Sympatric speciation without borders?

The biogeography of speciation remains a controversial issue and the process of allopatric speciation reigns. Sympatric speciation differs from allopatric speciation in terms of geographic setting and the role of selection in bringing about reproductive isolating mechanisms, making it a particularly fascinating and controversial subject for evolutionary biologists. Mayr (1947) explained the difference eloquently: for allopatric speciation, populations spatially diverge and then become reproductively isolated; for sympatric speciation, populations first become reproductively isolated and then diverge. Because of this, sympatric speciation is difficult to show empirically and most evolutionary biologists agree that strict ecological, evolutionary, and geographic criteria must be met ( Coyne & Orr 2004 ). In this issue, Crow et al. (2010) challenge us to expand the definition of sympatric speciation by studying species of marine fishes that they propose have arisen by sympatric speciation in a setting that does not appear to conform to the usual geographical criteria.     

Inter‐specific gene flow dynamics during the Pleistocene‐dated speciation of forest‐dependent mosquitoes in Southeast Asia

Tropical forests have undergone repeated fragmentation and expansion during Pleistocene glacial and interglacial periods, respectively. The effects of this repeated forest fragmentation in driving vicariance in tropical taxa have been well studied. However, relatively little is known about how often this process results in allopatric speciation, since it may be inhibited by recurrent gene flow during repeated secondary contact, or to what extent Pleistocene‐dated speciation results from ecological specialization in the face of gene flow. Here, divergence times and gene flow between three closely‐related mosquito species of the Anopheles dirus species complex endemic to the forests of Southeast Asia, are inferred using coalescent based Bayesian analysis. An Isolation with Migration model is applied to sequences of two mitochondrial and three nuclear genes, and 11 microsatellites. The divergence of An. scanloni has occurred despite unidirectional nuclear gene flow from this species into An. dirus. The inferred asymmetric gene flow may result from the unique evolutionary adaptation of An. scanloni to limestone karst habitat, and therefore the fitness advantage of this species over An. dirus in regions of sympatry. Mitochondrial introgression has led to the complete replacement of An. dirus haplotypes with those of An. baimaii through a recent (～62 kya) selective sweep. Speciation of An. baimaii and An. dirus is inferred to have involved allopatric divergence throughout much of the Pleistocene. Secondary contact and bidirectional gene flow has occurred only within the last 100 000 years, by which time the process of allopatric speciation seems to have been largely completed.     

DOES NICHE DIVERGENCE ACCOMPANY ALLOPATRIC DIVERGENCE IN APHELOCOMA JAYS AS PREDICTED UNDER ECOLOGICAL SPECIATION?: INSIGHTS FROM TESTS WITH NICHE MODELS

The role of ecology in the origin of species has been the subject of long‐standing interest to evolutionary biologists. New sources of spatially explicit ecological data allow for large‐scale tests of whether speciation is associated with niche divergence or whether closely related species tend to be similar ecologically (niche conservatism). Because of the confounding effects of spatial autocorrelation of environmental variables, we generate null expectations for niche divergence for both an ecological‐niche modeling and a multivariate approach to address the question: do allopatrically distributed taxa occupy similar niches? In a classic system for the study of niche evolution—the Aphelocoma jays—we show that there is little evidence for niche divergence among Mexican Jay (A. ultramarina) lineages in the process of speciation, contrary to previous results. In contrast, Aphelocoma species that exist in partial sympatry in some regions show evidence for niche divergence. Our approach is widely applicable to the many cases of allopatric lineages in the beginning stages of speciation. These results do not support an ecological speciation model for Mexican Jay lineages because, in most cases, the allopatric environments they occupy are not significantly more divergent than expected under a null model.     

Inferring the mode of speciation in Indo-West Pacific Conus (Gastropoda: Conidae)

