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Historical processes enhance patterns of diversity along latitudinal gradients.

One of the more vexing issues in ecology is how historical processes affect contemporary patterns of biodiversity. Accordingly, few models have been presented. Two corollary models (centre of origin, time-for-speciation) can be used to make quantitative predictions characterizing the tropical niche conservatism hypothesis and describe diversification as diffusion and subsequent cladogenesis of species away from the place of origin of a higher taxon in the tropics. Predictions derived from such models are: (i) species richness declines toward the periphery of the range of a higher taxon; (ii) taxa are more derived toward the periphery than the centre; (iii) ages of taxa are lower toward the periphery than the centre; and (iv) ages and measures of derivedness are less variable toward the periphery of the range of a higher taxon. I tested these predictions to better understand the formation of one of the most ubiquitous patterns of biodiversity-the latitudinal gradient in species richness. Results indicate well-supported predictions for New World leaf-nosed bats and that diversification has had strong influences on latitudinal gradients of species richness. A better understanding of how evolutionary diversification of taxa contributes to formation of patterns of species richness along environmental gradients is necessary to fully understand spatial variation in biodiversity.

Animals↗

Using phylogeographic analyses of gene trees to test species status and processes.

A gene tree is an evolutionary reconstruction of the genealogical history of the genetic variation found in a sample of homologous genes or DNA regions that have experienced little or no recombination. Gene trees have the potential of straddling the interface between intra- and interspecific evolution. It is precisely at this interface that the process of speciation occurs, and gene trees can therefore be used as a powerful tool to probe this interface. One application is to infer species status. The cohesion species is defined as an evolutionary lineage or set of lineages with genetic exchangeability and/or ecological interchangeability. This species concept can be phrased in terms of null hypotheses that can be tested rigorously and objectively by using gene trees. First, an overlay of geography upon the gene tree is used to test the null hypothesis that the sample is from a single evolutionary lineage. This phase of testing can indicate that the sampled organisms are indeed from a single lineage and therefore a single cohesion species. In other cases, this null hypothesis is not rejected due to a lack of power or inadequate sampling. Alternatively, this null hypothesis can be rejected because two or more lineages are in the sample. The test can identify lineages even when hybridization and lineage sorting occur. Only when this null hypothesis is rejected is there the potential for more than one cohesion species. Although all cohesion species are evolutionary lineages, not all evolutionary lineages are cohesion species. Therefore, if the first null hypothesis is rejected, a second null hypothesis is tested that all lineages are genetically exchangeable and/or ecologically interchangeable. This second test is accomplished by direct contrasts of previously identified lineages or by overlaying reproductive and/or ecological data upon the gene tree and testing for significant transitions that are concordant with the previously identified lineages. Only when this second null hypothesis is rejected is a lineage elevated to the status of cohesion species. By using gene trees in this manner, species can be identified with objective, a priori criteria with an inference procedure that automatically yields much insight into the process of speciation. When one or more of the null hypotheses cannot be rejected, this procedure also provides specific guidance for future work that will be needed to judge species status.

Animals↗

Protist biogeography.

Autecology (cellular organelles and secretions, encystment and dispersal abilities), environmental conditions (physiological tolerances and interaction with other organisms), and evolutionary history contribute to protist biogeography, which manifests ecological and historical aspects. Ecological biogeography is seen in the influence of geochemistry on the distribution of fresh-water phytoflagellates and algae, and of moisture and vegetation type on soil-litter protists. A temporal feature is often present because many protists encyst and respond only to certain ranges of temperature and organic content. Historical biogeography has occurred by radiative host evolution on symbiotic protozoa (e.g., termite flagellates and rumen ciliates), and by the isolating effects of water currents, temperature, and density gradients on oceanic protists (coccoliths, dinoflagellates, foraminifera, radiolaria, and tintinnines). These two aspects combine in protists living on animal surfaces. Humans affect protist biogeography by increasing parasite ranges through human migrations and by water pollution. They can diminish these situations by disease control and exploiting appropriate ciliates in sewage disposal. Many free-living protozoa appear to be cosmopolitan, but mating types and isoenzyme studies suggest that speciation with its geographic connotations may be more widespread than presently appreciated.

