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Toward an ecological synthesis: a case for habitat selection.

Habitat selection, and its associated density and frequency-dependent evolution, has a profound influence on such vital phenomena as population regulation, species interactions, the assembly of ecological communities, and the origin and maintenance of biodiversity. Different strategies of habitat selection, and their importance in ecology and evolution, can often be revealed simply by plots of density in adjacent habitats. For individual species, the strategies are closely intertwined with mechanisms of population regulation, and with the persistence of populations through time. For interacting species, strategies of habitat selection are not only responsible for species coexistence, but provide one of the most convenient mechanisms for measuring competition, and the various community structures caused by competitive interactions. Other kinds of interactions, such as those between predators and prey, demonstrate that an understanding of the coevolution of habitat-selection strategies among strongly interacting species is essential to properly interpret their spatial and temporal dynamics. At the evolutionary scale, the frequency dependence associated with habitat selection may often allow populations to diverge and diversify into separate species. Habitat selection thereby demonstrates how we can map microevolutionary strategies in behavior onto their population and community consequences, and from there, onto macroevolutionary patterns of speciation and adaptive radiation. We can anticipate that future studies of habitat selection will not only help us complete those maps, but that they will also continue to enrich the panoply of ideas that shape evolutionary ecology.

Animals↗

Differential strength of sex-biased hybrid inferiority in impeding gene flow may be a cause of Haldane's rule.

In animals, if one sex of the F1 hybrid between two species is sterile or inviable, it is usually the heterogametic (XY or WZ) sex. This phenomenon, known as Haldane's rule, is currently thought to be coincidentally caused by different mechanisms in separate entities. The following questions have never been asked: Are heterogametic and homogametic inferiority (sterility or inviability) equivalent as isolating mechanisms? Could discrepancies between them, if existing, produce Haldane's rule? Here I consider sex-biased hybrid inferiority strictly as an isolating mechanism, and quantitatively evaluate its strength in impeding gene flow. The comparison reveals that the ability of sex-biased inferiority to impede gene flow varies according to the sex and chromosome involved. Heterogametic inferiority is a weaker barrier when unidirectional and a much stronger one when in compound reciprocal directions, compared with homogametic inferiority. Such differential strength may affect divergence in speciation and produce Haldane's rule.

Animals↗

Ecologic niche modeling and differentiation of populations of Triatoma brasiliensis neiva, 1911, the most important Chagas' disease vector in northeastern Brazil (hemiptera, reduviidae, triatominae).

Ecologic niche modeling has allowed numerous advances in understanding the geographic ecology of species, including distributional predictions, distributional change and invasion, and assessment of ecologic differences. We used this tool to characterize ecologic differentiation of Triatoma brasiliensis populations, the most important Chagas' disease vector in northeastern Brazil. The species' ecologic niche was modeled based on data from the Fundação Nacional de Saúde of Brazil (1997-1999) with the Genetic Algorithm for Rule-Set Prediction (GARP). This method involves a machine-learning approach to detecting associations between occurrence points and ecologic characteristics of regions. Four independent "ecologic niche models" were developed and used to test for ecologic differences among T. brasiliensis populations. These models confirmed four ecologically distinct and differentiated populations, and allowed characterization of dimensions of niche differentiation. Patterns of ecologic similarity matched patterns of molecular differentiation, suggesting that T. brasiliensis is a complex of distinct populations at various points in the process of speciation.

Animals↗

Organotin speciation in environmental matrices by automated on-line hydride generation-programmed temperature vaporization-capillary gas chromatography-mass spectrometry detection.

