Ecological separation in a polymorphic terrestrial salamander
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1 When studying speciation, researchers commonly examine reproductive isolation in recently diverged populations. Polymorphic species provide an opportunity to examine the role of reproductive isolation in populations that may be in the process of divergence. 2 We examined a polymorphic population of Plethodon cinereus (red-backed salamanders) for evidence of sympatric ecological separation by colour morphology. Recent studies have correlated temperature and climate with colour morphology in this species, but no studies have looked at differences in diet or mate choice between colour morphs. We used artificial cover objects to assess salamander diet, mating preference and surface activity over a 2-year period at a field site in north-eastern Ohio. 3 We detected differences in diet between two colour morphs, striped and unstriped. The diets of striped individuals were significantly more diverse and were made up of more profitable prey than the diets of unstriped salamanders. 4 Opposite sex pairs were made up of individuals of the same colour morph and striped males were found more often with larger females than were unstriped males. 5 We corroborate findings of earlier studies suggesting that the unstriped form is adapted to warmer conditions. Unstriped individuals were the first to withdraw from the forest floor as temperatures fell in the late fall. We found no evidence that the colour morphs responded differently to abiotic factors such as soil moisture and relative humidity, and responses to surface temperatures were also equivocal. 6 We conclude that the two colour morphs exhibit some degree of ecological separation and tend to mate assortatively, but are unlikely to be undergoing divergence given the observed frequency of intermorph pairings.Keywords:
Reproductive isolation
Ecological speciation
Ecological speciation
Adaptive Radiation
Genetic algorithm
Reproductive isolation
Parapatric speciation
Character displacement
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HABITAT AVOIDANCE: OVERLOOKING AN IMPORTANT ASPECT OF HOST-SPECIFIC MATING AND SYMPATRIC SPECIATION?
Understanding speciation requires discerning how reproductive barriers to gene flow evolve between previously interbreeding populations. Models of sympatric speciation for phytophagous insects posit that reproductive isolation can evolve in the absence of geographic isolation as a consequence of an insect shifting and ecologically adapting to a new host plant. One important adaptation contributing to sympatric differentiation is host-specific mating. When organisms mate in preferred habitats, a system of positive assortative mating is established that facilitates sympatric divergence. Models of host fidelity generally assume that host choice is determined by the aggregate effect of alleles imparting positive preferences for different plant species. But negative effect genes for avoiding nonnatal plants may also influence host use. Previous studies have shown that apple and hawthorn-infesting races of Rhagoletis pomonella flies use volatile compounds emitted from the surface of fruit as key chemosensory cues to recognize and distinguish between their host plants. Here, we report results from field trials indicating that in addition to preferring the odor of their natal fruit, apple and hawthorn flies, and their undescribed sister species infesting flowering dogwood (Cornus florida), also avoid the odors of nonnatal fruit. We discuss the implications of nonnatal fruit avoidance for the evolutionary dynamics and genetics of sympatric speciation. Our findings reveal an underappreciated role for habitat avoidance as a potential postmating, as well as prezygotic, barrier to gene flow.