Animals↗

Ecology and evolution of galling thrips and their allies.

About 300 species of thrips belonging to 57 genera are known to form galls. Galls are caused by feeding, usually by one or more adults, on actively growing plant tissue. Most thrips genera with galling capabilities exploit multiple plant families, but there are several possible cases of thrips tracking the speciations of their host-plants. Gall morphology in thrips reflects insect phylogenetic relationships rather than those of plants. Galling species and their nongalling allies on Acacia in Australia exhibit a range of complex social behavior, including soldier castes, pleometrosis (i.e. joint colony founding), group foraging, and group defense, that is directly related to the nature of their domiciles. Galling thrips, by virtue of their haplodiploid genetic system and their ecological relationships with plants and natural enemies, are useful for analyzing a wide range of ecological, evolutionary, and behavioral questions.

Journal Article↗

On the heritability of geographic range sizes.

Within taxonomic groups, most species are restricted in their geographic range sizes, with only a few being widespread. The possibility that species-level selection on range sizes contributes to the characteristic form of such species-range size distributions has previously been raised. This would require that closely related species have similar range sizes, an indication of "heritability" of range sizes at the species level. Support for this view came from a positive correlation between the range sizes of closely related pairs of fossil mollusc species. We extend this analysis by considering the relationship between the geographic range sizes of 103 pairs of contemporary avian sister species. Range sizes in these sister species show no evidence of being more similar to each other than expected by chance. A reassessment of the mollusc data also suggests that the high correlation was probably overestimated because of the skewed nature of range size data. The fact that sister species tend to have similar life histories and ecologies suggests that any relationship between range sizes and biology is likely to be complicated and will be influenced by historical factors, such as mode of speciation and postspeciation range size transformations.

Animals↗

Global diversification rates of passerine birds.

The distribution of species richness in families of passerine birds suggests that the net rate of diversification was significantly higher than average in as many as 7 out of 47 families. However, the absence of excess species richness among the 106 tribes within these families indicates that these high rates were transient, perhaps associated in some cases with tectonic movements or dispersal events that extended geographical ranges. Thus, large clade size among passerine birds need not represent intrinsic key innovations that influence the rate of diversification. Approximately 17 families and 30 tribes have too few species relative to other passerine taxa. Many of these are ecologically or geographically marginal, being especially overrepresented in the Australasian region. Observed intervals between lineage splitting suggest that extinction has occurred ca. 90% as frequently as speciation (waiting times of 1.03 and 0.93 Myr) and that the 47 modern families comprising 5712 species descended from approximately 430 passerine lineages extant 24 Myr ago. Speciation and extinction rates among small, marginal families might be 1-2 orders of magnitude lower.

Animals↗

Aedes polynesiensis in the Society Islands: environmental correlates of isoenzyme differentiation.

Isoenzyme genetic differentiation of Aedes polynesiensis mosquitoes in Raiatea island, French Polynesia, was evaluated by two models of population structure based on seven gene-enzyme systems: Ak, Est, Got, Gpi, Hk, Mdh and Pgm. The ecological model tested whether genetic differentiation is congruent with habitat differences. The isolation model evaluated whether genetic differentiation is proportional to geographical distribution. The ecological model found no significant differentiation between populations of Ae.polynesiensis from beach and forest ecotopes, whereas the isolation model was consistent with the data. However, incipient speciation is opposed by the source-sink system of population dynamics in such small neighbouring islands, where Ae.polynensiensis extinction is readily followed by reinvasion causing considerable gene flow between island populations.

Aedes↗

The ecology of individuals: incidence and implications of individual specialization.

