosphoreum) and that the predicted ecological effect of polluted sediment is consistent with natural variability in aquatic ecosystems.

Animals↗

Natural selection and sympatric divergence in the apple maggot Rhagoletis pomonella.

In On the Origin of Species, Darwin proposed that natural selection had a fundamental role in speciation. But this view receded during the Modern Synthesis when allopatric (geographic) models of speciation were integrated with genetic studies of hybrid sterility and inviability. The sympatric hypothesis posits that ecological specialization after a host shift can result in speciation in the absence of complete geographic isolation. The apple maggot, Rhagoletis pomonella, is a model for sympatric speciation in progress. Hawthorn (Crataegus spp.) is the native host for R. pomonella in N. Americas. But in the mid-1800s, a new population formed on introduced, domesticated apple (Malus pumila). Recent studies have conferred 'host race' status on apple flies as a potentially incipient species, partially isolated from haw flies owing to host-related adaptation. However, the source of selection that differentiates apple and haw flies is unresolved. Here we document a gene-environment interaction (fitness trade-off) that is related to host phenology and that genetically differentiates the races.

Animals↗

Recent, allopatric, homoploid hybrid speciation: the origin of Senecio squalidus (Asteraceae) in the British Isles from a hybrid zone on Mount Etna, Sicily.

Homoploid hybrid speciation occurs through stabilization of a hybrid segregate (or segregates) isolated by premating and/or postmating barriers from parent taxa. Theory predicts that ecological and spatial isolation are of critical importance during homoploid hybrid speciation, and all confirmed homoploid hybrid species are ecologically isolated from their parents. Until recently, such species have been identified long after they originated, and consequently it has not been possible to determine the relative importance of spatial and ecological isolation during their origin. Here we present evidence for the recent origin (within the past 300 years) of a new homoploid hybrid species, Senecio squalidus (Asteraceae), in the British Isles, following long-distance dispersal of hybrid material from a hybrid zone between S. aethnensis and S. chrysanthemifolius on Mount Etna, Sicily, Italy. Historical records show that such hybrid material from Sicily was introduced to the Oxford Botanic Garden in Britain in the early part of the 18th century and that S. squalidus began to spread from there after approximately 90 years. A survey of randomly amplified polymorphic DNA/intersimple sequence repeats (RAPD/ISSR) marker variation demonstrated that S. squalidus is a diploid hybrid derivative of S. aethnensis and S. chrysanthemifolius that grow at high and low altitudes, respectively, on Mount Etna and that form a hybrid zone at intermediate altitudes. Senecio squalidus contained 11 of 13 RAPD/ISSR markers that were recorded at high frequency in S. chrysanthemifolius but were absent or occurred at low frequency in S. aethnensis, and 10 of 13 markers for which the reverse was true. Bayesian admixture analysis showed that all individuals of S. squalidus surveyed were of mixed ancestry with relatively high mean proportions of ancestry derived from both S. chrysanthemifolius and S. aethnensis (0.644 and 0.356, respectively). We argue that long-distance isolation of hybrid material from its parents on Mount Etna would have helped favor the origin and establishment of S. squalidus in the British Isles, regardless of whether the initial hybrid material introduced to Britain was preadapted to local conditions.

Altitude↗

Arthropods on islands: colonization, speciation, and conservation.

Islands have traditionally been considered to be any relatively small body of land completely surrounded by water. However, their primary biological characteristic, an extended period of isolation from a source of colonists, is common also to many situations on continents. Accordingly, theories and predictions developed for true islands have been applied to a huge array of systems, from rock pools, to single tree species in forests, to oceanic islands. Here, we examine the literature on islands in the broadest sense (i.e., whether surrounded by water or any other uninhabitable matrix) as it pertains to terrestrial arthropods. We categorize islands according to the features they share. The primary distinction between different island systems is "darwinian" islands (formed de novo) and "fragment" islands. In the former, the islands have never been in contact with the source of colonists and have abundant "empty" ecological niche space. On these islands, species numbers will initially increase through immigration, the rate depending on the degree of isolation. If isolation persists, over time species formation will result in "neo-endemics." When isolation is extreme, the ecological space will gradually be filled through speciation (rather than immigration) and adaptive radiation of neo-endemics. Fragment islands are fundamentally different. In these islands, the ecological space will initially be filled as a consequence of connection to the source of colonists prior to insularization. Species numbers will decrease following fragmentation through the process of relaxation. If these islands become more isolated, species will eventually arise through relictualization with the formation of "paleo-endemics." Given sufficient time, this process can result in generic level endemism on ancient fragment islands, a phenomenon well illustrated in Madagascar and New Zealand. Recognizing the distinction between the different kinds of islands is fundamental for understanding emerging patterns on each, in particular speciation, biodiversity (e.g., neo-endemics versus paleo-endemics), and conservation (e.g., naiveté in interactions with alien species).