In the present contribution, a new automated on-line hydride generation methodology was developed for dibutyltin and tributyltin speciation at the trace level, using a programmable temperature-vaporizing inlet followed by capillary gas chromatography coupled to mass spectrometry in the selected ion-monitoring mode acquisition (PTV-GC/MS(SIM)). The methodology involves a sequence defined by two running methods, the first one configured for hydride generation with sodium tetrahydroborate as derivatising agent and the second configured for speciation purposes, using a conventional autosampler and data acquisition controlled by the instrument's software. From the method-development experiments, it had been established that injector configuration has a great effect on the speciation of the actual methodology, particularly, the initial inlet temperature (-20 degrees C; He: 150 ml/min), injection volume (2 microl) and solvent characteristics using the solvent venting mode. Under optimized conditions, a remarkable instrumental performance including very good precision (RSD < 4%), excellent linear dynamic range (up to 50 microg/ml) and limits of detection of 0.12 microg/ml and 9 ng/ml, were obtained for dibutyltin and tributyltin, respectively. The feasibility of the present methodology was validated through assays upon in-house spiked water (2 ng/ml) and a certified reference sediment matrix (Community Bureau of Reference, CRM 462, Nr. 330 dibutyltin: 68+/-12 ng/g; tributyltin: 54+/-15 ng/g on dry mass basis), using liquid-liquid extraction (LLE) and solid-phase extraction (SPE) sample enrichment and multiple injections (2 x 5 microl) for sensitivity enhancement. The methodology evidenced high reproducibility, is easy to work-up, sensitive and showed to be a suitable alternative to replace the currently dedicated analytical systems for organotin speciation in environmental matrices at the trace level.

Automation↗

Bacterial species and speciation.

Bacteria are profoundly different from eukaryotes in their patterns of genetic exchange. Nevertheless, ecological diversity is organized in the same way across all of life: individual organisms fall into more less discrete clusters on the basis of their phenotypic, ecological, and DNA sequence characteristics. Each sequence cluster in the bacterial world appears to correspond to an "ecotype," defined as a population of cells in the same ecological niche, which would all be out-competed by any adaptive mutant coming from the population. Ecotypes, so defined, share many of the dynamic properties attributed to eukaryotic species: genetic diversity within an ecotype is limited by a force of cohesion (in this case, periodic selection); different ecotypes are free to diverge without constraint from one another; and ecotypes are ecologically distinct. Also, ecotypes can be discovered and classified as DNA sequence clusters, even when we are ignorant of their ecology. Owing to the rarity and promiscuity of bacterial genetic exchange, speciation in the bacterial world is expected to be much less constrained than in the world of animals and plants.

Bacteria↗

Spatial grain and the causes of regional diversity gradients in ants.

Gradients of species richness (S; the number of species of a given taxon in a given area and time) are ubiquitous. A key goal in ecology is to understand whether and how the many processes that generate these gradients act at different spatial scales. Here we evaluate six hypotheses for diversity gradients with 49 New World ant communities, from tundra to rain forest. We contrast their performance at three spatial grains from S(plot), the average number of ant species nesting in a m2 plot, through Fisher's alpha, an index that treats our 30 1-m2 plots as subsamples of a locality's diversity. At the smallest grain, S(plot), was tightly correlated (r2 = 0.99) with colony abundance in a fashion indistinguishable from the packing of randomly selected individuals into a fixed space. As spatial grain increased, the coaction of two factors linked to high net rates of diversification--warm temperatures and large areas of uniform climate--accounted for 75% of the variation in Fisher's alpha. However, the mechanisms underlying these correlations (i.e., precisely how temperature and area shape the balance of speciation to extinction) remain elusive.

Animals↗

Geographical range and speciation in fossil and living molluscs.

The notion of a positive relation between geographical range and speciation rate or speciation probability may go back to Darwin, but a negative relation between these parameters is equally plausible. Here, we test these alternatives in fossil and living molluscan taxa. Late Cretaceous gastropod genera exhibit a strong negative relation between the geographical ranges of constituent species and speciation rate per species per million years; this result is robust to sampling biases against small-bodied taxa and is not attributable to phylogenetic effects. They also exhibit weak inverse or non-significant relations between geographical range and (i) the total number of species produced over the 18 million year timeframe, and (ii) the number of species in a single timeplane. Sister-group comparisons using extant molluscan species also show a non-significant relation between median geographical range and species richness of genera. These results support the view that the factors promoting broad geographical ranges also tend to damp speciation rates. They also demonstrate that a strong inverse relation between per-species speciation rate and geographical range need not be reflected in analyses conducted within a single timeplane, underscoring the inadequacy of treating net speciation as a proxy for raw per-taxon rates.