Most empirical and theoretical studies of resource use and population dynamics treat conspecific individuals as ecologically equivalent. This simplification is only justified if interindividual niche variation is rare, weak, or has a trivial effect on ecological processes. This article reviews the incidence, degree, causes, and implications of individual-level niche variation to challenge these simplifications. Evidence for individual specialization is available for 93 species distributed across a broad range of taxonomic groups. Although few studies have quantified the degree to which individuals are specialized relative to their population, between-individual variation can sometimes comprise the majority of the population's niche width. The degree of individual specialization varies widely among species and among populations, reflecting a diverse array of physiological, behavioral, and ecological mechanisms that can generate intrapopulation variation. Finally, individual specialization has potentially important ecological, evolutionary, and conservation implications. Theory suggests that niche variation facilitates frequency-dependent interactions that can profoundly affect the population's stability, the amount of intraspecific competition, fitness-function shapes, and the population's capacity to diversify and speciate rapidly. Our collection of case studies suggests that individual specialization is a widespread but underappreciated phenomenon that poses many important but unanswered questions.

Animals↗

Phylogeny of the Baldratiina (Diptera: Cecidomyiidae) inferred from morphological, ecological and molecular data sources, and evolutionary patterns in plant-galler relationships.

The phylogeny of the gall-midge subtribe Baldratiina (Diptera: Cecidomyiidae) was reconstructed from molecular (partial sequence of the mitochondrial 12S rDNA), morphological and ecological data sets, using 16 representative species of most of the genera. The morphological and ecological data were combined in a single character matrix and analyzed separately from the molecular data, resulting in an eco-morphological cladogram and a molecular cladogram. Attributes of galls and host associations were superimposed on the molecular cladogram in order to detect possible trends in the evolution of these traits. The cladograms resulting from the two independent analyses were statistically incongruent, although both provide evidence for the monophyly of the genera Baldratia and Careopalpis and the paraphyly of the genera Stefaniola and Izeniola. The results suggest a minor impact of the morphological characters traditionally used in the classification of the Baldratiina, whereas ecological data had a major impact on the phylogenetic inference. Mapping of gall and host attributes on the molecular cladogram suggests that multi-chambered stem galls constitute the ancestral state in the subtribe, with several subsequent shifts to leaf galls. It is concluded that in contrast to other studied groups of gall insects, related baldratiine species induce different types of galls, attesting to speciation driven by gall-type shifts at least as often as host shifts.

Animals↗

Impact of large chromosomal inversions on the adaptation and evolution of Pseudomonas aeruginosa chronically colonizing cystic fibrosis lungs.

Pseudomonas aeruginosa chronically colonizing the lungs of cystic fibrosis (CF) patients undergoes fast evolution leading to clonal divergence. More than half of the genotypes of P. aeruginosa clone C isolates exclusively from CF lung infection exhibit large chromosomal inversions (LCIs). To analyse the impact of LCIs, as a novel mechanism of bacterial adaptation, the underlying molecular mechanism was examined. Analysis of inversion breakpoints suggested an IS6100-induced coupled insertion-inversion mechanism. A selective advantage was created by insertion of IS6100 into wbpM, pilB and mutS which leads to common CF phenotypes such as O-antigen and type IV pili deficiency and hypermutability. Speciation in bacteria is accompanied by LCIs. Therefore adaptation by LCIs that allows persistence of P. aeruginosa in the CF lung and species diversification in that new ecological niche can serve as a model for bacterial genome evolution.

Adaptation, Physiological↗

DNA sequence variation in the mitochondrial control region of subterranean mole rats, Spalax ehrenbergi superspecies, in Israel.

The complete mitochondrial control region was sequenced for 60 individuals representing different populations for each of the four species of the subterranean mole rat Spalax ehrenbergi superspecies in Israel: Spalax galili (2n = 52), S. golani (2n = 54), S. carmeli (2n = 58), and S. judaei (2n = 60). The control region of all species and populations is very similar both in length (979 to 983 bp) and in base composition. As in agreement with previous surveys on mitochondrial control regions on mammals, the mole rat control region can be divided into a central domain and two flanking domains, ETAS (extended termination associated sequences) and CSB (conserved sequence blocks). Along with the common conserved blocks found in these domains (ETAS1, ETAS2, CSB1, CSB2, and CSB3), we have also detected in all individuals an ETAS1-like and a CSB1-like element, both in the ETAS domain. The most conserved region was the central domain, followed by the CSB and ETAS domains, showing important differences in the four species analyzed. Phylogenetic analysis supported the existence of two clades. One clade contained individuals belonging to Spalax galili (2n = 52) and S. golani (2n = 54), separated in two different branches depending on the species. The other clade contained individuals belonging to S. carmeli (2n = 58) and S. judaei (2n = 60) mixed together, suggesting a more recent event of speciation. Within species we have observed a southward trend of increasing variability. These results have been explained as a consequence of the adaptation of the species to ecological factors such as aridity and temperature stresses.

