Natural and sexual selection against immigrants maintains differentiation among micro‐allopatric populations

Local adaptation to divergent environmental conditions can promote population genetic differentiation even in the absence of geographic barriers and hence lead to speciation. But what mechanisms contribute to reproductive isolation among diverging populations? We tested for natural and sexual selection against immigrants in a fish species inhabiting (and adapting to) nonsulphidic surface habitats, sulphidic surface habitats and a sulphidic cave. Gene flow is strong among sample sites situated within the same habitat type, but low among divergent habitat types. Our results indicate that females of both sulphidic populations discriminate against immigrant males during mate choice. Furthermore, using reciprocal translocation experiments, we document natural selection against migrants between nonsulphidic and sulphidic habitats, whereas migrants between sulphidic cave and surface habitats did not exhibit increased mortality within the same time period. Consequently, both natural and sexual selection may contribute to isolation among parapatric populations, and selection against immigrants may be a powerful mechanism facilitating speciation among locally adapted populations even over very small spatial distances.     

Divergence in trophic ecology characterizes colonization of extreme habitats

Extreme habitats are characterized by the presence of physio‐chemical stressors, but also differ in aspects of the biotic environment, such as resource availability or the presence of competitors. The present study quantifies variation in trophic ecology of a small livebearing fish (Poecilia mexicana, Poeciliidae) across four different habitats that included nonsulphidic and sulphidic surface waters, as well as a nonsulphidic and a sulphidic cave. Resource use in different habitat types was investigated using gut content analysis. Populations diverged in resource use from a diet dominated by algae and detritus in nonsulfidic surface habitats to a diet including invertebrate food items in the other habitats. Poecilia mexicana in cave habitats further exhibited a higher dietary niche width than conspecifics from surface habitats. The condition of P. mexicana was analysed using storage lipid extractions. Fish from sulphidic and cave habitats exhibited a very poor condition, suggesting resource limitation and/or high costs of coping with extreme conditions. Finally, divergence in resource use was correlated with variation in viscerocranial morphology. A common garden experiment indicated both a genetic and plastic basis to the morphological variation observed among field populations. It is suggested that the morphological diversification is an adaptation to the differential use of resources among populations. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 517–528.     

Convergent changes in the trophic ecology of extremophile fish along replicated environmental gradients

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Risk of predation as a promoting factor of species divergence in threespine sticklebacks (Gasterosteus aculeatus L.)

Icelandic freshwater systems are geologically young and contain only six species of freshwater fish. As these species colonized Icelandic fresh waters they were presented with a diversity of unique, uncontested habitats and food resources, promoting the evolution of new behaviour strategies crucial to the formation of new morphs and speciation. To determine the likelihood that predation threat could affect the antipredator behaviour and possibly the sympatric divergence of prey populations, we analysed antipredator behaviour of seven groups of Icelandic threespine sticklebacks ( Gasterosteus aculeatus ): two marine groups, one group from a lake without piscine predators, and two polymorphic lake populations, each with two groups occupying unique habitats. Shoaling cohesion, school formation and duration, and vigilance in predator inspection/avoidance behaviour varied greatly among groups. The differences appeared to be related to the risk of predation as well as to opportunities and constraints set by the different habitats. Antipredator behaviour was especially pronounced and differed extensively in two polymorphic forms from the lake Thingvallavatn, where predation risk is very high. By keeping the two morphs separate in their respective habitats, high predation risk may be a contributing factor in promoting the habitat-specific divergence of G. aculeatus seen in the lake. This suggests that in situations where refuge habitats are spatially separated, the risk of predation may contribute to the evolution of separate sympatric forms of small fish such as G. aculeatus .  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 82 , 189–203.     

Two endemic and endangered fishes, Poecilia sulphuraria (Alvarez, 1948) and Gambusia eurystoma Miller, 1975 (Poeciliidae, Teleostei) as only survivors in a small sulphidic habitat

In the Baños del Azufre in Tabasco, Mexico, only two poeciliid species, Poecilia sulphuraria and Gambusia eurystoma , were found in sulphidic habitats, whereas adjacent non-sulphidic habitats were characterized by a vastly different fish community. Exposure of fishes to water from a sulphidic spring showed that all species except for P. sulphuraria quickly lost motion control, probably because of the toxic properties of H₂S. It is suggested that the fishes endemic to the Baños del Azufre are highly specialized, and competitive exclusion allows them to survive only in this particular habitat type.     

