Founder-event Speciation Research Articles

Molecular phylogenetics suggests a New Guinean origin and frequent episodes of founder-event speciation in the nectarivorous lories and lorikeets (Aves: Psittaciformes)

The lories and lorikeets (Aves: Loriinae: Loriini) are a readily recognizable, discrete group of nectarivorous parrots confined to the Indo-Pacific region between Wallace’s Line and the Pitcairn Island group in the central-east Pacific Ocean. We present the first phylogenetic analysis of all currently recognized genera in the group using two mitochondrial and five nuclear loci. Our analyses suggest a New Guinean origin for the group at about 10million years ago (95% HPD 4.8–14.8) but this origin must be interpreted within the context of that island’s complicated, recent geological history. That is, the origin and early diversification of the group may have taken place as New Guinea’s Central Cordillera arose and the final constituent terranes that form present-day New Guinea were accreted. The latter activity may have promoted dispersal as a key element in the group’s history. We have detected several instances of dispersal out of New Guinea that we argue constitute instances of founder-event speciation. Some phenotypically cohesive genera are affirmed as monophyletic but other genera are clearly in need of taxonomic dismantlement and reclassification. We recognize Parvipsitta Mathews, 1916 for two species usually placed in Glossopsitta and we advocate transfer of Chalcopsitta cardinalis into Pseudeos Peters, 1935. Other non-monophyletic genera such as Charmosyna, Psitteuteles and, probably, Trichoglossus, require improved taxon sampling and further phylogenetic analysis before their systematics can be resolved. Cursory examination of trait mapping across the group suggests that many traits are ancestral and of little use in determining genus-level systematics.

Model selection in historical biogeography reveals that founder-event speciation is a crucial process in Island Clades.

Founder-event speciation, where a rare jump dispersal event founds a new genetically isolated lineage, has long been considered crucial by many historical biogeographers, but its importance is disputed within the vicariance school. Probabilistic modeling of geographic range evolution creates the potential to test different biogeographical models against data using standard statistical model choice procedures, as long as multiple models are available. I re-implement the Dispersal-Extinction-Cladogenesis (DEC) model of LAGRANGE in the R package BioGeoBEARS, and modify it to create a new model, DEC + J, which adds founder-event speciation, the importance of which is governed by a new free parameter, [Formula: see text]. The identifiability of DEC and DEC + J is tested on data sets simulated under a wide range of macroevolutionary models where geography evolves jointly with lineage birth/death events. The results confirm that DEC and DEC + J are identifiable even though these models ignore the fact that molecular phylogenies are missing many cladogenesis and extinction events. The simulations also indicate that DEC will have substantially increased errors in ancestral range estimation and parameter inference when the true model includes + J. DEC and DEC + J are compared on 13 empirical data sets drawn from studies of island clades. Likelihood-ratio tests indicate that all clades reject DEC, and AICc model weights show large to overwhelming support for DEC + J, for the first time verifying the importance of founder-event speciation in island clades via statistical model choice. Under DEC + J, ancestral nodes are usually estimated to have ranges occupying only one island, rather than the widespread ancestors often favored by DEC. These results indicate that the assumptions of historical biogeography models can have large impacts on inference and require testing and comparison with statistical methods.

Diversification in an Afro-Asian songbird clade (Erythropygia–Copsychus) reveals founder-event speciation via trans-oceanic dispersals and a southern to northern colonization pattern in Africa