Animals↗

Evolution of the mojavensis cluster of cactophilic Drosophila with descriptions of two new species.

The mojavensis cluster of the repleta species group of Drosophila (Drosophilidae: Diptera) consists of three species. One is newly described as D. navojoa. A second species, described here as D. arizonae, replaces D. arizonensis, which has become a junior subjective synonym for D. mojavensis, the third species in the cluster. A phylogeny of the three species is presented, based on chromosomal inversions, morphology, and the ability to produce hybrids. Breakage points are assigned for all inversions, and male genitalia are figured; 186 crosses were made from 225 possible combinations among 15 geographic strains from the southwestern United States, Mexico, and Guatemala. It is confirmed that D. mojavensis and D. arizonae are very closely related and shown that D. navojoa is more distantly related in regard to all criteria. This relationship is supported by the geographical positions of the ancestral gene sequences in each species, which show a sequential northwest movement (D. navojoa----D. arizonae----D. mojavensis) from southern Mexico to southern California and northern Arizona. The relationship is also supported by the fact that D. navojoa breeds in Opuntia cactus, an ancestral behavior, whereas the other two species breed chiefly in Stenocereus cacti, a derived behavior. The possible role of this host plant shift in speciation is discussed.

Animals↗

Biodiversity and the Lotka-Volterra theory of species interactions: open systems and the distribution of logarithmic densities.

Theoretical interest in the distributions of species abundances observed in ecological communities has focused recently on the results of models that assume all species are identical in their interactions with one another, and rely upon immigration and speciation to promote coexistence. Here we examine a one-trophic level system with generalized species interactions, including species-specific intraspecific and interspecific interaction strengths, and density-independent immigration from a regional species pool. Comparisons between results from numerical integrations and an approximate analytic calculation for random communities demonstrate good agreement, and both approaches yield abundance distributions of nearly arbitrary shape, including bimodality for intermediate immigration rates.

Biodiversity↗

Longevity and ageing: appraising the evolutionary consequences of growing old.

Senescence or ageing is an increase in mortality and/or decline in fertility with increasing age. Evolutionary theories predict that ageing or longevity evolves in response to patterns of extrinsic mortality or intrinsic damage. If ageing is viewed as the outcome of the processes of behaviour, growth and reproduction then it should be possible to predict mortality rate. Recent developments have shown that it is now possible to integrate these ecological and physiological processes and predict the shape of mortality trajectories. By drawing on the key exciting developments in the cellular, physiological and ecological process of longevity the evolutionary consequences of ageing are reviewed. In presenting these ideas an evolutionary demographic framework is used to argue how trade-offs in life-history strategies are important in the maintenance of variation in longevity within and between species. Evolutionary processes associated with longevity have an important role in explaining levels of biological diversity and speciation. In particular, the effects of life-history trait trade-offs in maintaining and promoting species diversity are explored. Such trade-offs can alleviate the effects of intense competition between species and promote species coexistence and diversification. These results have important implications for understanding a number of core ecological processes such as how species are divided among niches, how closely related species co-occur and the rules by which species assemble into food-webs. Theoretical work reveals that the proximate physiological processes are as important as the ecological factors in explaining the variation in the evolution of longevity. Possible future research challenges integrating work on the evolution and mechanisms of growing old are briefly discussed.

Aging↗

Behavioral syndromes: an ecological and evolutionary overview.

