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Dwarfism in insular sloths: biogeography, selection, and evolutionary rate.

The islands of Bocas del Toro, Panama, were sequentially separated from the adjacent mainland by rising sea levels during the past 10,000 years. Three-toed sloths (Bradypus) from five islands are smaller than their mainland counterparts, and the insular populations themselves vary in mean body size. We first examine relationships between body size and physical characteristics of the islands, testing hypotheses regarding optimal body size, evolutionary equilibria, and the presence of dispersal in this system. To do so, we conduct linear regressions of body size onto island area, distance from the mainland, and island age. Second, we retroactively calculate two measures of the evolutionary rate of change in body size (haldanes and darwins) and the standardized linear selection differential, or selection intensity (i). We also test the observed morphological changes against models of evolution by genetic drift. The results indicate that mean body size decreases linearly with island age, explaining up to 97% of the variation among population means. Neither island area nor distance from the mainland is significant in multiple regressions that include island age. Thus, we find no evidence for differential optimal body size among islands, or for dispersal in the system. In contrast, the dependence of body size on island age suggests uniform directional selection for small body size in the insular populations. Although genetic drift cannot be discounted as the cause for this evolution in body size, the probability is small given the consistent direction of evolution (repeated dwarfism). The insular sloths show a sustained rate of evolution similar to those measured in haldanes over tens of generations, appearing to unite micro- and macroevolutionary time scales. Furthermore, the magnitude and rate of this example of rapid differentiation fall within predictions of theoretical models from population genetics. However, the linearity of the relationship between body size and island age is not predicted, suggesting that either more factors are involved than those considered here, or that theoretical advances are necessary to explain constant evolutionary rates over long time spans in new selective environments.

Animals↗

The history of a nearctic colonization: molecular phylogenetics and biogeography of the Nearctic toads (Bufo).

Previous hypotheses of phylogenetic relationships among Nearctic toads (Bufonidae) and their congeners suggest contradictory biogeographic histories. These hypotheses argue that the Nearctic Bufo are: (1) a polyphyletic assemblage resulting from multiple colonizations from Africa; (2) a paraphyletic assemblage resulting from a single colonization event from South America with subsequent dispersal into Eurasia; or (3) a monophyletic group derived from the Neotropics. We obtained approximately 2.5 kb of mitochondrial DNA sequence data for the 12S, 16S, and intervening valine tRNA gene from 82 individuals representing 56 species and used parametric bootstrapping to test hypotheses of the biogeographic history of the Nearctic Bufo. We find that the Nearctic species of Bufo are monophyletic and nested within a large clade of New World Bufo to the exclusion of Eurasian and African taxa. This suggests that Nearctic Bufo result from a single colonization from the Neotropics. More generally, we demonstrate the utility of parametric bootstrapping for testing alternative biogeographic hypotheses. Through parametric bootstrapping, we refute several previously published biogeographic hypotheses regarding Bufo. These previous studies may have been influenced by homoplasy in osteological characters. Given the Neotropical origin for Nearctic Bufo, we examine current distributional patterns to assess whether the Nearctic-Neotropical boundary is a broad transition zone or a narrow boundary. We also survey fossil and paleogeographic evidence to examine potential Tertiary and Cretaceous dispersal routes, including the Paleocene Isthmian Link, the Antillean and Aves Ridges, and the current Central American Land Bridge, that may have allowed colonization of the Nearctic.

Animals↗

Pattern and process in the ecological biogeography of European freshwater fish.