Animals↗

Francolin phylogenetics: molecular, morphobehavioral, and combined evidence.

The phylogenetics of francolins (Francolinus species) were reassessed by obtaining 660 bp of sequence of the mitochondrial DNA (mtDNA) cytochrome b gene from 20 species, the Common Quail Coturnix coturnix africana, and the Madagascar Partridge Margaroperdix madagarensis. Published sequences of the Japanese Quail C. c. japonica, Alectoris partridges, and the Junglefowl Gallus gallus were also included. Separate analysis of the 200 phylogenetically informative cytochrome b characters and the 25 informative morphobehavioral characters, as well as a combined analysis of molecular and morphobehavioral data, do not support francolin monophyly but provide strong evidence for two previously suggested clades--the quail-francolins (or partridges) and the partridge-francolins (pheasants/francolins). The quail-francolin clade comprises three groups of African francolins and three Asian species that were previously considered more closely related to the partridge-francolins. The partridge-francolin clade, which includes four groups of African francolins, forms a sister group to the Coturnix quails, the Madagascar Partridge, and the Alectoris partridges. The molecular data suggest that the two francolin clades diverged approximately 3-6 MYA. Climatic fluctuations of the past 2.5 MYA may have led to the diversification of the ecologically different francolin species groups and speciation within them.

Animals↗

Genetic population structure as indirect measure of dispersal ability in a Lake Tanganyika cichlid.

Diversification and speciation processes are influenced by intrinsic (ecological specialization, dispersal) and extrinsic (habitat structure and instability) factors, but the effect of ecological characteristics on dispersal is difficult to assess. This study uses mitochondrial control region sequences to investigate the population structure and demographic history of the endemic Lake Tanganyika cichlid Neolamprologus caudopunctatus with a preference for the rock-sand interface along two stretches of continuous, rocky shoreline, and across a sandy bay representing a potential dispersal barrier. Populations along uninterrupted habitat were not differentiated; whereas, the sandy bay separated two reciprocally monophyletic clades. The split between the two clades between 170,000 and 260,000 years BP coincides with a period of rising water level following a major lowstand, and indicates that clades remained isolated throughout subsequent lake level fluctuations. Low long-term effective population sizes were inferred from modest genetic diversity estimates, and may be due to recent population expansions starting from small population sizes 45,000-60,000 years BP. Comparisons with available data from specialized rock-dwelling species of the same area suggest that habitat structure and lake level fluctuations determine phylogeographic patterns on large scales, while fine-scale population structure and demography are modulated by species-specific ecologies.

Africa, Eastern↗

Vicious circles: positive feedback in major evolutionary and ecological transitions.

Evolutionary biologists and ecologists often focus on equilibrium states that are subject to forms of negative feedback, such as optima for phenotypic traits or regulation of population sizes. However, recent theoretical and empirical studies show how positive feedback can be instrumental in driving many of the most important and spectacular processes in evolutionary ecology, including the evolution of sex and genetic systems, mating systems, life histories, complex cooperation in insects and humans, ecological specialization, species diversity, species ranges, speciation and extinction. Taken together, this work suggests that positive feedback is more common than is generally appreciated, and that its self-reinforcing dynamics generate the conditions for changes that might otherwise be difficult or impossible for selection or other mechanisms to achieve. Testing for positive feedback requires analysing each causal link in feedback loops, tracking genetic, character and population-dynamic changes across generations, and elucidating the conditions that can result in self-reinforcing change.

Journal Article↗

Re-assessment of monophyly, evolution of myrmecophytism, and rapid radiation in Neonauclea s.s. (Rubiaceae).