Erythropygia scrub-robins and their allies are distributed throughout Africa, Europe, Southeast Asia, India, Madagascar and the Seychelles. This broad distribution, as well as the distribution of Erythropygia taxa across Africa, presents an interesting opportunity to explore the mechanisms by which this biogeographic distribution was achieved. Multilocus sequence data (3310 base pairs from two mitochondrial and two nuclear genes) were generated for all species of Erythropygia and Cercotrichas scrub-robins, as well as from genera previously shown to render Erythropygia paraphyletic. Using model-based phylogenetic methods and molecular clock dating, we constructed a time-calibrated molecular phylogenetic hypothesis for the lineage. Ancestral area reconstructions were performed on the phylogeny using probabilistic approaches implemented in LaGrange and BioGeoBEARS. Our results confirm that Erythropygia is not monophyletic, and that one of the two Erythropygia clades is more closely related to a clade of Asian and Indian Ocean islands distributed species. Overall, the Erythropygia and allies clade originated in Africa in the late Miocene c. 6.9 Ma. Subsequently, a number of overwater dispersals occurred to include an initial colonization of Southeast Asia, and an ensuing progression of colonizations from Southeast Asia to the Seychelles, from there to Madagascar, and from these Indian Ocean islands back to Southeast Asia. Within the two clades of Erythropygia, ancestral area reconstructions within Africa indicate a Southern Africa origin, with subsequent lineage divergence in each clade indicating northward colonization. Overall, this clade of non-migratory songbirds shows a remarkable number of trans-oceanic colonization events, that were possibly facilitated by wind-driven dispersal; repeated Africa to Asia colonizations, two of which occur in this clade, are exceptionally rare in birds. Also rare is our finding that colonization patterns in Africa indicate a southern to northern progression.

Probabilistic historical biogeography: new models for founder-event speciation, imperfect detection, and fossils allow improved accuracy and model-testing

Historical biogeography has been characterized by a large diversity of methods and unresolved debates about which processes, such as dispersal or vicariance, are most important for explaining distributions. A new R package, BioGeoBEARS, implements many models in a common likelihood framework, so that standard statistical model selection procedures can be applied to let the data choose the best model. Available models include a likelihood version of DIVA (“DIVALIKE”), LAGRANGE’s DEC model, and BAYAREA, as well as “+J” versions of these models which include founder-event speciation, an important process left out of most inference methods. I use BioGeoBEARS on a large sample of island and non-island clades (including two fossil clades) to show that founder-event speciation is a crucial process in almost every clade, and that most published datasets reject the non-J models currently in widespread use. BioGeoBEARS is open-source and freely available for installation at the Comprehensive R Archive Network at http://CRAN.R-project.org/package=BioGeoBEARS. A step-by-step tutorial is available at http://phylo.wikidot.com/biogeobears.

Probabilistic historical biogeography: new models for founder-event speciation, imperfect detection, and fossils allow improved accuracy and model-testing

Historical biogeography has been characterized by a large diversity of methods and unresolved debates about which processes, such as dispersal or vicariance, are most important for explaining distributions. A new R package, BioGeoBEARS, implements many models in a common likelihood framework, so that standard statistical model selection procedures can be applied to let the data choose the best model. Available models include a likelihood version of DIVA (“DIVALIKE”), LAGRANGE’s DEC model, and BAYAREA, as well as “+J” versions of these models which include founder-event speciation, an important process left out of most inference methods. I use BioGeoBEARS on a large sample of island and non-island clades (including two fossil clades) to show that founder-event speciation is a crucial process in almost every clade, and that most published datasets reject the non-J models currently in widespread use. BioGeoBEARS is open-source and freely available for installation at the Comprehensive R Archive Network at http://CRAN.R-project.org/package=BioGeoBEARS. A step-by-step tutorial is available at http://phylo.wikidot.com/biogeobears.

Testing Founder Effect Speciation: Divergence Population Genetics of the Spoonbills Platalea regia and Pl. minor (Threskiornithidae, Aves)

Although founder effect speciation has been a popular theoretical model for the speciation of geographically isolated taxa, its empirical importance has remained difficult to evaluate due to the intractability of past demography, which in a founder effect speciation scenario would involve a speciational bottleneck in the emergent species and the complete cessation of gene flow following divergence. Using regression-weighted approximate Bayesian computation, we tested the validity of these two fundamental conditions of founder effect speciation in a pair of sister species with disjunct distributions: the royal spoonbill Platalea regia in Australasia and the black-faced spoonbill Pl. minor in eastern Asia. When compared with genetic polymorphism observed at 20 nuclear loci in the two species, simulations showed that the founder effect speciation model had an extremely low posterior probability (1.55 × 10(-8)) of producing the extant genetic pattern. In contrast, speciation models that allowed for postdivergence gene flow were much more probable (posterior probabilities were 0.37 and 0.50 for the bottleneck with gene flow and the gene flow models, respectively) and postdivergence gene flow persisted for a considerable period of time (more than 80% of the divergence history in both models) following initial divergence (median = 197,000 generations, 95% credible interval [CI]: 50,000-478,000, for the bottleneck with gene flow model; and 186,000 generations, 95% CI: 45,000-477,000, for the gene flow model). Furthermore, the estimated population size reduction in Pl. regia to 7,000 individuals (median, 95% CI: 487-12,000, according to the bottleneck with gene flow model) was unlikely to have been severe enough to be considered a bottleneck. Therefore, these results do not support founder effect speciation in Pl. regia but indicate instead that the divergence between Pl. regia and Pl. minor was probably driven by selection despite continuous gene flow. In this light, we discuss the potential importance of evolutionarily labile traits with significant fitness consequences, such as migratory behavior and habitat preference, in facilitating divergence of the spoonbills.