1. Species lists for regions of Europe defined by Illies (1978, Limnofauna Europaea, 2nd edn. Gustav Fischer Verlag, Stuttgart), and augmented by information from Maitland (2000, Guide to Freshwater Fish of Britain and Europe. Hamlyn, London), are used to describe patterns in freshwater fish species richness and to examine the contribution of habitat preference, migration, body size and glacial history to these patterns. 2. The number of non-endemic species declines to the north and west, with increasing distance from the Ponto-Caspian region, the main source area, whereas endemic species richness declines only with latitude. 3. Habitat generalists tend to be migratory while riverine specialists are usually resident. Similar numbers of riverine species and generalists occur in Europe as a whole but generalists dominate in regional faunas and, to an increasing extent, in more isolated, formerly glaciated areas. Very few lacustrine specialists were found, reflecting the geologically ephemeral nature of lakes. Only 8% of riverine species have colonized glaciated areas, compared with more than half the generalist species, and the number declines rapidly with increasing distance from the source area. 4. Diadromous species show no geographical variation in species richness but potamodromous and resident species are affected by glaciation and by mountain and marine barriers. 5. The mean body size of regional faunas increases with latitude because there are relatively fewer small species in more distant, glaciated areas. 6. About half the species occurring in Europe are restricted to one region and the majority of these endemics occur in barrier regions with Mediterranean climates. Species in glaciated regions have much larger range sizes. Habitat preference and migration type, not body size, are the main determinants of range size. 7. Freshwater habitat availability varies across Europe with glaciated areas having more lakes of a given size than unglaciated areas. Catchment size is greatest at mid-latitudes. For a given catchment size rivers in glaciated areas are shorter. 8. The results support the notion that habitat variability, on both short and long time scales, favours colonization ability, which requires large body size. 9. As a result of their limited vagility northern fish faunas are depauperate, show high levels of plasticity and polymorphism and may show elevated speciation rates. The isolated southern faunas of the Iberian and Italian peninsulas and the Balkans are rich in endemic species but may be subject to extinctions because of the spread of the highly seasonal Mediterranean climate.

Animal Migration↗

A molecular analysis of relationships and biogeography within a species complex of Holarctic fish (genus Osmerus).

Episodes of trans-Arctic faunal exchange and isolation between the north Pacific and Atlantic ocean basins have been implicated as important historic geological events contributing to extant patterns of genetic diversity and structure in Holarctic faunas. We made a further test of the significance of such biogeographic events by examining mitochondrial DNA (mtDNA) restriction fragment length and cytochrome b sequence polymorphism among north Pacific and Arctic, north-western Atlantic (north-eastern North American), and north-eastern Atlantic (European) regional forms of the boreal smelt, genus Osmerus. Our analyses also assessed whether the regional forms within this 'species complex': (i) represent a single widely distributed and polytypic species, or is composed of three geographically distinct species, and (ii) resulted from a single split from north Pacific ancestral Osmerus or two independent Pacific-Atlantic divergences. MtDNA sequence divergence estimates among forms ranged from 5.6-8.9% and from 6.1-8.5% based on restriction fragment and 300 base pairs of cytochrome b sequencing, respectively. Divergence within forms averaged less than 0.5% for fragment analysis and no differences were detected from sequence analysis. Provisional dating of lineage separations in Osmerus based on our sequence divergence estimates suggested a mid-Pliocene to early Pleistocene time frame for diversification among the forms. These estimated lineage separation dates support the idea that geological events in 'Beringia' and the surrounding trans-Arctic area (e.g. opening of the Bering Seaway, Pleistocene glacial advances), occurring over a similar time frame, have influenced radiation in Osmerus. Phenetic and parsimony analyses of the sequence divergence estimates and of sequence polymorphisms suggested that the north Pacific/Arctic form and the north-western Atlantic form shared a common ancestor more recently than either has with the north-eastern Atlantic form, thus supporting the hypothesis that the species complex has arisen from two independent Pacific-Atlantic divergences probably beginning during the mid-Pliocene.

Animals↗

Molecular biogeography of clonal lineages in a high-arctic apomictic Daphnia complex.