The biologically interesting ant-plant association, myrmecophytism, occurs in ca. 140 of the 11,000 species and 22 of the 630 genera of the coffee family (Rubiaceae). These myrmecophytic Rubiaceae species are predominantly distributed in Southeast Asia, especially the Malesian region, with comparatively few species in mainland Africa and the Neotropics. The mostly Southeast Asian genus Neonauclea s.s is one of the three Rubiaceae genera with extensive radiation of myrmecophytes and also the most speciose genus of the tribe Naucleeae s.l. We perform parsimony phylogenetic analyses of Neonauclea s.s., previously resolved as paraphyletic, and its allied genera using both ETS and ITS sequencing data to test: (1) the paraphyly of Neonauclea s.s.; (2) the phylogenetic relationships within the Ludekia-Myrmeconauclea-Neonauclea complex; and (3) the evolution of myrmecophytism within the complex. The earlier proposed paraphyly of Neonauclea s.s. appears to be the result of the combined effects of parallel substitutions in Metadina trichotoma and the sampled ITS putative pseudogenes of Neonauclea longipedunculata and losses of some synapomorphies of Neonauclea s.s. in the latter. The analyses present strong support for the monophyly of Myrmeconauclea and Neonauclea s.s. and their sister-group relationships. Our findings additionally favor the hypothesis of multiple origins of myrmecophytism in the Bornean Neonauclea, which have independently been exploited by at least three Cladomyrma ant species. Furthermore, we interpret the low levels of variation in both the ETS and ITS sequences as indication of a recent and rapid radiation for Neonauclea s.s. (with 65 species) and a recent and slow radiation for Myrmeconauclea (with three species). We argue that the rapid diversification of Neonauclea s.s. is partly associated with the nature of its fruits and its ability to colonize a wide range of habitats. We postulate that both ecological and geographical events may have been responsible for the radiation of the non-myrmecophytic Neonauclea species. Finally, we argue that the acquisition of the pseudo-multiple fruits and long-tailed seeds has allowed Myrmeconauclea to specialize on rheophytic habitats but its narrow ecological tolerance may have hindered its speciation.

Animals↗

Comparative analysis of morphological traits among Drosophila melanogaster and D. simulans: genetic variability, clines and phenotypic plasticity.

The two sibling cosmopolitan species, Drosophila melanogaster and D. simulans, are able to proliferate under very different climatic conditions. This has resulted in local adaptations, which are often arranged in latitudinal clines. Such clines are documented for body weight, wing and thorax length, sternopleural and abdominal bristle number, ovariole number and thoracic pigmentation. The overall magnitude of geographical differentiation is, however, much less in D. simulans than in D. melanogaster, and latitudinal clines are less pronounced. The fact that natural populations live under different climates raises the problem of interaction between temperature and phenotype. The reaction norms of morphometrical traits have been investigated as a function of growth temperature. The shapes of the response curves vary according to the investigated trait. They are generally curvilinear and can be described by calculating characteristic values after polynomial adjustments. For a given trait, the reaction norms of the two species are similar in their shape, although some significant differences may be observed. Within each species, significant differences are also observed between geographic populations: reaction norms are not parallel and the divergence is better marked when more distant populations (e.g., temperate and tropical) are compared. It thus appears that besides mean trait value, phenotypic plasticity is also a target of natural selection. A specific analysis of wing shape variation according to growth temperature was also undertaken. Reaction norms with different shapes may be observed in various parts of the wing: the major effect is found between the basis and the tip of the wing, but in a similar way in the two species. By contrast, some ratios, called wing indices by taxonomists, may exhibit completely different reaction norms in the two species. For a single developmental temperature (25 degrees C) the phenotypic variability of morphometrical traits is generally similar in the two species, and also the genetic variability, estimated by the intraclass correlation. A difference exists, however, for the ovariole number which is less variable in D. simulans. Variance parameters may vary according to growth temperature, and a detailed analysis was made on wing dimensions. An increase of environmental variability at extreme, heat or cold temperatures, has been found in both species. Opposite trends were, however, observed for the genetic variability: a maximum heritability in D. simulans at middle temperatures, corresponding to a minimum heritability in D. melanogaster. Whether such a difference exists for other traits and in other populations deserves further investigations. In conclusion, morphometrical analyses reveal a large amount of significant differences which may be related to speciation and to the divergence of ecological niches. Within each species, numerous geographic variations are also observed which, in most cases, reflect some kinds of climatic adaptation.

Animals↗