Aim This study aims to initially explore the mode of speciation in Indo‐West Pacific Conus. Location The Indo‐West Pacific island arc, Indian and Pacific Oceans. Methods Relating evolutionary divergence in a molecular phylogeny [T.F. Duda & S.R. Palumbi (1999) Proceedings of the National Academy of Science USA, 96 , 10272] using node height with modern range extents as a possible measure of allopatric or sympatric speciation following that of T.G. Barraclough, A.P. Vogler & P.H. Harvey [(1999) Evolution of Biological Diversity. Oxford University Press, Oxford] models of sympatric and allopatric speciation. Results The analysis seems to indicate that the relationship of sympatry with node height is not informative. Species that have diverged quite recently show 100% sympatry with the sister species. A clearer signal of recent allopatric speciation is observed in species whose distribution is at the edge of the Indian and Pacific Ocean basins. In the widely distributed Conus ebraeus clade, the relationships of node heights and range extents of the member species support a key prediction of sympatric speciation. In highly ecologically specialized species, there is a smaller degree of sympatry than those species that are less specialized. Main conclusions The modes of speciation models presented in this study are not informative. This suggests that there had been large and possibly rapid changes in range size after speciation in the various clades. This could have been due to the fact that the wide dispersal life‐history strategy in the genus had been largely conserved in Conus evolution. There is evidence of sympatric and parapatric speciation in one Conus clade. Overall, the patterns of phylogeny and range distribution when related to the timing of speciation lend circumstantial support to a Neogene centre of origin hypothesis but not to speciation on the Pacific Plate. Speciation is likely to have been associated with the Tethys Sea closure event, with rapid speciation occurring after closure.     

Spatial and ecological population genetic structures within two island‐endemic Aeonium species of different niche width

The Crassulacean genus Aeonium is a well‐known example for plant species radiation on oceanic archipelagos. However, while allopatric speciation among islands is documented for this genus, the role of intra‐island speciation due to population divergence by topographical isolation or ecological heterogeneity has not yet been addressed. The aim of this study was to investigate intraspecific genetic structures and to identify spatial and ecological drivers of genetic population differentiation on the island scale. We analyzed inter simple sequence repeat variation within two island‐endemic Aeonium species of La Palma: one widespread generalist that covers a large variety of different habitat types (Ae. davidbramwellii) and one narrow ecological specialist (Ae. nobile), in order to assess evolutionary potentials on this island. Gene pool differentiation and genetic diversity patterns were associated with major landscape structures in both species, with phylogeographic implications. However, overall levels of genetic differentiation were low. For the generalist species, outlier loci detection and loci–environment correlation approaches indicated moderate signatures of divergent selection pressures linked to temperature and precipitation variables, while the specialist species missed such patterns. Our data point to incipient differentiation among populations, emphasizing that ecological heterogeneity and topographical structuring within the small scales of an island can foster evolutionary processes. Very likely, such processes have contributed to the radiation of Aeonium on the Canary Islands. There is also support for different evolutionary mechanisms between generalist and specialist species.     

Patterns and processes in the evolutionary history of parrotfishes (Family Labridae)

Phylogenetic reconstruction of the evolutionary relationships among 61 of the 70 species of the parrotfish genera Chlorurus and Scarus (Family Labridae) based on mitochondrial and nuclear gene sequences retrieved 15 well‐supported clades with mid Pliocene/Pleistocene diversification. Twenty‐two reciprocally monophyletic sister‐species pairs were identified: 64% were allopatric, and the remainder were sympatric. Age of divergence was similar for allopatric and sympatric species pairs. Sympatric sister pairs displayed greater divergence in morphology, ecology, and sexually dimorphic colour patterns than did allopatric pairs, suggesting that both genetic drift in allopatric species pairs and ecologically adaptive divergence between members of sympatric pairs have played a role in diversification. Basal species typically have small geographical ranges and are restricted to geographically and ecologically peripheral reef habitats. We found little evidence that a single dominant process has driven diversification, nor did we detect a pattern of discrete, sequential stages of diversification in relation to habitat, ecology, and reproductive biology. The evolution of Chlorurus and Scarus has been complex, involving a number of speciation processes. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 107 , 529–557.     

Ecological divergence and speciation between lemur (Eulemur) sister species in Madagascar

Understanding ecological niche evolution over evolutionary timescales is crucial to elucidating the biogeographic history of organisms. Here, we used, for the first time, climate‐based ecological niche models (ENMs) to test hypotheses about ecological divergence and speciation processes between sister species pairs of lemurs (genus Eulemur) in Madagascar. We produced ENMs for eight species, all of which had significant validation support. Among the four sister species pairs, we found nonequivalent niches between sisters, varying degrees of niche overlap in ecological and geographic space, and support for multiple divergence processes. Specifically, three sister‐pair comparisons supported the null model that niches are no more divergent than the available background region. These findings are consistent with an allopatric speciation model, and for two sister pairs (E. collaris–E. cinereiceps and E. rufus–E. rufifrons), a riverine barrier has been previously proposed for driving allopatric speciation. However, for the fourth sister pair E. flavifrons–E. macaco, we found support for significant niche divergence, and consistent with their parapatric distribution on an ecotone and the lack of obvious geographic barriers, these findings most strongly support a parapatric model of speciation. These analyses thus suggest that various speciation processes have led to diversification among closely related Eulemur species.     