Animals↗

An integrated approach to taphonomy and faunal change in the Shungura formation (Ethiopia) and its implication for hominid evolution.

Environmental and faunal changes through time have been recorded for many African Plio-Pleistocene sites. Fossil evidence suggests that there is a continuous, if not uniform, transformation of the fauna and flora from the Pliocene through the end of Pleistocene. However, discerning major biotic turnovers and linking them to global and regional climatic changes have been complicated by many factors, notably taphonomy and discontinuity of the fossil evidence, notwithstanding the considerable work of some researchers (e.g., Vrba, E.S., 1988. Late Pliocene climatic events and hominid evolution, in: Grine, F. (Ed.), Evolutionary History of the "Robust" Australopithecines. De Gruyter, New York, pp. 405-426, Vrba, E.S., 1995. The fossil record of African (Mammalia, Bovidae) in relation to human evolution and paleoclimate, in: Vrba, E.S., Denton, G.H., Partridge, T.C., Burkle, L.H. (Eds.), Paleoclimate and Evolution, with Emphasis on Human Origins. Yale University Press, New Haven, pp. 385-424). A sample of over 22,000 fossils collected by the French Omo Expedition, from the Shungura Formation of Ethiopia, was analyzed using an integrated approach to investigate taphonomic and faunal change patterns. The following results are obtained: (1) Univariate and multivariate studies support continuous faunal change from Member A through Member G of the Shungura sequence; (2) Correspondence analysis (CA) on extant bovids in African game parks shows that bovid tribes and genera are generally characterized by habitat specificity; (3) Taphonomic studies demonstrate that the relative abundance of different skeletal elements varies according to depositional environment; (4) CA on 73 localities of the Shungura Formation and 19 mammalian taxa points to a major faunal change around the base of Member G dated to ca. 2.3 Ma. This transformation is characterized by a change to open and edaphic grassland as a dominant type of environment; (5) This major faunal change correlates in time with the appearance of A. boisei. It is tentatively suggested that this major biome change is associated with an anagenetic speciation from A. aethiopicus to A. boisei.

Animals↗

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↗

Perspective: Reproductive isolation caused by natural selection against immigrants from divergent habitats.

The classification of reproductive isolating barriers laid out by Dobzhansky and Mayr has motivated and structured decades of research on speciation. We argue, however, that this classification is incomplete and that the unique contributions of a major source of reproductive isolation have often been overlooked. Here, we describe reproductive barriers that derive from the reduced survival of immigrants upon reaching foreign habitats that are ecologically divergent from their native habitat. This selection against immigrants reduces encounters and thus mating opportunities between individuals from divergently adapted populations. It also reduces the likelihood that successfully mated immigrant females will survive long enough to produce their hybrid offspring. Thus, natural selection against immigrants results in distinctive elements of premating and postmating reproductive isolation that we hereby dub "immigrant inviability." We quantify the contributions of immigrant inviability to total reproductive isolation by examining study systems where multiple components of reproductive isolation have been measured and demonstrate that these contributions are frequently greater than those of traditionally recognized reproductive barriers. The relevance of immigrant inviability is further illustrated by a consideration of population-genetic theory, a review of selection against immigrant alleles in hybrid zone studies, and an examination of its participation in feedback loops that influence the evolution of additional reproductive barriers. Because some degree of immigrant inviability will commonly exist between populations that exhibit adaptive ecological divergence, we emphasize that these barriers play critical roles in ecological modes of speciation. We hope that the formal recognition of immigrant inviability and our demonstration of its evolutionary importance will stimulate more explicit empirical studies of its contributions to speciation.

Animals↗

Simulating a heavy metal spill under estuarine conditions: effects on the clam Scrobicularia plana.

We describe the effect of heavy metals Zn, Cd, Pb and Cu on the induction of methallothioneins on the clam Scrobicularia plana along a salinity gradient simulated under laboratory conditions. The clams were exposed to constant heavy metal concentrations in a dynamic estuary simulator during a 15-day assay to investigate possible induction of metal-binding proteins in them. The concentration of heavy metals in water was analysed. Clams were analysed for methallothionein concentrations. The speciation of Zn, Cd, Pb and Cu along the salinity gradient was modelled. Zn showed the highest concentrations and its prevalent species was the free ion. Intersite differences have been observed in methallothionein concentration and related to the salinity gradient. It seems that synthesis of methallothioneins is the result of physiological forces acting in concert with the changes in the chemical speciation of metals, owing to the trace metals uptake is controlled by means of an interaction of physiology and physicochemistry.