Diversification at high latitudes: speciation of buntings in the genusPlectrophenaxinferred from mitochondrial and nuclear markers

High-latitude diversification is a process characterized by speciation and extinction due to climatically driven vicariance and dispersal events. McKay's buntings (Plectrophenax hyperboreus) are high-latitude island endemic songbirds, and their global range is restricted to Beringia. Snow buntings (P. nivalis), their closest relatives, are distributed throughout the Holarctic, breeding in available habitat surrounding the island range of McKay's buntings. We sequenced 1123 base pairs of mitochondrial DNA for 40 individuals of each species and analysed a total of 913 AFLPs for 57 individuals. Both marker types suggested weak but significant genetic differentiation. Analysis of sequence data indicated divergence occurring when the current breeding range of McKay's buntings was a hill on the Beringian steppe (approximately 18,400 to approximately 73,700 years before present), suggesting that snow buntings were restricted to lower latitudes by ice sheets. Ancestral effective population size estimates indicate a founder event in McKay's buntings followed by an expansion and then a reduction in effective size. Rising sea levels and asymmetric hybridization from McKay's buntings into the postglacially-colonizing population of snow buntings could account for this reduction. Reproductive isolation is likely maintained through differential arrival dates on breeding grounds and the high breeding density of McKay's buntings. This recent, high-latitude divergence best fits a model of founder event speciation driven by vicariance and oscillations in habitat due to climate change.

Nucleotide Variation, Linkage Disequilibrium and Founder-Facilitated Speciation in Wild Populations of the Zebra Finch (Taeniopygia guttata)

The zebra finch has long been an important model system for the study of vocal learning, vocal production, and behavior. With the imminent sequencing of its genome, the zebra finch is now poised to become a model system for population genetics. Using a panel of 30 noncoding loci, we characterized patterns of polymorphism and divergence among wild zebra finch populations. Continental Australian populations displayed little population structure, exceptionally high levels of nucleotide diversity (pi = 0.010), a rapid decay of linkage disequilibrium (LD), and a high population recombination rate (rho approximately 0.05), all of which suggest an open and fluid genomic background that could facilitate adaptive variation. By contrast, substantial divergence between the Australian and Lesser Sunda Island populations (K(ST) = 0.193), reduced genetic diversity (pi = 0.002), and higher levels of LD in the island population suggest a strong but relatively recent founder event, which may have contributed to speciation between these populations as envisioned under founder-effect speciation models. Consistent with this hypothesis, we find that under a simple quantitative genetic model both drift and selection could have contributed to the observed divergence in six quantitative traits. In both Australian and Lesser Sundas populations, diversity in Z-linked loci was significantly lower than in autosomal loci. Our analysis provides a quantitative framework for studying the role of selection and drift in shaping patterns of molecular evolution in the zebra finch genome.

Plant speciation in continental island floras as exemplified byNigellain the Aegean Archipelago