The role of head shape and trophic variation in the diversification of the genus Herichthys in sympatry and allopatry

In the present study we evaluated the putative cases of sympatric speciation in the genus Herichthys by studying the variation in head shape using principal component analysis, phylomorphospace and reconstructions of the ancestral states of feeding preferences. Herichthys includes both allopatric and sympatric sister species, as well as sympatric unrelated species and thus offers great potential for evolutionary studies of putatively sympatric speciation. Herichthys is the northernmost group of cichlids in America and one of the most ecologically disparate genera within Middle American cichlids. Fifteen anatomical points were recorded on the heads of 293 specimens of the 11 species recognized within the genus. The results show that in spite of having wide variation in consumed diets, most species of Herichthys are close in morphospace. However, morphological variation was great among the two pairs of sympatric sister species in agreement with the suggested sympatric model of speciation.     

Genomic signatures of divergent selection and speciation patterns in a ‘natural experiment’, the young parallel radiations of Nicaraguan crater lake cichlid fishes

Divergent selection is the main driving force in sympatric ecological speciation and may also play a strong role in divergence between allopatric populations. Characterizing the genome‐wide impact of divergent selection often constitutes a first step in unravelling the genetic bases underlying adaptation and ecological speciation. The Midas cichlid fish (Amphilophus citrinellus) species complex in Nicaragua is a powerful system for studying evolutionary processes. Independent colonizations of isolated young crater lakes by Midas cichlid populations from the older and great lakes of Nicaragua resulted in the repeated evolution of adaptive radiations by intralacustrine sympatric speciation. In this study we performed genome scans on two repeated radiations of crater lake species and their great lake source populations (1030 polymorphic AFLPs, n ～ 30 individuals per species). We detected regions under divergent selection (0.3% in the crater lake Xiloá flock and 1.7% in the older crater lake Apoyo radiation) that might be responsible for the sympatric diversifications. We find no evidence that the same genomic regions have been involved in the repeated evolution of parallel adaptations across crater lake flocks. However, there is some genetic parallelism apparent (seven out of 51 crater lake to great lake outlier loci are shared; 13.7%) that is associated with the allopatric divergence of both crater lake flocks. Interestingly, our results suggest that the number of outlier loci involved in sympatric and allopatric divergence increases over time. A phylogeny based on the AFLP data clearly supports the monophyly of both crater lake species flocks and indicates a parallel branching order with a primary split along the limnetic‐benthic axis in both radiations.     

Recurrent adaptation in a low‐dispersal trait

The study of natural populations from contrasting environments has greatly enhanced our understanding of ecological‐dependent selection, adaptation and speciation. Cases of parallel evolution in particular have facilitated the study of the molecular and genetic basis of adaptive variation. This includes the type and number of genes underlying adaptive traits, as well as the extent to which these genes are exchanged among populations and contribute repeatedly to parallel evolution. Yet, surprisingly few studies provide a comprehensive view on the evolutionary history of adaptive traits from mutation to widespread adaptation. When did key mutations arise, how did they increase in frequency, and how did they spread? In this issue of Molecular Ecology, Van Belleghem et al. ( 2015 ) reconstruct the evolutionary history of a gene associated with wing size in the salt marsh beetle Pogonus chalceus. Screening the entire distribution range of this species, they found a single origin for the allele associated with the short‐winged ecotype. This allele seemingly evolved in an isolated population and rapidly introgressed into other populations. These findings suggest that the adaptive genetic variation found in sympatric short‐ and long‐winged populations has an allopatric origin, confirming that allopatric phases may be important at early stages of speciation.     

