Sympatric speciation by allochrony?

Sympatric speciation was once thought most improbable, but careful study of some systems, particularly the apple maggot (Rhagoletis pomonella) and related Rhagoletis species, has led to its reinstatement as a likely mode of speciation in some cases. Different species and host races in this clade of flies often have highly specialized host preference, and along with frequent evolutionary shifts to different fruit species between sister taxa, there is a likely effect of the timing of adult emergence that follows host fruiting phenology. This is known as “allochronic” isolation (from the Greek meaning “different timing”). This overview covers recent discoveries by Inskeep et al. (Molecular Ecology, 2021) showing how allochrony is a major factor in preventing gene flow between a pair of sister species of Rhagoletis on different host fruits. Although the authors do not claim to prove sympatric speciation, it does seem very likely, and the work clearly underscores how readily host shifts via allochrony can aid sympatric speciation

Extensive range overlap between heliconiine sister species: evidence for sympatric speciation in butterflies?

BACKGROUND: Sympatric speciation is today generally viewed as plausible, and some well-supported examples exist, but its relative contribution to biodiversity remains to be established. We here quantify geographic overlap of sister species of heliconiine butterflies, and use age-range correlations and spatial simulations of the geography of speciation to infer the frequency of sympatric speciation. We also test whether shifts in mimetic wing colour pattern, host plant use and climate niche play a role in speciation, and whether such shifts are associated with sympatry. RESULTS: Approximately a third of all heliconiine sister species pairs exhibit near complete range overlap, and analyses of the observed patterns of range overlap suggest that sympatric speciation contributes 32%-95% of speciation events. Müllerian mimicry colour patterns and host plant choice are highly labile traits that seem to be associated with speciation, but we find no association between shifts in these traits and range overlap. In contrast, climatic niches of sister species are more conserved. CONCLUSIONS: Unlike birds and mammals, sister species of heliconiines are often sympatric and our inferences using the most recent comparative methods suggest that sympatric speciation is common. However, if sister species spread rapidly into sympatry (e.g. due to their similar climatic niches), then assumptions underlying our methods would be violated. Furthermore, although we find some evidence for the role of ecology in speciation, ecological shifts did not show the associations with range overlap expected under sympatric speciation. We delimit species of heliconiines in three different ways, based on "strict and " "relaxed" biological species concepts (BSC), as well as on a surrogate for the widely-used "diagnostic" version of the phylogenetic species concept (PSC). We show that one reason why more sympatric speciation is inferred in heliconiines than in birds may be due to a different culture of species delimitation in the two groups. To establish whether heliconiines are exceptional will require biogeographic comparative studies for a wider range of animal taxa including many more invertebrates

The Amazon river is a suture zone for a polyphyletic group of co-mimetic heliconiine butterflies

The Amazon basin contains few obvious geographic barriers, yet it is the most biodiverse region on Earth. One hypothesis to explain its diversity is that the very large rivers promote allopatric divergence. Consistent with this, maps of heliconiine butterflies made from museum specimens show high subspecies richness close to the Amazon river, suggesting that it may produce or maintain intra-specific phenotypic variability. However, museum data are subject to strong spatial biases (the ‘Wallacean shortfall' of distribution data), raising the possibility that this pattern is a sampling artefact. To test this, we systematically collected along a ~900 km north–south transect running through central Amazonia. We found a significant association between phenotypic diversity and major rivers, with distance from the Amazon river explaining 61% of the variance in the mean polymorphism of 25 species. This association is partly because many species exhibit different phenotypes on either side of the river. Nonetheless, we also find sites with high polymorphism close to the river, indicating continual cross-river dispersal. Our results strongly suggest the presence of a suture zone (a region where multiple species have hybrid zones) near the city of Manaus. However, the effect of the river on spatial patterns of intra-specific phenotypic diversity depends on a species' mimetic phenotype. Rather than being absolute barriers, our results support the idea that rivers can act as partial barriers that trap moving hybrid zones, resulting in a suture zone. As such, the wide Amazonian rivers help generate and maintain colour pattern diversity, but to date there is no evidence that they lead to speciation in our study group