Recent studies suggest that populations and species often exhibit behavioral syndromes; that is, suites of correlated behaviors across situations. An example is an aggression syndrome where some individuals are more aggressive, whereas others are less aggressive across a range of situations and contexts. The existence of behavioral syndromes focuses the attention of behavioral ecologists on limited (less than optimal) behavioral plasticity and behavioral carryovers across situations, rather than on optimal plasticity in each isolated situation. Behavioral syndromes can explain behaviors that appear strikingly non-adaptive in an isolated context (e.g. inappropriately high activity when predators are present, or excessive sexual cannibalism). Behavioral syndromes can also help to explain the maintenance of individual variation in behavioral types, a phenomenon that is ubiquitous, but often ignored. Recent studies suggest that the behavioral type of an individual, population or species can have important ecological and evolutionary implications, including major effects on species distributions, on the relative tendencies of species to be invasive or to respond well to environmental change, and on speciation rates. Although most studies of behavioral syndromes to date have focused on a few organisms, mainly in the laboratory, further work on other species, particularly in the field, should yield numerous new insights.

Journal Article↗

Polyploid incidence and evolution.

Changes in ploidy occurred early in the diversification of some animal and plant lineages and represent an ongoing phenomenon in others. While the prevalence of polyploid lineages indicates that this phenomenon is a common and successful evolutionary transition, whether polyploidization itself has a significant effect on patterns and rates of diversification remains an open question. Here we review evidence for the creative role of polyploidy in evolution. We present new estimates for the incidence of polyploidy in ferns and flowering plants based on a simple model describing transitions between odd and even base chromosome numbers. These new estimates indicate that ploidy changes may represent from 2 to 4% of speciation events in flowering plants and 7% in ferns. Speciation via polyploidy is likely to be one of the more predominant modes of sympatric speciation in plants, owing to its potentially broad-scale effects on gene regulation and developmental processes, effects that can produce immediate shifts in morphology, breeding system, and ecological tolerances. Theoretical models support the potential for increased adaptability in polyploid lineages. The evidence suggests that polyploidization can produce shifts in genetic systems and phenotypes that have the potential to result in increased evolutionary diversification, yet conclusive evidence that polyploidy has changed rates and patterns of diversification remains elusive.

Animals↗

Karyotype differentiation in Chromaphyosemion killifishes (Cyprinodontiformes, Nothobranchiidae). II: cytogenetic and mitochondrial DNA analyses demonstrate karyotype differentiation and its evolutionary direction in C. riggenbachi.

African killifishes of the genus Chromaphyosemion show a high degree of phenotypic and karyotypic diversity. The latter is especially pronounced in C. riggenbachi, a morphologically defined species restricted to a small distribution area in Cameroon. This study presents a detailed reconstruction of karyotype differentiation within C. riggenbachi using conventional Giemsa staining and sequential chromosome banding as well as a phylogenetic analysis based on part of the mitochondrial (mt) cytochrome b gene from eleven populations. The cytogenetic analysis revealed differences in chromosome morphology, banding patterns and/or diploid chromosome number (2n) among all populations examined. Diploid number ranged from 2n = 20 to 2n = 36 and varied mainly among populations, while C-banding patterns and NOR phenotypes showed fixed differences among populations as well as some variability within populations. The mtDNA analysis disclosed five clearly differentiated haplotype groups. Mapping the karyotype data onto the mtDNA dendrogram revealed a decrease in 2n from the most basal to the most derived groups, thus demonstrating a reduction of 2n during their evolutionary history. Our results indicate that karyotype differentiation involved Robertsonian fusions as well as non-Robertsonian processes. Causes of the high karyotypic variability may include an elevated chromosomal mutation rate as well as certain features of the ecology and mating system that could facilitate the fixation of chromosomal rearrangements. The pattern of karyotype and haplotype differentiation and the results of previous crossing experiments suggest incipient speciation in C. riggenbachi.

Africa↗

Continuing the debate on the role of Quaternary environmental change for macroevolution.

The Quaternary has been a period of dramatic environmental change for the past 1.8 Myr, with major shifts in distributions and abundances of terrestrial and marine organisms. The evolutionary consequences of this have been debated since the nineteenth century. However, the lack of accurate relative and absolute time-scales for evolutions and environmental change inhibited progress. We do now have an understanding of time-scales. Palaeoecology has demonstrated the individualistic nature of species' response to environmental change, but lacks a means of determining ancestry. DNA characterization of modern populations in relation to their distributions nicely complements palaeoecological results by contributing ancestry. The chance to understand how species originate and the causal factors of speciation (environmental change or otherwise) may be within reach.

Animals↗