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Recombination in Wolbachia.

Wolbachia are widely distributed intracellular bacteria that cause a number of reproductive alterations in their eukaryotic hosts. Such alterations include the induction of parthenogenesis, feminization, cytoplasmic incompatibility, and male killing [1-11]. These important bacteria may play a role in rapid speciation in insects [12-14], and there is growing interest in their potential uses as tools for biological control and genetic manipulation of pests and disease vectors [15-16]. Here, we show recombination in the Wolbachia outer surface protein gene (wsp) between strains of Wolbachia. In addition, we find a possible ecological context for this recombination. Evidence indicates either genetic exchange between Wolbachia in a parasitoid wasp and in the fly that it parasitizes or horizontal transfer of Wolbachia between the parasitoid and the fly, followed by a recombination event. Results have important implications for the evolution of these bacteria and the potential use of Wolbachia in biological control.

Animals↗

Phylogeny of Palaearctic wheatears (genus Oenanthe)--congruence between morphometric and molecular data.

Wheatears of the genus Oenanthe are birds specialized to desert ecosystems in the Palaearctic region from Morocco to China. Although they have been the subject of many morphological and ecological studies, no molecular data have been used to elucidate their phylogenetic relationships, and, their relationships are still debated. Here we use DNA sequences of 1180 bp of two mitochondrial genes, 16S rRNA and cytochrome oxidase subunit I, from 32 individuals from Middle East and North Africa, and Bayesian methods to derive a phylogeny for 11 species of Oenanthe. The resulting tree supported three major clades: (A) O. alboniger, O. chrysopygia, O. lugens, O. finschii, O. leucopyga, O. picata, O. moesta, (B) O. deserti and O. pleschanka; and (C) O. isabellina and O. oenanthe. These results largely differ from previous hypotheses based on analysis of morphological and chromatic characters. However, the two clades (B) and (C) were also supported by a phenetic analysis of new morphometric data presented here, indicating that characters related to colouration and ecology in Oenanthe are more strongly influenced by homoplasy than those of body shape.

Africa, Northern↗

Are flightless Galapaganus weevils older than the Galápagos Islands they inhabit?

The 15 species in the weevil genus Galapaganus Lanteri 1992 (Entiminae: Curculionidae: Coleoptera) are distributed on coastal Perú and Ecuador and include 10 flightless species endemic to the Galápagos islands. These beetles thus provide a promising system through which to investigate the patterns and processes of evolution on Darwin's archipelago. Sequences of the mtDNA locus encoding cytochrome oxidase subunit I (COI) were obtained from samples of seven species occurring in different ecological zones of the oldest south-eastern islands: San Cristóbal, Española and Floreana, and the central island Santa Cruz. The single most parsimonious tree obtained shows two well-supported clades that correspond to the species groups previously defined by morphological characters. Based on a mtDNA clock calibrated for arthropods, the initial speciation separating the oldest species, G. galapagoensis (Linell) on the oldest island, San Cristóbal, from the remaining species in the Galápagos occurred about 7.2 Ma. This estimate exceeds geological ages of the extant emerged islands, although it agrees well with molecular dating of endemic Galápagos iguanas, geckos and lizards. An apparent explanation for the disagreement between geological and molecular time-frames is that about 7 Ma there were emerged islands which subsequently disappeared under ocean waters. This hypothesis has gained support from the recent findings of 11-Myr-old submarine seamounts (sunken islands), south-east of the present location of the archipelago. Some species within the darwini group may have differentiated on the extant islands, 1-5 Ma.

Animals↗

Reproductive isolation between Pogonomyrmex rugosus and two lineages with genetic caste determination.

Hybrid speciation occurs when combination of two interspecific genomes results in individuals that are of high fitness but reproductively incompatible with the parental species. Although hybrid speciation is a relatively common source of new species in plants, it appears to be a much rarer occurrence in animal taxa. Here we report on reproductive isolation and range overlap between the rough harvester ant Pogonomyrmex rugosus and two lineages with hybrid genotypes (H 1 and H2). Both lineages obligately interbreed and produce genetically distinct queen and worker offspring, a phenomenon referred to as genetic caste determination (GCD). Diploid offspring produced by gametes of the same lineage develop only into queens, whereas diploid offspring derived from gametes of distinct lineages develop into workers. We investigated small-scale patterns of gene flow between the parent and the two H lineages by sampling along an 80-km transect between a pure P. rugosus population and a two-lineage population. Microsatellite and mitochondrial markers both indicated virtually no gene flow between the parent species and either lineage even at sites where parental and H-lineage colonies co-occurred. The geographic ranges of the parental species and the two-lineage population were essentially parapatric, with a surprisingly narrow band of overlap and evidence of spatial structuring even at microgeographic scales within the transition zone. This suggests that ecological competition with the parent species plays a significant role in determining the evolutionary persistence and current distribution of the hybrid lineages and the genetic caste system.