Clypeal patterning in the paper wasp Polistes dominulus: no evidence of adaptive value in the wild

Status signals function in a number of species to communicate competitive ability to conspecific rivals during competition for resources. In the paper wasp Polistes dominulus, variable black clypeal patterns are thought to be important in mediating competition among females. Results of previous behavioral experiments in the lab indicate that P. dominulus clypeal patterns provide information about an individual's competitive ability to rivals during agonistic interactions. To date, however, there has been no detailed examination of the adaptive value of clypeal patterns in the wild. To address this, we looked for correlations between clypeal patterning and various fitness measures, including reproductive success, hierarchical rank, and survival, in a large, free-living population of P. dominulus in southern Spain. Reproductive success over the nesting season was not correlated with clypeal patterning. Furthermore, there was no relationship between a female's clypeal patterning and the rank she achieved within the hierarchy or her survival during nest founding. Overall, we found no evidence that P. dominulus clypeal patterns are related to competitive ability or other aspects of quality in our population. This result is consistent with geographical variation in the adaptive value of clypeal patterns between P. dominulus populations; however, data on the relationship between patterning and fitness from other populations are required to test this hypothesi

Clypeal patterning in the paper wasp Polistes dominulus: no evidence of adaptive value in the wild

Status signals function in a number of species to communicate competitive ability to conspecific rivals during competition for resources. In the paper wasp Polistes dominulus, variable black clypeal patterns are thought to be important in mediating competition among females. Results of previous behavioral experiments in the lab indicate that P. dominulus clypeal patterns provide information about an individual's competitive ability to rivals during agonistic interactions. To date, however, there has been no detailed examination of the adaptive value of clypeal patterns in the wild. To address this, we looked for correlations between clypeal patterning and various fitness measures, including reproductive success, hierarchical rank, and survival, in a large, free-living population of P. dominulus in southern Spain. Reproductive success over the nesting season was not correlated with clypeal patterning. Furthermore, there was no relationship between a female's clypeal patterning and the rank she achieved within the hierarchy or her survival during nest founding. Overall, we found no evidence that P. dominulus clypeal patterns are related to competitive ability or other aspects of quality in our population. This result is consistent with geographical variation in the adaptive value of clypeal patterns between P. dominulus populations; however, data on the relationship between patterning and fitness from other populations are required to test this hypothesi

Geographic contrasts between pre- and postzygotic barriers are consistent with reinforcement in Heliconius butterflies.

Identifying the traits causing reproductive isolation and the order in which they evolve isfundamental to understanding speciation. Here, we quantify prezygotic and intrinsicpostzygotic isolation between allopatric, parapatric and sympatric populations of thebutterflies Heliconius elevatus and Heliconius pardalinus. Sympatric populations from theAmazon (H. elevatus and H. p. butleri) exhibit strong prezygotic isolation and rarely mate incaptivity; however, hybrids are fertile. Allopatric populations from the Amazon(H. p. butleri) and Andes (H. p. sergestus) mate freely when brought together in captivity, butthe female F1 hybrids are sterile. Parapatric populations (H. elevatus and H. p. sergestus)exhibit both assortative mating and sterility of female F1s. Assortative mating in sympatricpopulations is consistent with reinforcement in the face of gene flow, where the driving force,selection against hybrids, is due to disruption of mimicry and other ecological traits ratherthan hybrid sterility. In contrast, the lack of assortative mating and hybrid sterility observedin allopatric populations suggests that geographic isolation enables the evolution of intrinsicpostzygotic reproductive isolation. Our results show how the types of reproductive barriersthat evolve between species may depend on geography

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Cryptic speciation associated with geographic and ecological divergence in two Amazonian Heliconius butterflies

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