Animals↗

Studies on karyotype evolution in higher primates in relation to human chromosome 14 and 9 by comparative mapping of immunoglobulin C epsilon genes with fluorescence in situ hybridization.

Karyotypic homologies in relation to human chromosome 14 and 9 were studied through comparative mapping of the immunoglobulin C epsilon genes in higher primates by fluorescence in situ hybridization (FISH) technique. The C epsilon genes will be suitable probes for the analysis of evolutionary rearrangements due to that the multiple recombinational events such as gene duplications and deletions have occurred repeatedly in the immunoglobulin CH gene family (IGH@) during the course of primate evolution. IGH@ locating on the terminal region of human chromosome 14 (HSA14), at band HSA14q32.33, has generated multiple pseudogenes and among subclasses of IGH@ the C epsilon genes have shown most dynamic changes with generating both truncated type (C epsilon 2) and processed type (C epsilon 3) pseudogenes. In this study, chromosomal homologies and rearrangements on HSA14 (C epsilon 1) and HSA9 (C epsilon 3) in relation to the evolutionary genesis of their primate homologous chromosomes in speciation were investigated by comparative mapping with FISH and chromosome painting (ZOO-FISH) techniques. Comparative mapping of the C epsilon 1 gene at HSA14q32.33 was carried out in seven species of nonhuman primates: common chimpanzee (PTR), pygmy chimpanzee (PPA), gorilla (GGO), orangutan (PPY), white-handed gibbon (HLA), agile gibbon (HAG), and Japanese macaque (MFU). The C epsilon 1 gene was assigned to the telomeric region of HSA14 homologues in each species, namely, PTR15q32, PPA15q32, GGO18q16, PPY15q32, HLA17qter, HAG17qter, and MFU7q29, respectively. These results suggested that HSA14 has high degree of syntenic organization with its primate homologues confirmed by ZOO-FISH. Concerning HSA9, comparative mapping of the C epsilon 3 gene at HSA9p24.2-->p24.1 was performed. The mapped positions indicated the HSA9 homologous regions detected by ZOO-FISH in each species, namely, PTR11q34, PPA11q34, GGO13q22, PPY13q16, HLA8qter, HAG8qter, and MFU14q22, respectively, suggesting that several dynamic chromosomal rearrangements including at least twice pericentric inversions have occurred during the course of hominoid evolution. The comparison of syntenic groups and painting results has provided a hypothesis of the evolutionary genesis of HSA9 and its homologues with defined breakpoints on the present chromosomes. Likewise, studies on karyotype evolution will be promoted by combining comparative mapping with ZOO-FISH that can more clearly define the chromosomal rearrangements among species.

Animals↗

Importance of natural soil processes relative to atmospheric deposition in the mobility of aluminium in forested watersheds of the Black Forest.

The dynamics of aqueous aluminium in the ARINUS experimental watersheds at Schluchsee (granite) and Villingen (quartz sandstone), Black Forest (South-west Germany), were studied in order to detect the processes and factors controlling its mobility. Aluminium speciation was performed in the seepage of typical soils (podsol, acidic brown earth, stagnogley) at 3 depths (organic layer, 30 cm and 80 cm of the mineral soil) as well as in streamwater. The studies concentrated on the variability in time and space of inorganic monomeric Al (Ali), and organic monomeric Al (Alo). Furthermore, the equilibrium speciation model WATEQF was used to calculate the distribution of inorganic Al species. Natural soil properties and processes, such as DOC mobilization and excess mineralization of NO3(2-) and SO4(2-), appeared to have great influence and outweigh the deposition effects upon Al mobilization in these systems which receive only low to moderate loads of acidic deposition.

Journal Article↗

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↗

The effects of intraspecific competition and stabilizing selection on a polygenic trait.