Animals↗

Phylogenetics of Cancer crabs (Crustacea: Decapoda: Brachyura).

We used morphological, mitochondrial DNA sequence, paleontological, and biogeographical information to examine the evolutionary history of crabs of the genus Cancer. Phylogenies inferred from adult morphology and DNA sequence of the cytochrome oxidase I (COI) gene were each well resolved and well supported, but differed substantially in topology. Four lines of evidence suggested that the COI data set accurately reflected Cancer phylogeny: (1) in the phylogeny inferred from morphological data, each Atlantic species was sister taxon to an ecologically similar Pacific species, suggesting convergence in morphology; (2) a single trans-Arctic dispersal event, as indicated by the phylogeny inferred from COI, is more parsimonious than two such dispersal events, as inferred from morphology; (3) test and application of a maximum likelihood molecular clock to the COI data yielded estimates of origin and speciation times that fit well with the fossil record; and (4) the tree inferred from the combined COI and morphology data was closely similar to the trees inferred from COI, although notably less well supported by the bootstrap. The phylogeny inferred from maximum likelihood analysis of COI suggested that Cancer originated in the North Pacific in the early Miocene, that the Atlantic species arose from a North Pacific ancestor, and that Cancer crabs invaded the Atlantic from the North Pacific 6-12 mya. This inferred invasion time is notably prior to most estimates of the date of submergence of the Bering Strait and the trans-Arctic interchange, but it agrees with fossil evidence placing at least one Cancer species in the Atlantic about 8 mya.

Animals↗

Identification of a host-associated species complex using molecular and morphometric analyses, with the description of Gyrodactylus rugiensoides n. sp. (Gyrodactylidae, Monogenea).

Gyrodactylus rugiensis was originally described as a parasite occurring on the marine gobies Pomatoschistus minutus and Pomatoschistus microps. In our preliminary survey this species was also frequently found on Pomatoschistus pictus and Pomatoschistus lozanoi. Subsequent molecular analysis of the internal transcribed spacers rDNA region revealed that this parasite actually represents a complex of two apparently cryptic species, one restricted to P. microps and the other shared by P. minutus, P. lozanoi and P. pictus. Morphometric analyses were conducted on 17 features of the opisthaptoral hard parts of specimens collected from all four host species. Standard discriminant analysis showed a clear separation of both genotypes by significant differences in marginal hook and ventral bar features. Statistical classifiers (linear discriminant analysis and nearest neighbours) resulted in an estimated misclassification rate of 4.7 and 3.1%, respectively. Based on molecular, morphological and statistical analyses a new species, Gyrodactylus rugiensoides is described. This species seems to display a lower host-specificity than generally observed for Gyrodactylus species as it infects three sympatric host species.However, seasonal and host-dependent morphometric variation is shown for G. rugiensoides collected on P. pictus. Host-switching and gene flow might be important factors preventing speciation on closely related and sympatric host species. The presence of host associated species complexes in this Gyrodactylus-Pomatoschistus system is also confirmed by the presence of two host-dependent genotypes within G. micropsi found on P. minutus and P. lozanoi, and P. microps, respectively. By comparing host and parasite phylogeny, phylogenetic and ecological factors influencing host-specificity are discussed.

Animals↗

Raciation and speciation in house mice from the Alps: the role of chromosomes.

There are at least 24 different karyotypic races of house mouse in the central Alps, each characterized by a different complement of ancestral acrocentric and derived metacentric chromosomes; altogether 55 different metacentric chromosomes have been described from the region. We argue that this chromosome variation largely arose in situ. If these races were to make contact, in most cases they would produce F1 hybrids with substantial infertility (sometimes complete sterility), due to nondisjunction and germ cell death associated with the formation of long-chain and/or ring configurations at meiosis. We present fertility estimates to confirm this for two particular hybrid types, one of which demonstrates male-limited sterility (in accordance with Haldane's Rule). As well as a model for speciation in allopatry, the Alpine mouse populations are of interest with regards speciation in parapatry: we discuss a possible reinforcement event. Raciation of house mice appears to have happened on numerous occasions within the central Alps. To investigate one possible source of new karyotypic races, we use a two-dimensional stepping stone model to examine the generation of recombinant races within chromosomal hybrid zones. Using field-derived ecological data and laboratory-derived fertility estimates, we show that hybrid karyotypic races can be generated at a reasonable frequency in simulations. Our model complements others developed for flowering plants that also emphasize the potential of chromosomal hybrid zones in generating new stable karyotypic forms.