An electrophoretic survey of 81 populations of arctic Daphnia pulex from around the Svalbard archipelago revealed the presence of 49 unique allozyme clones (N = 3357). Two closely related clones accounted for 66% of the total sample, and were widespread across the archipelago. Restriction fragment length polymorphisms (RFLPs) of a 2.1-kb fragment of mtDNA (NADH-4 and NADH-5 subunits), amplified using the polymerase chain reaction (PCR), revealed the presence of eight mtDNA haplotypes. One haplotype was particularly widespread, and the two most abundant allozyme clones shared this haplotype. Nonrandom distribution patterns of clones were observed, and are most likely the result of historical events (i.e. founder effects) related to the past glacial history of the archipelago. The data are discussed with reference to past glaciation events, and attempts are made to discern the colonization history of this apomictic complex.

Animals↗

Molecular characterization of clonal population structure and biogeography of arctic apomictic Daphnia from Greenland and Iceland.

As part of a continuing international project to characterize the population genetic (clonal) structure of arctic members of the Daphnia pulex complex, 34 populations from western Iceland (N = 1373), and 76 populations from western Greenland (N = 2917), were surveyed for allozymic variation at six polymorphic enzyme loci. Mean clonal richness (+/- SE) was 1.91 +/- 0.19 and 1.50 +/- 0.12 for Iceland and Greenland populations, respectively. Mean clonal diversity (+/-1 SE) was 1.20 +/- 0.07 and 1.13 +/- 0.04 for Iceland and Greenland populations, respectively. Four widely distributed clones constituted 92.2% of the total animals surveyed from Iceland, while three locally abundant clones constituted 80.1% of the total animals collected primarily from Disko Island, western Greenland. Selected populations were screened for mitochondrial (mt)DNA variation using PCR-based RFLP analysis of a 2100 bp fragment containing part of the ND4 and ND5 genes. One mtDNA haplotype was very widespread in both western Greenland and western Iceland, although, a number of mutational derivatives were also detected. These data indicate the potential for long-distance dispersal of mtDNA lineages, of the order of hundreds or thousands of kilometers across the arctic. Phylogenetic analysis of the sequence of a 254 base pair (bp) fragment of the control region of the mtDNA molecule revealed two major clades one of which consisted solely of non-melanic lineages, and the other of which consisted almost exclusively of melanic lineages (i.e. one non-melanic lineage also clustered in this clade). Sequence divergence between the two clades averaged 7.3%. Both mitochondrial analyses did not reveal any distinct intraregional clustering of lineages. We discuss our results in reference to previous molecular work done on this arctic Daphnia complex, and we attempt to infer phylogeographic patterning based on geological/glaciological historical events in this region of the arctic.

Animals↗

Phylogeny and historical biogeography of African ground squirrels: the role of climate change in the evolution of Xerus.

We used phylogenetic and phylogeographical methods to infer relationships among African ground squirrels of the genus Xerus. Using Bayesian, maximum-parsimony, nested clade and coalescent analyses of cytochrome b sequences, we inferred interspecific relationships, evaluated the specific distinctness of Cape (Xerus inauris) and mountain (Xerus princeps) ground squirrels, and tested hypotheses for historical patterns of gene flow within X. inauris. The inferred phylogeny supports the hypothesized existence of an 'arid corridor' from the Horn of Africa to the Cape region. Although doubts have been raised regarding the specific distinctness of X. inauris and X. princeps, our analyses show that each represents a distinct well-supported, monophyletic lineage. Xerus inauris includes three major clades, two of which are geographically restricted. The distributions of X. inauris populations are concordant with divergences within and disjunctions between other taxa, which have been interpreted as results of Plio-Pleistocene climate cycles. Nested clade analysis, coalescent analyses, and analyses of genetic structure support allopatric fragmentation as the cause of the deep divergences within this species.

Africa↗

Biogeography of the túngara frog, Physalaemus pustulosus: a molecular perspective.