The equilibrium properties of an additive multilocus model of a quantitative trait under frequency- and density-dependent selection are investigated. Two opposing evolutionary forces are assumed to act: (i) stabilizing selection on the trait, which favors genotypes with an intermediate phenotype, and (ii) intraspecific competition mediated by that trait, which favors genotypes whose effect on the trait deviates most from that of the prevailing genotypes. Accordingly, fitnesses of genotypes have a frequency-independent component describing stabilizing selection and a frequency- and density-dependent component modeling competition. We study how the equilibrium structure, in particular, number, degree of polymorphism, and genetic variance of stable equilibria, is affected by the strength of frequency dependence, and what role the number of loci, the amount of recombination, and the demographic parameters play. To this end, we employ a statistical and numerical approach, complemented by analytical results, and explore how the equilibrium properties averaged over a large number of genetic systems with a given number of loci and average amount of recombination depend on the ecological and demographic parameters. We identify two parameter regions with a transitory region in between, in which the equilibrium properties of genetic systems are distinctively different. These regions depend on the strength of frequency dependence relative to pure stabilizing selection and on the demographic parameters, but not on the number of loci or the amount of recombination. We further study the shape of the fitness function observed at equilibrium and the extent to which the dynamics in this model are adaptive, and we present examples of equilibrium distributions of genotypic values under strong frequency dependence. Consequences for the maintenance of genetic variation, the detection of disruptive selection, and models of sympatric speciation are discussed.

Biological Evolution↗

Chromosome-Level Genome Assembly of Eden's Whale Clarifies the Taxonomy and Speciation of Bryde's Whale Complex.

Eden's whale (Balaenoptera edeni), a poorly understood baleen cetacean, has long been shrouded in taxonomic ambiguity due to limited genomic resources, obscuring its distinction from closely related species and its position within the cetacean Tree of Life. In this paper, we present a high-quality chromosomal-level genome of B. edeni and conduct comparative genomic analyses to address long-standing taxonomic confusion and elucidate speciation of balaenopterids. Our phylogenomic analysis and demographic reconstruction reveal that B. edeni is a distinct sister to Bryde's whale (Balaenoptera brydei), sharing a common ancestor that diverged approximately 7.84 million years ago during the late Miocene. Their genetic divergence exceeds typical intraspecific variation in whales, supporting the reinstatement of B. brydei as a valid species. Chromosomal syntenic analyses suggest that macro-fragment inversions contributed to speciation in balaenopterid whales and uncover unexpected large-scale complex genome rearrangements in Bryde's whale, offering novel insights into cetacean genome evolution. Functional enrichment analysis of inverted regions between B. edeni and Balaenoptera musculus indicates their predominant association with metabolism and biosynthesis, as well as responses to various substances, stress, and stimuli. These genomic resources for B. edeni not only lay a critical foundation for comparative genetic and evolutionary research of cetaceans but also advance our understanding of the taxonomy and evolutionary dynamics of the Bryde's whale complex, with broader implications for baleen whale conservation and biodiversity.

Animals↗

The dynamic herpesvirus DNA genome: the case of MDV-1 and HSV-1.

Herpesviruses evolved from an ancestral viral genome that contained five blocks of genes which provide the members of this family of viruses with structural and enzymatic properties. These genes allow the herpesviruses to infect a host by entering into the nuclei of the cells, the site of replication and transcription of the viral DNA. The viral mRNAs are released into the cell cytoplasm where synthesis of enzymatic and structural proteins occurs. The latter proteins are responsible for the formation of the infectious virions. Herpesviruses that were able to adapt to different hosts during the evolution of the species (speciation) had acquired additional genes from transposons or retrotransposons that allowed them to successfully maintain their hold in the specific vertebrate host. The present overview deals with molecular differences between Marek's disease virus type 1 (MDV-1) and herpes simplex virus type 1 (HSV-1) and the specialized genes that differentiate MDV-1 from HSV-1, the promoters of the viral genes that control gene expression and the nuclear localization signals. Dynamic changes in the viral genomes that may occur during viral DNA replication and recombination and their effects on virus pathogenicity and genome evolution will be discussed.

Animals↗