Continental shelf island systems, created by rising sea levels, provide a premier setting for studying the effects of geographical isolation on non-adaptive radiation and allopatric speciation brought about by genetic drift. The Aegean Archipelago forms a highly fragmented complex of mostly continental shelf islands that have become disconnected from each other and the mainland in relatively recent geological times (ca <5.2 Ma). These ecologically fairly homogenous islands thus provide a suitable biogeographic context for assessing the relative influences of past range fragmentation, colonization, gene flow and drift on taxon diversification. Indeed, recent molecular biogeographic studies on the Aegean Nigella arvensis complex, combining phylogenetic, phylogeographic and population level approaches, exemplify the importance of allopatry and genetic drift coupled with restricted gene flow in driving plant speciation in this continental archipelago at different temporal and spatial scales. While the recent (Late Pleistocene) radiation of Aegean Nigella, as well as possible instances of incipient speciation (in the Cyclades), is shown to be strongly conditioned by (palaeo)geographic factors (including changes in sea level), shifts in breeding system (selfing) and associated isolating mechanisms have also contributed to this radiation. By contrast, founder event speciation has probably played only a minor role, perhaps reflecting a migratory situation typical for continental archipelagos characterized by niche pre-emption because of a long established resident flora. Overall, surveys of neutral molecular markers in Aegean Nigella have so far revealed population genetic processes that conform remarkably well to predictions raised by genetic drift theory. The challenge is now to gain more direct insights into the relative importance of the role of genetic drift, as opposed to natural selection, in the phenotypic and reproductive divergence among these Aegean plant species.

Signatures of seaway closures and founder dispersal in the phylogeny of a circumglobally distributed seahorse lineage.

BackgroundThe importance of vicariance events on the establishment of phylogeographic patterns in the marine environment is well documented, and generally accepted as an important cause of cladogenesis. Founder dispersal (i.e. long-distance dispersal followed by founder effect speciation) is also frequently invoked as a cause of genetic divergence among lineages, but its role has long been challenged by vicariance biogeographers. Founder dispersal is likely to be common in species that colonize remote habitats by means of rafting (e.g. seahorses), as long-distance dispersal events are likely to be rare and subsequent additional recruitment from the source habitat is unlikely. In the present study, the relative importance of vicariance and founder dispersal as causes of cladogenesis in a circumglobally distributed seahorse lineage was investigated using molecular dating. A phylogeny was reconstructed using sequence data from mitochondrial and nuclear markers, and the well-documented closure of the Central American seaway was used as a primary calibration point to test whether other bifurcations in the phylogeny could also have been the result of vicariance events. The feasibility of three other vicariance events was explored: a) the closure of the Indonesian Seaway, resulting in sister lineages associated with the Indian Ocean and West Pacific, respectively; b) the closure of the Tethyan Seaway, resulting in sister lineages associated with the Indo-Pacific and Atlantic Ocean, respectively, and c) continental break-up during the Mesozoic followed by spreading of the Atlantic Ocean, resulting in pairs of lineages with amphi-Atlantic distribution patterns.ResultsComparisons of pairwise genetic distances among the seahorse species hypothesized to have diverged as a result of the closure of the Central American Seaway with those of published teleost sequences having the same distribution patterns show that the seahorses were among the last to diverge. This suggests that their cladogenesis was associated with the final closure of this seaway. Although two other divergence events in the phylogeny could potentially have arisen as a result of the closures of the Indonesian and Tethyan seaways, respectively, the timing of the majority of bifurcations in the phylogeny differed significantly from the dates of vicariance events suggested in the literature. Moreover, several divergence events that resulted in the same distribution patterns of lineages at different positions in the phylogeny did not occur contemporaneously. For that reason, they cannot be the result of the same vicariance events, a result that is independent of molecular dating.ConclusionInterpretations of the cladogenetic events in the seahorse phylogeny based purely on vicariance biogeographic hypotheses are problematic. We conclude that the evolution of the circumglobally distributed seahorse lineage was strongly influenced by founder dispersal, and suggest that this mode of speciation may be particularly important in marine organisms that lack a pelagic dispersal phase and instead disperse by means of rafting.

A Test of Founder Effect Speciation Using Multiple Loci in the Auklets (Aethia spp.)

Whether speciation results more frequently from the genetic consequences of founder events or from gradual genetic divergence of large populations is a matter of debate. In this study, multiple analyses were applied to data from three loci (cytochrome b, alpha-enolase intron VIII, and MHC class II B) to test for founder effects associated with speciation in Aethia (Aves: Alcidae), a genus of seabirds thought to have undergone a rapid founder-induced radiation. Effective population sizes (N(e)) were derived from estimators of based on allelic diversity and the coalescent and from data on trans-species polymorphism. Results indicated that N(e) has been on the order of 10(5)-10(6) individuals throughout the evolutionary histories of least and crested auklets (A. pusilla and A. cristatella, respectively) and that N(e) of the ancestral species was at least 16,000 individuals. Computer simulations of MHC evolution indicated that a single-generation bottleneck at speciation could not have involved <85 individuals for each species. More moderate simulation scenarios indicated that population size could not have dropped below 2000 individuals at the time of species founding. Demographic history appears to have been stable for the auklets throughout the past several million years, and a founder effect associated with their speciation is unlikely.