Alleles↗

Ecotoxicology of selenium in freshwater systems.

The toxicology, environmental impacts and risk assessment of Se in freshwater systems are a high priority for research and regulatory agencies. However, understanding Se in freshwater systems is a challenging endeavor. The accurate risk assessment and determination of a water-quality criterion for any freshwater ecosystem are difficult for many reasons. First, the understanding of the structure and energy dynamics in ecosystems is limited. Second, knowledge of Se cycling and transformations in aquatic ecosystems is rudimentary. Third, the role of various environmental factors affecting its bioaccumulation, biotransformation, and toxicology in aquatic ecosystems is largely unknown. Fourth, the extrapolation and use of existing data in commonly used formulas for risk assessment and calculations of water quality criteria is difficult because of the demonstrated species differences in the bioaccumulation, metabolism, and tolerance of Se. There are many aspects of Se ecotoxicology that need to be addressed to develop more accurate environmental risk assessments and determine appropriate water-quality criteria to protect aquatic ecosystems. Studies evaluating the biochemical speciation of Se in aquatic ecosystems, and determination of the role of microbial communities in its cycling, bioaccumulation, transformation, transfer, and toxicity in aquatic systems, appear to be priorities for future research. These would include a broad exploration of Se effects on ecosystems, e.g., exposure regime, direct and indirect biologic effects, and ecosystem level effects. There are, however, ecological uncertainties that tend to confound such endeavors, e.g., insufficient data, extrapolation issues, and environmental stochasticity (Harwell and Harwell 1989). There are several concepts concerning the ecotoxicology of Se that can be stated. Elevated concentrations have degraded many freshwater ecosystems throughout the United States, and additional systems are expected to be affected as anthropogenic activities increasingly mobilize Se into aquatic systems. Se is a very toxic essential trace element. Toxic threshold concentrations in water, dietary items, and tissues are only 2-5 times normal background concentrations. Toxicity in freshwater ecosystems is the result of bioaccumulation, biotransformation, and cycling of Se in aquatic food chains. Thus, environmental risk assessment and development of effective water-quality criteria to protect freshwater ecosystems become formidable tasks.

Animals↗

Closing of the Tethys Sea and the phylogeny of Eurasian killifishes (Cyprinodontiformes: Cyprinodontidae).

To test vicariant speciation hypotheses derived from geological evidence of the closing of the Tethys Sea, we reconstruct phylogenetic relationships of the predominantly fresh-water killifish genus Aphanius using 3263 aligned base pairs of mitochondrial DNA from samples representing 49 populations of 13 species. We use additional 11 cyprinodontid species as outgroup taxa. Genes analysed include those encoding the partial 12S and 16S ribosomal RNAs; transfer RNAs for valine, leucine, isoleucine, glutamine, methionine, tryptophan, alanine, asparagine, cysteine and tyrosine; and complete nicotinamide adenine dinucleotide dehydrogenase subunit I and II. Molecular substitution rate for this DNA region is estimated at of 8.6 +/- 0.1 x 10(-9) substitutions base pair(-1) year(-1), and is derived from a well dated transgression of the Red Sea into the Wadi Sirhan of Jordan 13 million years ago; an alternate substitution rate of 1.1 +/- 0.2 x 10(-8) substitutions base pair(-1) year(-1) is estimated from fossil evidence. Aphanius forms two major clades which correspond to the former eastern and western Tethys Sea. Within the eastern clade Oligocene divergence into a fresh-water clade inhabiting the Arabian Peninsula and an euhaline clade inhabiting coastal area from Pakistan to Somalia is observed. Within the western Tethys Sea clade we observe a middle Oligocene divergence into Iberian Peninsula and Atlas Mountains, and Turkey and Iran sections. Within Turkey we observe a large amount of genetic differentiation correlated with late Miocene orogenic events. Based on concordance of patterns of phylogenetic relationships and area relationships derived from geological and fossil data, as well as temporal congruence of these patterns, we support a predominantly vicariant-based speciation hypothesis for the genus Aphanius. An exception to this pattern forms the main clade of A. fasciatus, an euhaline circum-Mediterranean species, which shows little genetic differentiation or population structuring, thus providing no support for the hypothesis of vicariant differentiation associated with the Messinian Salinity Crisis. The two phylogenetically deepest events were also likely driven by ecological changes associated with the closing of the Tethys Sea.