Genetic Tests for Ecological and Allopatric Speciation in Anoles on an Island Archipelago

From Darwin''s study of the Galapagos and Wallace''s study of Indonesia, islands have played an important role in evolutionary investigations, and radiations within archipelagos are readily interpreted as supporting the conventional view of allopatric speciation. Even during the ongoing paradigm shift towards other modes of speciation, island radiations, such as the Lesser Antillean anoles, are thought to exemplify this process. Geological and molecular phylogenetic evidence show that, in this archipelago, Martinique anoles provide several examples of secondary contact of island species. Four precursor island species, with up to 8 mybp divergence, met when their islands coalesced to form the current island of Martinique. Moreover, adjacent anole populations also show marked adaptation to distinct habitat zonation, allowing both allopatric and ecological speciation to be tested in this system. We take advantage of this opportunity of replicated island coalescence and independent ecological adaptation to carry out an extensive population genetic study of hypervariable neutral nuclear markers to show that even after these very substantial periods of spatial isolation these putative allospecies show less reproductive isolation than conspecific populations in adjacent habitats in all three cases of subsequent island coalescence. The degree of genetic interchange shows that while there is always a significant genetic signature of past allopatry, and this may be quite strong if the selection regime allows, there is no case of complete allopatric speciation, in spite of the strong primae facie case for it. Importantly there is greater genetic isolation across the xeric/rainforest ecotone than is associated with any secondary contact. This rejects the development of reproductive isolation in allopatric divergence, but supports the potential for ecological speciation, even though full speciation has not been achieved in this case. It also explains the paucity of anole species in the Lesser Antilles compared to the Greater Antilles.     

PROVINCIALISM IN PLANKTON: ENDEMISM AND ALLOPATRIC SPECIATION IN AUSTRALIAN DAPHNIA

Zooplankton species are generally considered poor candidates for allopatric speciation because of their broad distributions and capabilities for long-distance dispersal. We examined the validity of this conclusion by determining both species distributions and the extent of gene frequency divergence in the Daphnia fauna of southeastern Australia, a mature landscape dominated by members of the carinata complex. Although delimitation of species boundaries was complicated by the prevalence of interspecific hybrids and variation in breeding systems, allozyme analysis of 187 populations indicated the presence of at least seven species. All of these species had restricted distributions, and several were narrowly endemic. Gene frequency divergence was often apparent between populations separated by only a few kilometers but was least prominent in species from inland areas. The extent of regional gene frequency shifts varied among species—two narrowly distributed (projecta, thomsoni) and one broadly distributed (carinata) species showed little divergence between sites, but two other common species (cephalata, longicephala) showed marked gene frequency shifts coincident with physiographic barriers. Together, the limited species distributions and regional gene-pool fragmentation suggest that allopatric speciation has played an important role in the origin of taxon diversity in the Daphnia carinata complex.     

HISTORICAL DIVERGENCE AND GENE FLOW: COALESCENT ANALYSES OF MITOCHONDRIAL,AUTOSOMAL AND SEX‐LINKED LOCI IN PASSERINA BUNTINGS

Quantifying the role of gene flow during the divergence of closely related species is crucial to understanding the process of speciation. We collected DNA sequence data from 20 loci (one mitochondrial, 13 autosomal, and six sex‐linked) for population samples of Lazuli Buntings (Passerina amoena) and Indigo Buntings (Passerina cyanea) (Aves: Cardinalidae) to test explicitly between a strict allopatric speciation model and a model in which divergence occurred despite postdivergence gene flow. Likelihood ratio tests of coalescent‐based population genetic parameter estimates indicated a strong signal of postdivergence gene flow and a strict allopatric speciation model was rejected. Analyses of partitioned datasets (mitochondrial, autosomal, and sex‐linked) suggest the overall gene flow patterns are driven primarily by autosomal gene flow, as there is no evidence of mitochondrial gene flow and we were unable to reject an allopatric speciation model for the sex‐linked data. This pattern is consistent with either a parapatric divergence model or repeated periods of allopatry with gene flow occurring via secondary contact. These results are consistent with the low fitness of female avian hybrids under Haldane's rule and demonstrate that sex‐linked loci likely are important in the initial generation of reproductive isolation, not just its maintenance.     

The relative importance of geography and ecology in species diversification: evidence from a tropical marine intertidal snail (Nerita)