Untangling Coevolutionary History

Organisms in nature exist in complex environments where they interact with a diversity of other species. Just as interactions between species are fundamental to ecological studies, they also are integral to understanding many macroevolutionary patterns of diversification. Chief among these are the coevolutionary patterns apparent between closely interacting groups, such as hosts and parasites, or mutualists. A variety of historical processes play a role in the diversification of interacting taxa. These processes occur on a variety of scales, ranging from the intracellular to the community level, and across a diverse array of taxa, ranging from viruses to vertebrates. Recent phylogenetic studies reveal that the phylogenies of interacting taxa are often very similar, sometimes identical. This similarity is most often generated by repeated instances of cospeciation, the simultaneous speciation of interacting taxa. Alternatively, a number of processes can reduce the congruence between phylogenies of closely interacting taxa. These processes include switching, duplication, sorting events, and failure of one lineage to speciate in response to speciation in the other lineage. Recent cophylogenetic studies have revealed a diversity of patterns, ranging from complete congruence to no significant similarity between phylogenies of interacting taxa (Page, 2003). Coevolutionary biologists can help provide explanations for this diversity by integrating cophylogenetic studies with knowledge of the ecological and physiological mechanisms underlying species interactions. Research on coevolution often merges ecology, systematics, population genetics, physiology, and molecular biology. At the 2002 joint Society of Systematic Biologists (SSB)/Society for the Study of Evolution (SSE) annual meeting we had an opportunity to bring together coevolutionary biologists working on a diverse array of taxa. These workers have summarized their presentations in this issue of Systematic Biology. We are grateful to the SSB for providing support for the symposium and its publication. Our goal has been to inspire comparisons across studies to understand the processes responsible for different patterns of coevolutionary history. As background for this compilation of empirical studies, we review here the processes believed to be important in coevolutionary history (see also Page and Charleston, 1998; Page, 2003). For ease of description, we use hosts and parasites to illustrate these processes, but any pair of interacting taxa could be used. We provide specific examples from a single host-parasite system consisting of pigeons and doves (Aves: Columbiformes) and their parasitic lice (Insecta: Phthiraptera). The first process, cospeciation, is the simultaneous speciation of hosts and parasites. Repeated instances of cospeciation generally increase the congruence between host and parasite phylogenies; cospeciation is usually inferred directly from such congruence. For tightly interacting organisms, some workers consider cospeciation to be a null model of sorts, because forces isolating host populations also tend to isolate their parasite populations. An example of cospeciation can be seen in body lice (Physconelloides) found on New World doves (Fig. 1). One of the processes that reduces the congruence between host and parasite phylogenies is host switching. Incomplete host switching is simply an expansion of a parasite's host range, such as when a parasite colonizes an additional species of host (Fig. 2). Incomplete host switching reduces the parasite's level of host specificity; where a parasite species once occurred on a single host species, it now occurs on two host species. Host switching can be completed either by extinction of the parasite on the original host, speciation between the parasite populations on the new host and those on the original host, or both. Incomplete host switching has occurred in body lice (Physconelloides) of small New World ground doves (Fig. 2). Sorting events are another group of factors that, in combination with other processes, can reduce the similarity between host and parasite phylogenies. Sorting can involve extinction, in which a parasite lineage is lost from a host lineage, or the boat, in which a parasite lineage is missing from a host lineage during founder event speciation. An apparent sorting event has occurred in the wing lice (Columbicola) of the New Zealand pigeon (Hemiphaga novaeseelandiae). Wing lice have never been found on New Zealand pigeons despite intensive sampling (Paterson et al., 1999; Clayton, unpubl. data), whereas the closest relatives of the New Zealand pigeon do have wing lice (Adams, 2002). Yet another factor that can reduce phylogenetic congruence is duplication, which occurs when a parasite lineage speciates in the absence of host speciation. Duplication can occur via sympatric speciation of parasites or via allopatric speciation of parasite populations across the geographic range of a host. Although there are few examples of recent duplication events in lice (Johnson and Clayton, 2003), a duplication event seems to have occurred in the body lice of the Band-tailed