Evolution in extreme environments: replicated phenotypic differentiation in livebearing fish inhabiting sulfidic springs

We investigated replicated ecological speciation in the livebearing fish Poecilia mexicana and P. sulphuraria (Poeciliidae), which inhabit freshwater habitats and have also colonized multiple sulfidic springs in southern Mexico. These springs exhibit extreme hypoxia and high concentrations of hydrogen sulfide, which is lethal to most metazoans. We used phylogenetic analyses to test whether springs were independently colonized, performed phenotypic assessments of body and gill morphology variation to identify convergent patterns of trait differentiation, and conducted an eco-toxicological experiment to detect differences in sulfide tolerances among ecotypes. Our results indicate that sulfidic springs were colonized by three different lineages, two within P. mexicana and one representing P. sulphuraria. Colonization occurred earlier in P. sulphuraria, whereas invasion of sulfidic springs in P. mexicana was more recent, such that each population is more closely related to neighboring populations from adjacent nonsulfidic habitats. Sulfide spring fish also show divergence from nonsulfidic phenotypes and a phenotypic convergence toward larger heads, larger gills, and increased tolerance to H(2) S. Together with previous studies that indicated significant reproductive isolation between fish from sulfidic and nonsulfidic habitats, this study provides evidence for repeated ecological speciation in the independent sulfide spring populations of P. mexicana and P. sulphuraria.     

Female mating preferences and male coloration covary with water transparency in a Lake Victoria cichlid fish

Rapid speciation in Lake Victoria cichlid fish of the genus Pundamilia may be facilitated by sexual selection: female mate choice exerts sexual selection on male nuptial coloration within species and maintains reproductive isolation between species. However, declining water transparency coincides with increasingly dull coloration and increasing hybridization. In the present study, we investigated the mechanism underlying this pattern in Pundamilia nyererei, a species that interbreeds with a sister species in turbid but not in clear water. We compared measures of intraspecific sexual selection between two populations from locations that differ in water transparency. First, in laboratory mate‐choice experiments, conducted in clear water and under broad‐spectrum illumination, we found that females originating from turbid water have significantly weaker preferences for male coloration than females originating from clear water. Second, both the hue and body coverage of male coloration differ between populations, which is consistent with adaptation to different photic habitats. These findings suggest that the observed relationship between male coloration and water transparency is not mediated by environmental variation alone. Rather, female mating preferences are indicated to have changed in response to this variation, constituting the first evidence for intraspecific preference‐trait co‐evolution in cichlid fish. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 398–406.     

Fine-scale genetic structure, estuarine colonization and incipient speciation in the marine silverside fish Odontesthes argentinensis

The identification of incipient ecological species represents an opportunity to investigate current evolutionary process where adaptive divergence and reproductive isolation are associated. In this study we analysed the genetic structure of marine and estuarine populations of the silverside fish Odontesthes argentinensis using nine microsatellite loci and 396 bp of the mitochondrial DNA (mtDNA) control region. Our main objective was to investigate the relationship among estuarine colonization, divergent selection and speciation in silversides. Significant genetic structure was detected among all marine and estuarine populations. Despite the low phylogeographic structure in mtDNA haplotypes, there was clear signal of local radiations of haplotypes in more ancient populations. Divergence among marine populations was interpreted as a combined result of homing behaviour, isolation by distance and drift. On the other hand, ecological shifts due to the colonization of estuarine habitats seem to have promoted rapid adaptive divergence and reproductive isolation in estuarine populations, which were considered as incipient ecological species. This conclusion is supported by the existence of a set of environmental factors required for successful reproduction of estuarine ecotypes. The pattern of genetic structure indicates that phenotypic and reproductive divergence evolved in the face of potential gene flow between populations. We suggest that the 'divergence-with-gene-flow' model of speciation may account for the diversification of estuarine populations. The approach used can potentially identify 'incipient estuarine species', being relevant to the investigation of the evolutionary relationships of silversides in several coastal regions of the world.     

Studying phenotypic plasticity: the advantages of a broad approach

Human evolution has clearly been shaped by gene–culture interactions, and there is growing evidence that similar processes also act on populations of non‐human animals. Recent theoretical studies have shown that culture can be an important evolutionary mechanism because of the ability of cultural traits to spread rapidly both vertically, obliquely, and horizontally, resulting in decreased within‐group variance and increased between‐group variance. Here, we collate the extensive literature on population divergence in killer whales (Orcinus orca), and argue that they are undergoing ecological speciation as a result of dietary specializations. Although we cannot exclude the possibility that cultural divergence pre‐dates ecological divergence, we propose that cultural differences in the form of learned behaviours between ecologically divergent killer whale populations have resulted in sufficient reproductive isolation even in sympatry to lead to incipient speciation. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 1–17.     

Phenotype and gene flow in a marsupial (Antechinus flavipes) in contrasting habitats

Ecological factors are important drivers of phenotypic divergence, which may lead to incipient speciation. A variety of habitats should be preserved to maintain evolutionary potential. We used the marsupial, the yellow-footed antechinus ( Antechinus flavipes ) as a model species for investigating phenotypic differentiation between animals inhabiting two habitat types in south-eastern Australia: flood-plain river red gum and box–ironbark forests. All tested phenotypic characteristics varied between years at the same sites and therefore were not useful for investigating morphological specialization that may lead to speciation. Males generally were significantly heavier when antechinus densities were lower, but exceptions were found, possibly related to food availability. Teat-number variation recently has been shown to be associated with habitat specialization and incipient speciation within Antechinus agilis . We investigated genetic differentiation associated with this trait in A. flavipes . Population genetic analyses of microsatellite genotypes and mitochondrial DNA haplotypes revealed that sympatric 12-, 13- and 14-teat females in Chiltern forest were part of one freely interbreeding population. Our parentage analyses found two cases where 13-teat mothers produced 12-teat daughters. This suggests either plasticity or paternal genetic influence on the offspring's teat-number phenotype. Laboratory matings may be required to resolve the extent to which teat number is heritable in A. flavipes .  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 303–314.     