Physalaemus pustulosus, a small leptodactylid frog with South American affinities, ranges across northern South America through Middle America to southern Mexico. To investigate its geographic variation and evolutionary origins, we analysed the presumptive gene products of 14 allozyme loci and sequenced a portion of the mitochondrial COI gene from individuals sampled throughout the distribution. Generally, allozyme dissimilarities and sequence divergences are correlated with each other and with geographic proximity. The greatest discontinuity in genetic variation was found between populations in Middle America vs. South America + Panama. Based on two Bayesian MCMC (Markov chain Monte Carlo) divergence time estimates involving two independent temporal constraints, the timing of the separation of northern and southern túngara frog lineages is significantly older than the time since completion of the current Panama land bridge. P. pustulosus first invaded Middle America from South America about 6-10 million years ago giving rise to the northern lineage. The southern lineage then invaded Panama independently after land bridge completion. Despite millions of years of independent evolution, the multilocus allozyme data revealed that western Panama populations represent a contact zone containing individuals with alleles from both groups present.

Animals↗

Phylogeny and biogeography of Croton alabamensis (Euphorbiaceae), a rare shrub from Texas and Alabama, using DNA sequence and AFLP data.

Croton alabamensis (Euphorbiaceae s.s.) is a rare plant species known from several populations in Texas and Alabama that have been assigned to var. texensis and var. alabamensis, respectively. We performed maximum parsimony, maximum likelihood, and Bayesian analyses of DNA sequences from the nuclear ribosomal internal transcribed spacer (ITS) and 5.8S regions and chloroplast trnL-trnF regions from collections of the two varieties of C. alabamensis and from outgroup taxa. C. alabamensis emerges alone on a long branch that is sister to Croton section Corylocroton and the Cuban endemic genus Moacroton. Molecular clock analysis estimates the split of C. alabamensis from its closest relatives in sect. Corylocroton at 41 million years ago, whereas the split of the two varieties of C. alabamensis occurred sometime in the Quaternary. Amplified fragment length polymorphism (AFLP) analyses were performed using two selective primer pairs on a larger sampling of accessions (22 from Texas, 17 from Alabama) to further discriminate phylogenetic structure and quantify genetic diversity. Using both neighbour joining and minimum evolution, the populations from the Cahaba and Black Warrior watersheds in Alabama form two well-separated groups, and in Texas, geographically distinct populations are recovered from Fort Hood, Balcones Canyonlands, and Pace Bend Park. Most of the molecular variance is accounted for by variance within populations. Approximately equal variance is found among populations within states and between states (varieties). Genetic distance between the Texas populations is significantly less than genetic distance between the Alabama populations. Both sequence and AFLP data support the same relationships between the varieties of C. alabamensis and their outgroup, while the AFLP data provide better resolution among the different geographical regions where C. alabamensis occurs. The conservation implications of these findings are discussed.

Alabama↗

Biogeography of the marine actinomycete Salinispora.

Marine actinomycetes belonging to the genus Salinispora were cultured from marine sediments collected at six geographically distinct locations. Detailed phylogenetic analyses of both 16S rRNA and gyrB gene sequences reveal that this genus is comprised of three distinct but closely related clades corresponding to the species Salinispora tropica, Salinispora arenicola and a third species for which the name 'Salinispora pacifica' is proposed. Salinispora arenicola was cultured from all locations sampled and provides clear evidence for the cosmopolitan distribution of an individual bacterial species. The co-occurrence of S. arenicola with S. tropica and S. pacifica suggests that ecological differentiation as opposed to geographical isolation is driving speciation within the genus. All Salinispora strains cultured to date share greater than 99% 16S rRNA gene sequence identity and thus comprise what has been described as a microdiverse ribotype cluster. The description of this cluster as a new genus, containing multiple species, provides clear evidence that fine-scale 16S rDNA sequence analysis can be used to delineate among closely related species and that more conservative operational taxonomic unit values may significantly underestimate global species diversity.

Biodiversity↗

Biogeography of the yeasts of ephemeral flowers and their insects.