Phylogenetic and biogeographic relationships of the mouse opossum Thylamys (Didelphimorphia, Didelphidae) in southern South America

Nucleotide sequence data from the mitochondrial cytochrome b gene were used to evaluate the phylogenetic relationships among mouse opossum species of the genus Thylamys. Based on approximately 1000 bp in five of the six species of the genus and including different localities for some of the species, we concluded that T. macrura from the subtropical forests of eastern Paraguay is the most primitive taxon. Subsequent radiation of the genus is explained mainly via founder effect speciation. This evolutionary scenario would account for the speciation of T. pusilla, T. venusta, T. pallidior, and T. elegans in the Chaco, southern Bolivia and northern Argentina, the Andean Altiplano, the Coastal Desert of Chile, and coastal Perú, respectively. Calibration of a molecular clock set the Pleistocene as the period for the differentiation of Thylamys species. The molecular results confirm the strong genetic connection between populations that inhabit the “pre-cordillera” of northern Chile ( T. pallidior) and the canyons that run through the Atacama Desert to the lowlands in northern Chile. Our results confirm the occurrence of two Thylamys species in Chile, T. pallidior and T. elegans, within and south to the Atacama Desert, respectively.

The speciation revolution

I predict the years 1990±2010 will be seen as a revolution in the study of speciation. One person's punctuated equilibrium is another's gradual change, and the current revolution is in any case paltry compared with Darwin's own. Even so, many previously accepted beliefs about speciation are now doubted, and features of a classic scienti®c revolution are evident. To see just how much has changed, consider what experts were saying until recently. Coyne (1994), for example, listed four major achievements due to Ernst Mayr since the 1940s. These were: (1) an appreciation of the reality of species (as compared with, say, the unreality of subspecies or genera); (2) the reproductive isolation de®nition of species (the `biological species concept'); (3) the generality of allopatric speciation; and (4) founder-effect speciation. Coyne (1994) argued that Mayr's fourth achievement, the founder effect, was probably incorrect, but regarded the other three as completely in tune with the current view of speciation. What is extraordinary about this list is that all these `achievements' are now, only 7 years later, rejected by major groups of evolutionary biologists. Softening on points (1) and (4) are found today even in papers coauthored by Coyne himself (Kliman et al., 2000; Turelli et al., 2001). The opinions under attack date from the 1930s to the 1950s, and are identi®ed strongly with Theodosius Dobzhansky and Ernst Mayr; however, virtually all evolutionary biologists and most textbooks supported these ideas until the late 1980s. In what follows, I discuss each of Coyne's points in turn.

Genealogical portraits of speciation in montane grasshoppers (genus Melanoplus) from the sky islands of the Rocky Mountains.

Grasshoppers in the genus Melanoplus have undergone a radiation in the 'sky islands' of western North America, with many species originating during the Pleistocene. Despite their recent origins, phylogenetic analyses indicate that all the species exhibit monophyletic or paraphyletic gene trees. The objectives of this study were to determine whether the monophyletic genealogies are the result of a bottleneck at speciation and to investigate the extent to which the different phylogenetic states of eight species (i.e. monophyletic versus paraphyletic gene trees) can be ascribed to the effects of speciation. A coalescent simulation was used to test for a bottleneck at speciation in each species. The effective population sizes and demographic histories of species were compared across taxa to evaluate the possibility that the paraphyly versus monophyly of the species reflects differential rates of lineage loss rather than speciation mode. While coalescent analyses indicate that the monophyly of Melanoplus species might not be indicative of bottlenecks at speciation, the results suggest that the paraphyletic gene trees may reflect the demography of speciation, involving localized divergences in the ancestral species. With respect to different models of Pleistocene divergence, the data do not support a model of founder-effect speciation but are compatible with divergence in allopatric refugia.

EPISTASIS AND THE EVOLUTION OF ADDITIVE GENETIC VARIANCE IN POPULATIONS THAT PASS THROUGH A BOTTLENECK.