Aim In this study, I examined the relative contributions of geography and ecology to species diversification within the genus Nerita, a prominent clade of marine snails that is widely distributed across the tropics and intertidal habitats. Specifically, I tested whether geographical patterns of speciation correspond primarily to allopatric or sympatric models, and whether habitat transitions have played a major role in species diversification. Location Indo‐West Pacific, eastern Pacific, Atlantic, tropical marine intertidal. Methods I used a previously reconstructed molecular phylogeny of Nerita as a framework to assess the relative importance of geographical and ecological factors in species diversification. To evaluate whether recently diverged clades exhibit patterns consistent with allopatric or sympatric speciation, I mapped the geo‐graphical distribution of each species onto the species‐level phylogeny, and examined the relationship between range overlap and time since divergence using age–range correlation analyses. To determine the relative contribution of habitat transitions to divergence, I traced shifts in intertidal substrate affinity and vertical zonation across the phylogeny using parsimony, and implemented randomization tests to evaluate the resulting patterns of ecological change. Results Within the majority of Nerita clades examined, age–range correlation analysis yielded a low intercept and a positive slope, similar to that expected under allopatric speciation. Approximately 75% of sister species pairs have maintained allopatric distributions; whereas more distantly related sister taxa often exhibited complete or nearly complete geographical overlap. In contrast, only 19% of sister species occupy distinct habitats. For both substrate and zonation, habitat transitions failed to concentrate towards either the tips or the root of the phylogeny. Instead, habitat shifts have occurred throughout the history of Nerita, with a general transition from the lower and mid‐littoral towards the upper and supra‐littoral zones, and multiple independent shifts from hard (rock) to softer substrates (mangrove, mud and sand). Main conclusions Both geography and ecology appear to have influenced diversification in Nerita, but to different extents. Geography seems to play a principal role, with allopatric speciation driving the majority of Nerita divergences. Habitat transitions appear insignificant in shaping the early and recent history of speciation, and promoting successive diversification in Nerita; however, shifts may have been important for respective divergences (i.e. those that correspond to the transitions) and enhancing diversity throughout the clade.     

THE RATE TEST OF SPECIATION: ESTIMATING THE LIKELIHOOD OF NON‐ALLOPATRIC SPECIATION FROM REPRODUCTIVE ISOLATION RATES IN DROSOPHILA

Among the most debated subjects in speciation is the question of its mode. Although allopatric (geographical) speciation is assumed the null model, the importance of parapatric and sympatric speciation is extremely difficult to assess and remains controversial. Here I develop a novel approach to distinguish these modes of speciation by studying the evolution of reproductive isolation (RI) among taxa. I focus on the Drosophila genus, for which measures of RI are known. First, I incorporate RI into age‐range correlations. Plots show that almost all cases of weak RI are between allopatric taxa whereas sympatric taxa have strong RI. This either implies that most reproductive isolation (RI) was initiated in allopatry or that RI evolves too rapidly in sympatry to be captured at incipient stages. To distinguish between these explanations, I develop a new “rate test of speciation” that estimates the likelihood of non‐allopatric speciation given the distribution of RI rates in allopatry versus sympatry. Most sympatric taxa were found to have likely initiated RI in allopatry. However, two putative candidate species pairs for non‐allopatric speciation were identified (5% of known Drosophila). In total, this study shows how using RI measures can greatly inform us about the geographical mode of speciation in nature.     

Biogeography and systematics of endemic island damselflies: The Nesobasis and Melanesobasis (Odonata: Zygoptera) of Fiji

The study of island fauna has greatly informed our understanding of the evolution of diversity. We here examine the phylogenetics, biogeography, and diversification of the damselfly genera Nesobasis and Melanesobasis, endemic to the Fiji Islands, to explore mechanisms of speciation in these highly speciose groups. Using mitochondrial (COI, 12S) and nuclear (ITS) replicons, we recovered Garli ‐part maximum likelihood and Mrbayes Bayesian phylogenetic hypotheses for 26 species of Nesobasis and eight species/subspecies of Melanesobasis. Biogeographical patterns were explored using Lagrange and Bayes ‐Lagrange and interpreted through beast relaxed clock dating analyses. We found that Nesobasis and Melanesobasis have radiated throughout Fiji, but are not sister groups. For Nesobasis, while the two largest islands of the archipelago—Viti Levu and Vanua Levu—currently host two distinct species assemblages, they do not represent phylogenetic clades; of the three major groupings each contains some Viti Levu and some Vanua Levu species, suggesting independent colonization events across the archipelago. Our Beast analysis suggests a high level of species diversification around 2–6 Ma. Our ancestral area reconstruction (Rasp ‐Lagrange ) suggests that both dispersal and vicariance events contributed to the evolution of diversity. We thus conclude that the evolutionary history of Nesobasis and Melanesobasis is complex; while inter‐island dispersal followed by speciation (i.e., peripatry) has contributed to diversity, speciation within islands appears to have taken place a number of times as well. This speciation has taken place relatively recently and appears to be driven more by reproductive isolation than by ecological differentiation: while species in Nesobasis are morphologically distinct from one another, they are ecologically very similar, and currently are found to exist sympatrically throughout the islands on which they are distributed. We consider the potential for allopatric speciation within islands, as well as the influence of parasitic endosymbionts, to explain the high rates of speciation in these damselflies.     