Reproductive isolation
Ecological speciation
Assortative mating
Genetic algorithm
Mating preferences
Incipient speciation
Disruptive selection
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Amongst several theories of speciation, sympatric speciation has been the most controversial but it is now widely accepted that populations can become reproductively isolated without being separated geographically. One problem with the acceptance of the theory of sympatric speciation, however, has been the lack of supporting empirical data and it is still believed that geographical isolation is responsible for the majority of speciation events. Here the example of species pairs in lampreys suggests that sympatric speciation in a whole taxonomic group could occur throughout its worldwide range. Lampreys occur in two ecologically distinct forms: parasitic mostly anadromous species that forage on tissue and body fluids of host fishes, and non‐parasitic forms that, apart from a short adult life when they cease feeding, spend their entire life as filter feeders in the substratum of stream beds. Both forms occur in sympatric species pairs throughout the range of lampreys that occur in Eurasia, North America and Australia and it is widely acknowledged that non‐parasitic forms derive from parasitic forms. The larvae of both forms can be distinguished by their potential fecundity and therefore, it is argued that the mode of life is not a consequence of different ecological conditions. Furthermore, as lampreys prefer to choose mates of similar sizes and fertilization success decreases with increasing difference in body size, there is a strong disruptive selection between the two forms and they are therefore reproductively isolated. Besides theoretical aspects, the similarity of the species pairs, together with their occurrence in sympatry, the occurrence of forms with intermediate characteristics, and examples where speciation might be in progress, hints at the possibility that speciation also occurred in sympatry. The difference between lampreys and other examples of sympatric speciation is that there seems to be a trend towards sympatric speciation events throughout the worldwide range of lampreys which is neither restricted to relatively small localities nor caused by human disturbance. Species pairs in lampreys therefore offer a unique possibility of studying the process of sympatric speciation on a large scale.
Ecological speciation
Reproductive isolation
Genetic algorithm
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Abstract The process of speciation generates biodiversity. According to the null model of speciation, barriers between populations arise in allopatry, where, prior to biology, geography imposes barriers to gene flow. On the other hand, sympatric speciation requires that the process of speciation happen in the absence of a geographical barrier, where the members of the population have no spatial, temporal barriers. Several attempts have been made to theoretically identify the conditions in which speciation can occur in sympatry. However, these efforts suffer from several limitations. We propose a model for sympatric speciation based on adaptation for resource utilization. We use a genetics-based model to investigate the relative roles of prezygotic and postzygotic barriers, from the context of ecological disruptive selection, sexual selection, and genetic architecture, in causing and maintaining sympatric speciation. Our results show that sexual selection that acts on secondary sexual traits does not play any role in the process of speciation in sympatry and that assortative mating based on an ecologically relevant trait forces the population to show an adaptive response. We also demonstrate that understanding the genetic architecture of the trait under ecological selection is very important and that it is not required for the strength of ecological disruptive selection to be very high in order for speciation to occur in sympatry. Our results provide an insight into the kind of scenarios in which sympatric speciation can be demonstrated in the lab.
Disruptive selection
Ecological speciation
Genetic algorithm
Incipient speciation
Assortative mating
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Species delineation has a long and contentious history, yet most agree that sympatric populations exhibiting high levels of reproductive isolation and evolving independently are species. In an opinion piece, Hill and Powers (2021; hereafter H&P) claim that several recognized species of crossbills ( Loxia spp.) do not represent species because by no measure are they discrete, the vocalizations used to categorize crossbills are learned, modified and can switch to that of a different species, and reproductive isolation is incomplete and weak. We argue that the behavioral and genetic evidence indicate that Cassia crossbills L. sinesciuris , which we focus on because the data relevant to species status are more diverse and extensive, are genetically discrete; call modification rarely leads to crossbill misclassification and overwhelmingly results in call divergence and enhanced discrimination; and are nearly completely reproductively isolated with little evidence of introgression from sympatric red crossbills. The differences in our conclusions result in part from H&P mischaracterizing and misconstruing the ecology of Cassia crossbills, geographic context of their divergence, and evidence for reproductive isolation. H&P seemingly require that crossbills must adhere to the typical model of bird speciation–protracted divergence in allopatry, followed by a gradual increase in sympatry if reproductive isolation and ecological divergence allow–and require evidence such as initial long periods of allopatry, F ST values > 0.2, divergent mtDNA and intrinsic postzygotic isolation. Although such evidence commonly distinguishes bird species, an increasing number of studies show that such criteria are not necessary to indicate sympatric, evolutionarily independent lineages.
Reproductive isolation
Parapatric speciation
Divergence (linguistics)
Introgression
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HABITAT AVOIDANCE: OVERLOOKING AN IMPORTANT ASPECT OF HOST-SPECIFIC MATING AND SYMPATRIC SPECIATION?