Traditional models of genetic drift predict a linear decrease in additive genetic variance for populations passing through a bottleneck. This perceived lack of heritable variance limits the scope of founder-effect models of speciation. We produced 55 replicate bottleneck populations maintained at two male-female pairs through four generations of inbreeding (average F = 0.39). These populations were formed from an F2 intercross of the LG/J and SM/J inbred mouse strains. Two contemporaneous control strains maintained with more than 60 mating pairs per generation were formed from this same source population. The average level of within-strain additive genetic variance for adult body weight was compared between the control and experimental lines. Additive genetic variance for adult body weight within experimental bottleneck strains was significantly higher than expected under an additive genetic model This enhancement of additive genetic variance under inbreeding is likely to be due to epistasis, which retards or reverses the loss of additive genetic variance under inbreeding for adult body weight in this population. Therefore, founder-effect speciation processes may not be constrained by a loss of heritable variance due to population bottlenecks.

Genetic differentiation and speciation in Stomion (Coleoptera: Tenebrionidae): flightless beetles of the Galápagos Islands, Ecuador

Abstract Stomion is a swarm of 13 tenebrionid beetle taxa endemic to the Galapagos Islands. The taxa show marked morphological variation, and all are flightless. They can be divided into three divergent morphological groups based on external sculpturing of the elytra. Eleven taxa were examined electrophoretically at 20 enzyme loci to measure the extent of genetic differentiation which accompanied morphological differentiation and speciation. Stomion are shown to be ideal candidates for allopatric and founder event speciation, with their limited dispersal ability, narrow distributions, and incidence of single-island endemics. Accordingly, individual heterozygosities were low in most taxa, and gene pool divergence was apparent among populations of single taxa occupying different islands. Both interspecific and intraspecific genetic identities were high, with an average I=0.88 and I=0.96, respectively. The relatively young age of the genus and the allopatric mode of speciation may account for high genetic relatedness. No clear relationship was found between allozyme phenotypes and the three morphological groups, thus, the evolution of these traits may have been facilitated by similar selection among islands of similar age and geography. The geographical distributions of the morphological groups can be used to suggest a general pattern of speciation for the group.

Genetic differentiation and speciation inStomion(Coleoptera: Tenebrionidae): flightless beetles of the Galápagos Islands, Ecuador

Stomion is a swarm of 13 tenebrionid beetle taxa endemic to the Galápagos Islands. The taxa show marked morphological variation, and all are flightless. They can be divided into three divergent morphological groups based on external sculpturing of the elytra. Eleven taxa were examined electrophoretically at 20 enzyme loci to measure the extent of genetic differentiation which accompanied morphological differentiation and speciation. Stomion are shown to be ideal candidates for allopatric and founder event speciation, with their limited dispersal ability, narrow distributions, and incidence of single-island endemics. Accordingly, individual heterozygosities were low in most taxa, and gene pool divergence was apparent among populations of single taxa occupying different islands. Both interspecific and intraspecific genetic identities were high, with an average I=0.88 and I=0.96, respectively. The relatively young age of the genus and the allopatric mode of speciation may account for high genetic relatedness. No clear relationship was found between allozyme phenotypes and the three morphological groups, thus, the evolution of these traits may have been facilitated by similar selection among islands of similar age and geography. The geographical distributions of the morphological groups can be used to suggest a general pattern of speciation for the group.

Allelic Diversity and Gene Genealogy at the Self-Incompatibility Locus in the Solanaceae

The self-incompatibility (S) locus of flowering plants offers an example of extreme polymorphism maintained by balancing selection. Estimates of recent and long-term effective population size (Ne) were determined for two solanaceous species by examination of S-allele diversity. Estimates of recent Ne in two solanaceous species differed by an order of magnitude, consistent with differences in the species' ecology. In one species, the evidence was consistent with historical population restriction despite a large recent Ne. In the other, no severe bottleneck was indicated over millions of years. Bottlenecks are integral to founder-event speciation, and loci that are subject to balancing selection can be used to evaluate the frequency of this mode of speciation.

Natural selection and random genetic drift as causes of evolution on islands

The evolutionary processes responsible for adaptation and speciation on islands differ in several ways from those on the mainland. Most attention has been given to the random genetic drift that arises when a population is founded from just a few colonizing genomes. Theoretical obstacles to "founder effect speciation' are discussed, together with recent proposals for avoiding them. It is argued that although certain kinds of epistasis can facilitate the evolution of strong reproductive isolation, this favours divergence by selection as much as by random drift.