Animals↗

Evolution of MHC class II beta chain-encoding genes in the Lake Tana barbel species flock (Barbus intermedius complex).

Major histocompatibility complex (MHC) class II protein polymorphism is maintained in allelic lineages which evolve in a trans-specific manner, passing from one species to descendant species. Selection pressure on peptide binding residues should be greatest during speciation, when organisms move into new environments and their MHC molecules encounter new pathogens. The isolation of MHC genes from teleost fishes, the most diverse group of vertebrates, has created possibilities for testing this hypothesis. The large barbels of Lake Tana have undergone an adaptive radiation within the last 5 million years, producing 14 morphotypes which inhabit different ecological niches within the lake. We studied the variability in class II beta chain-encoding genes of four of these morphotypes using polymerase chain reaction amplification and DNA sequencing. The sequences obtained were orthologous to four of the known class II genes from the common carp, from which barbels diverged approximately 32 million years ago. When subjected to phylogenetic analysis, the 48 sequences clustered into groups which represent allelic lineages. A comparison of nonsynonymous and synonymous substitutions between the peptide binding region codons and non-peptide binding region codons of these sequences revealed that they are under strong selective pressure.

Amino Acid Sequence↗

Experimental infections of the monogenean Gyrodactylus turnbulli indicate that it is not a strict specialist.

Parasites represent a threat to endangered fish species, particularly when the parasite can host switch and the new host is vulnerable. If the parasite is highly host specific then successful host switching should be a rare occurrence; however, the host range of many parasites which are assumed to be specialists has never been tested. This includes the monogenean Gyrodactylus turnbulli, a well-studied ectoparasite found caudally on its known host, the guppy, Poecilia reticulata. In this study, we monitored parasite establishment and reproduction on a range of poeciliids and more distantly related fish. Individually maintained fish were experimentally infected with a single parasite and monitored daily to establish whether G. turnbulli could survive and reproduce on other fish species. Gyrodactylus turnbulli can infect a wider range of hosts than previously considered, highlighting the fact that host specificity can never be assumed unless experimentally tested. Our findings also have significant implications for parasite transmission to novel hosts and provide further insight into the evolutionary origins of this ubiquitous group of fish pathogens. Previous molecular evidence indicates that host switching is the key mechanism for speciation within the genus Gyrodactylus. Until recently, most Gyrodactylus spp. were assumed to be narrowly host specific. However, our findings suggest that even so-called specialist species, such as G. turnbulli, may represent a threat to vulnerable fish stocks. In view of the potential importance of host switching under artificial conditions, we propose to describe this as 'artificial ecological transfer' as opposed to 'natural ecological transfer', host switching under natural conditions.

Animals↗

Evolution driven by differential dispersal within a wild bird population.

Evolutionary theory predicts that local population divergence will depend on the balance between the diversifying effect of selection and the homogenizing effect of gene flow. However, spatial variation in the expression of genetic variation will also generate differential evolutionary responses. Furthermore, if dispersal is non-random it may actually reinforce, rather than counteract, evolutionary differentiation. Here we document the evolution of differences in body mass within a population of great tits, Parus major, inhabiting a single continuous woodland, over a 36-year period. We show that genetic variance for nestling body mass is spatially variable, that this generates different potential responses to selection, and that this diversifying effect is reinforced by non-random dispersal. Matching the patterns of variation, selection and evolution with population ecological data, we argue that the small-scale differentiation is driven by density-related differences in habitat quality affecting settlement decisions. Our data show that when gene flow is not homogeneous, evolutionary differentiation can be rapid and can occur over surprisingly small spatial scales. Our findings have important implications for questions of the scale of adaptation and speciation, and challenge the usual treatment of dispersal as a force opposing evolutionary differentiation.

Aging↗

Resource partition history and evolutionary specialization of subunits in complex systems.

Specialization for part of the resources available is a common adaptation leading to diversification in biological systems. In complex ecological communities, the history of diversification of parts of a system correlates with the structure of the system in present times. Abundances and sequences of resource partition of species expected by a sequential broken stick model are compared with observed abundances of species and their sequences of evolutionary divergence. Where resources are abundant, first resource partitions correspond to most ancient speciations, and last to recent speciations. The inverse is true when resources are scarce, agreeing with the occupation of extreme habitats by recent species. Diverging species seem to specialize by dividing the niche of common ancestors. Similar trends exist inside genera as well as between families, suggesting similar patterns in diversification and specialization at other levels of biological organization, such as evolution of functional complexes of macromolecules and organs embryogenesis.