Understanding speciation requires discerning how reproductive barriers to gene flow evolve between previously interbreeding populations. Models of sympatric speciation for phytophagous insects posit that reproductive isolation can evolve in the absence of geographic isolation as a consequence of an insect shifting and ecologically adapting to a new host plant. One important adaptation contributing to sympatric differentiation is host‐specific mating. When organisms mate in preferred habitats, a system of positive assortative mating is established that facilitates sympatric divergence. Models of host fidelity generally assume that host choice is determined by the aggregate effect of alleles imparting positive preferences for different plant species. But negative effect genes for avoiding nonnatal plants may also influence host use. Previous studies have shown that apple and hawthorn‐infesting races of Rhagoletis pomonella flies use volatile compounds emitted from the surface of fruit as key chemosensory cues to recognize and distinguish between their host plants. Here, we report results from field trials indicating that in addition to preferring the odor of their natal fruit, apple and hawthorn flies, and their undescribed sister species infesting flowering dogwood (Cornus florida), also avoid the odors of nonnatal fruit. We discuss the implications of nonnatal fruit avoidance for the evolutionary dynamics and genetics of sympatric speciation. Our findings reveal an under appreciated role for habitat avoidance as a potential postmating, as well as prezygotic, barrier to gene flow.
Reproductive isolation
Ecological speciation
Assortative mating
Mating preferences
Genetic algorithm
Incipient speciation
Disruptive selection
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Citations (72)
Ecological speciation
Genetic algorithm
Reproductive isolation
Incipient speciation
Disruptive selection
Genetic divergence
Divergence (linguistics)
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Reproductive isolation
Ecological speciation
Genetic algorithm
Etheostoma
Incipient speciation
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Abstract The process of speciation is the source of biodiversity. The most popularly accepted mode of speciation is allopatric speciation, where geography imposes the initial barrier to gene flow, and then biological barriers come up. On the other hand, sympatric speciation, which was not accepted as a possibility for long, requires that the process of speciation happen in the absence of a geographical barrier, in a well-mixed population. Several attempts have been made to theoretically identify the conditions in which speciation can occur in sympatry, but have several problems associated with them. We propose a model for sympatric speciation based on adaptation for resource utilization. We use this genetics- based model to investigate the relative roles of prezygotic and postzygotic barriers, from the context of ecological disruptive selection, sexual selection, and genetic architecture, in causing and maintaining sympatric speciation. We show that sexual selection that acts on secondary sexual traits does not play any role in the process of speciation in sympatry, and that assortative mating based on an ecologically relevant trait forces the population to show an adaptive response. We also demonstrate that understanding the genetic architecture of the trait under ecological selection is very important, and that it is not required for the strength of ecological disruptive selection to be very high in order for speciation to occur in sympatry. With this, we provide an insight into the kind of scenarios in which sympatric speciation can be demonstrated in lab.
Disruptive selection
Ecological speciation
Genetic algorithm
Incipient speciation
Assortative mating
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Knowledge on interspecific pre- and post-zygotic isolation mechanisms provides insights into speciation patterns. Using crosses (F(1) and backcrosses) of two closely related flea beetles species, Altica fragariae and A. viridicyanea, specialized on different hosts in sympatry, we measured: (a) the type of reproductive isolation and (b) the inheritance mode of preference and host-specific performance, using a joint-scaling test. Each species preferred almost exclusively its host plant, creating strong prezygotic isolation between them, and suggesting that speciation may occur at least partly in sympatry. Reproductive isolation was intrinsic between females of A. fragariae and either A. viridicyanea or F(1) males, whereas the other crosses showed ecologically dependent reproductive isolation, suggesting ecological speciation. The genetic basis of preference and performance was at least partially independent, and several loci coded for preference, which limits the possibility of sympatric speciation. Hence, both ecological and intrinsic factors may contribute to speciation between these species.
Reproductive isolation
Ecological speciation
Genetic algorithm
Incipient speciation
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Citations (26)