We studied specific yeast communities vectored by beetles, drosophilids, and bees that visit ephemeral flowers, mostly in the genus Hibiscus and in the families Convolvulaceae and Cactaceae, in the Neotropical, Nearctic, and Australian biogeographic regions. The communities consist mostly of yeasts in four clades centered around the genera Metschnikowia, Kodamaea, Wickerhamiella, and Starmerella. The largest geographic discontinuity occurs as a function of the nitidulid beetle species that dominate the non-pollinator insect visitors of the flowers. This partitions the New World, where the dominant beetle is in the genus Conotelus, from the Australian biogeographic region, dominated by species of Aethina. Distinct but sympatric insects may also carry radically different yeast communities.

Animals↗

Genetic diversity and biogeography of haloalkaliphilic sulphur-oxidizing bacteria belonging to the genus Thioalkalivibrio.

A group of 85 isolates of haloalkaliphilic obligately chemolithoautotrophic sulphur-oxidizing bacteria belonging to the genus Thioalkalivibrio were recently obtained from soda lakes in Mongolia, Kenya, California, Egypt and Siberia. They have been analyzed by repetitive extragenic palindromic (rep)-PCR genomic fingerprinting technique with BOX- and (GTG)5-primer set. Cluster analysis was performed using combined fingerprint profiles and a dendrogram similarity value (r) of 0.8 was used to define the same genotype. Fifty-six genotypes were found among the isolates, revealing a high genetic diversity. The strains can be divided into two major clusters, including isolates from the Asiatic (Siberia and Mongolia) and the African (Kenya and Egypt) continents, respectively. The majority (85.9%) of the genotypes were found in only one area, suggesting an endemic character of the Thioalkalivibrio strains. Furthermore, a correlation between fingerprint clustering, geographic origin and the characteristics of the lake of origin was found.

Africa↗

Phylogeny of the 1-aminocyclopropane-1-carboxylic acid deaminase-encoding gene acdS in phytobeneficial and pathogenic Proteobacteria and relation with strain biogeography.

Deamination of the ethylene precursor 1-aminocyclopropane-1-carboxylic acid (ACC) is a key plant-beneficial trait found in plant growth-promoting rhizobacteria (PGPR) and phytosymbiotic bacteria, but the diversity of the corresponding gene (acdS) is poorly documented. Here, acdS sequences were obtained by screening putative ACC deaminase sequences listed in databases, based on phylogenetic properties and key residues. In addition, acdS was sought in 71 proteobacterial strains by PCR amplification and/or hybridization using colony dot blots. The presence of acdS was confirmed in established AcdS+ bacteria and evidenced noticeably in Azospirillum (previously reported as AcdS-), in 10 species of Burkholderia and six Burkholderia cepacia genomovars (which included PGPR, phytopathogens and opportunistic human pathogens), and in five Agrobacterium genomovars. The occurrence of acdS in true and opportunistic pathogens raises new questions concerning their ecology in plant-associated habitats. Many (but not all) acdS+ bacteria displayed ACC deaminase activity in vitro, including two Burkholderia clinical isolates. Phylogenetic analysis of partial acdS and deduced AcdS sequences evidenced three main phylogenetic clusters, each gathering pathogens and plant-beneficial strains of contrasting geographic and habitat origins. The acdS phylogenetic tree was only partly congruent with the rrs tree. Two clusters gathered both Betaprotobacteria and Gammaproteobacteria, suggesting extensive horizontal transfers of acdS, noticeably between plant-associated Proteobacteria.

Carbon-Carbon Lyases↗

Evolution and biogeography of deep-sea vent and seep invertebrates.

Deep-sea hydrothermal vents and cold seeps are submarine springs where nutrient-rich fluids emanate from the sea floor. Vent and seep ecosystems occur in a variety of geological settings throughout the global ocean and support food webs based on chemoautotrophic primary production. Most vent and seep invertebrates arrive at suitable habitats as larvae dispersed by deep-ocean currents. The recent evolution of many vent and seep invertebrate species (<100 million years ago) suggests that Cenozoic tectonic history and oceanic circulation patterns have been important in defining contemporary biogeographic patterns.

Animals↗