Animals↗

Chloroplast DNA evidence of colonization, adaptive radiation, and hybridization in the evolution of the Macaronesian flora.

Most evolutionary studies of oceanic islands have focused on the Pacific Ocean. There are very few examples from the Atlantic archipelagos, especially Macaronesia, despite their unusual combination of features, including a close proximity to the continent, a broad range of geological ages, and a biota linked to a source area that existed in the Mediterranean basin before the late Tertiary. A chloroplast DNA (cpDNA) restriction site analysis of Argyranthemum (Asteraceae: Anthemideae), the largest endemic genus of plants of any volcanic archipelago in the Atlantic Ocean, was performed to examine patterns of plant evolution in Macaronesia. cpDNA data indicated that Argyranthemum is a monophyletic group that has speciated recently. The cpDNA tree showed a weak correlation with the current sectional classification and insular distribution. Two major cpDNA lineages were identified. One was restricted to northern archipelagos--e.g., Madeira, Desertas, and Selvagens--and the second comprised taxa endemic to the southern archipelago--e.g., the Canary Islands. The two major radiations identified in the Canaries are correlated with distinct ecological habitats; one is restricted to ecological zones under the influence of the northeastern trade winds and the other to regions that are not affected by these winds. The patterns of phylogenetic relationships in Argyranthemum indicate that interisland colonization between similar ecological zones is the main mechanism for establishing founder populations. This phenomenon, combined with rapid radiation into distinct ecological zones and interspecific hybridization, is the primary explanation for species diversification.

Journal Article↗

Evolution in Darwin's finches: a review of a study on Isla Daphne Major in the Galápagos archipelago.

This paper reviews research pertaining to the problem of speciation of the finches on the Galápagos archipelago carried out by assistants, several colleagues, Peter Grant and myself. I give a brief history of the radiation, examine the process of divergence by natural selection over time, and then consider the nature of the reproductive barrier to gene flow between closely related species. Fluctuating climatic conditions have produced a continuously changing ecological landscape and altered feeding conditions for the finches over the last 30 years. Finch populations tracked these changes by natural selection and evolutionary responses to the main events. At each event significant morphological change occurred from one generation to the next generation. As a consequence of these accumulated changes, the mean bill shape and body size of the Geospiza fortis and G. scandens populations differed markedly from 1973 to 2002. Song, a learned culturally transmitted trait, acted as a barrier to reproduction between these species. Rare incidences of misimprinting on song led to hybridization and introgression. Low levels of gene flow from one species to another increased genetic variation on which selection acted. Although the major driving force of diversification was ecological change, the process of diversification involved a subtle interplay between ecology, genetic evolution and learned culturally transmitted traits. An important message for conservation is that neither the environment nor species are fixed entities, therefore a wise strategy for conserving endangered species should keep them capable of further change.

Journal Article↗

Genome evolution and biodiversity in teleost fish.

Teleost fish, which roughly make up half of the extant vertebrate species, exhibit an amazing level of biodiversity affecting their morphology, ecology and behaviour as well as many other aspects of their biology. This huge variability makes fish extremely attractive for the study of many biological questions, particularly of those related to evolution. New insights gained from different teleost species and sequencing projects have recently revealed several peculiar features of fish genomes that might have played a role in fish evolution and speciation. There is now substantial evidence that a round of tetraploidization/rediploidization has taken place during the early evolution of the ray-finned fish lineage, and that hundreds of duplicate pairs generated by this event have been maintained over hundreds of millions of years of evolution. Differential loss or subfunction partitioning of such gene duplicates might have been involved in the generation of fish variability. In contrast to mammalian genomes, teleost genomes also contain multiple families of active transposable elements, which might have played a role in speciation by affecting hybrid sterility and viability. Finally, the amazing diversity of sex determination systems and the plasticity of sex chromosomes observed in teleost might have been involved in both pre- and postmating reproductive isolation. Comparison of data generated by current and future genome projects as well as complementary studies in other species will allow one to approach the molecular and evolutionary mechanisms underlying genome diversity in fish, and will certainly significantly contribute to our understanding of gene evolution and function in humans and other vertebrates.

Animals↗