Strong reproductive barriers in a narrow hybrid zone of West-Mediterranean green toads (<Emphasis Type="Italic">Bufo viridis</Emphasis>subgroup) with Plio-Pleistocene divergence

Background  

One key question in evolutionary biology deals with the mode and rate at which reproductive isolation accumulates during allopatric speciation. Little is known about secondary contacts of recently diverged anuran species. Here we conduct a multi-locus field study to investigate a contact zone between two lineages of green toads with an estimated divergence time of 2.7 My, and report results from preliminary experimental crosses.     

ADAPTIVE RADIATION OF DAY-GECKOS (PHELSUMA) IN THE SEYCHELLES ARCHIPELAGO: A PHYLOGENETIC ANALYSIS

A phylogenetic analysis using characters derived from mitochondrial DNA was used to show that the species of Phelsuma in the Seychelles Islands represent a single, monophyletic lineage that has diversified as a result of both historical and ecological factors. In the distant past, the Seychelles archipelago was physically invaded by a single species of Phesluma. Separate eustatic sea level changes likely led first to allopatric speciation and then to the secondary contact of these sister species. Differences in the relative timing of the secondary contact between island groups resulted in P. sundbergi evolving an intermediate body size in the group of islands associated with Mahé and a large body size, while sympatric with P. astriata, in the group of islands associated with Praslin. Ecological information was used to support the conclusion that the actual evolutionary mechanism for the body size shift was a response to frequency dependent natural selection of P. sundbergi in single-species and two-species competitive regimes.     

A geographical model of high species diversity

Several cases of high species diversity, for example in tropical rain forests, imply that speciation has been frequent or rapid. However, how speciation could proceed so frequently as to generate extraordinary diversity still remains unsolved, despite recent advancements of diverse theories of allopatric and sympatric speciation. This paper presents a theoretical model that demonstrates the process of frequent speciation by means of geographical fragmentation. We focus on allopatric speciation and explore the evolutionary effect of fragmentation and extinction of demes (subpopulations) in a widespread species or species group. After a large contagious population of a single species is fragmented into demes, extinction of some demes could result in isolation of multiple demes. Thus, several demes could become good species simultaneously through the process of allopatric speciation. We apply the random extinction method to this fragmentation process where demes become randomly extinct. The present model illustrates that frequent speciation could occur in communities where large environmental changes frequently take place.     

Geographic patterns of postzygotic isolation between two closely related widespread dung fly species (Sepsis cynipsea and Sepsis neocynipsea; Diptera: Sepsidae)

Identifying the contribution of pre‐ and postzygotic barriers to gene flow is a key goal of speciation research. The widespread dung fly species Sepsis cynipsea and Sepsis neocynipsea offer great potential for studying the speciation process over a range of opportunities for gene exchange within and across sister species (cross‐continental allopatry, continental parapatry and sympatry). We examined the role of postcopulatory isolating barriers by comparing female fecundity and egg‐to‐adult viability of F₁ and F₂ hybrids, as well as backcrosses of F₁ hybrids with the parental species, via replicated crosses of sym‐, para‐ and allopatric populations. Egg‐to‐adult viability was strongly but not totally suppressed in hybrids, and offspring production approached nil in the F₂ generation (hybrid breakdown), indicating yet unspecified intrinsic incompatibilities. Viable F₁ hybrid offspring showed almost absolute male (the heterogametic sex) sterility while females remained largely fertile, in accordance with Haldane's rule. Hybridization between the two species in European areas of sympatry (Swiss Alps) indicated only minor reinforcement based on fecundity traits. Crossing geographically isolated European and North American S. neocynipsea showed similar albeit weaker isolating barriers that are most easily explained by random genetic drift. We conclude that in this system with a biogeographic continuum of reproductive barriers, speciation is mediated primarily by genetic drift following dispersal of flies over a wide (allopatric) geographic range, with some role of natural or sexual selection in incidental or direct reinforcement of incompatibility mechanisms in areas of European sympatry. S(ubs)pecies status of continental S. neocynipsea appears warranted.