Morphological convergence during pregnancy among predator and nonpredator populations of the livebearing fish Brachyrhaphis rhabdophora (Teleostei: Poeciliidae)

Predation can drive morphological divergence in prey populations, although examples of divergent selection are typically limited to nonreproductive individuals. In livebearing females, shape often changes drastically during pregnancy, reducing speed and mobility and enhancing susceptibility to predation. In the present study, we document morphological divergence among populations of nonreproductive female livebearing fish (Brachyrhaphis rhabdophora) in predator and nonpredator environments. We then test the hypothesis that shape differences among nonreproductive females are maintained among reproductive females between predator and nonpredator environments. Nonreproductive females in predator environments had larger caudal regions and more fusiform bodies than females in nonpredator environments; traits that are associated with burst speed in fish. Shape differences were maintained in reproductive females, although the magnitude of this difference declined relative to nonreproductive females, suggesting morphological convergence during pregnancy. Phenotypic change vector analysis revealed that females in predator environments became more similar to females in nonpredator environments in the transition from nonreproductive to reproductive. Furthermore, the level of reproductive allocation affected shape similarly between predator environments. These results suggest a life‐history constraint on morphology, in which predator‐driven morphological divergence among nonreproductive B. rhabdophora is not maintained at the same level during pregnancy. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 386–392.     

Reproductive isolating barriers between colour‐differentiated populations of an African annual killifish,Nothobranchius korthausae (Cyprinodontiformes)

Allopatric populations separated by vicariance events are expected to evolve reproductive isolating mechanisms as a result of disparate selection pressures and genetic drift. The appearance of reproductive isolating mechanisms may vary across taxa with differences in the opportunity for mate choice, and may be asymmetrical. In addition, premating barriers may be affected by individual mating experience. We used choice and no‐choice experiments to investigate reproductive isolation between two allopatric (island and mainland) and colour‐differentiated populations of an African annual fish, Nothobranchius korthausae. Assortative mating under experimental conditions was limited and asymmetrical. Preference for sympatric males was only expressed in nonvirgin females from one population. Virgin fish from both populations mated indiscriminately. No difference in the number of eggs laid, fertilization rate and hatching success was detected in no‐choice experiments. All mating combinations produced viable offspring and no postmating barriers were detected in terms of the performance and fertility of F1 hybrids. Overall, we found little evidence for significant reproductive isolation, which is in contrast with the related killifish taxa in which assortative mating can be strong, even among allopatric populations with no colour differentiation. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 62–72.     

Genetic detection of sperm from forced copulations between sympatric populations of Poecilia reticulata and Poecilia picta

The reproductive success of the sneaky mating tactic is difficult to determine in fish with internal fertilization. We approached this problem by developing an assay that allows the recovery of sperm DNA from the reproductive tract of females. This assay was used to test whether sperm transfer occurs between sympatric populations of the guppy, Poecilia reticulata , and its putative sister species, Poecilia picta . Six hundred females of both species from two sympatric sites in Trinidad were collected, and then the contents of the females' reproductive tracts were analysed using reciprocally diagnostic microsatellite markers. Conspecific sperm occurred in approximately 86% of females, whereas heterospecific sperm were found in only 4% of females. Because females of either species do not mate willingly with heterospecific males, the results indicate that sneaky mating results in the transfer of sperm. The data are consistent with the idea that sexual conflict can result in differences in the strength of behavioural isolation between the sexes, and they suggest that behavioural isolation is unlikely to have driven speciation between the guppy and P. picta .  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 397–402.     

Parallel speciation in Astyanax cave fish (Teleostei) in Northern Mexico

We investigated differentiation processes in the Neotropical fish Astyanax that represents a model system for examining adaptation to caves, including regressive evolution. In particular, we analyzed microsatellite and mitochondrial data of seven cave and seven surface populations from Mexico to test whether the evolution of the cave fish represents a case of parallel evolution. Our data revealed that Astyanax invaded northern Mexico across the Trans-Mexican Volcanic Belt at least three times and that populations of all three invasions adapted to subterranean habitats. Significant differentiation was found between the cave and surface populations. We did not observe gene flow between the strongly eye and pigment reduced old cave populations (Sabinos, Tinaja, Pachon) and the surface fish, even when syntopically occurring like in Yerbaniz cave. Little gene flow, if any, was found between cave populations, which are variable in eye and pigmentation (Micos, Chica, Caballo Moro caves), and surface fish. This suggests that the variability is due to their more recent origin rather than to hybridization. Finally, admixture of the young Chica cave fish population with nuclear markers from older cave fish demonstrates that gene flow between populations that independently colonized caves occurs. Thus, all criteria of parallel speciation are fulfilled. Moreover, the microsatellite data provide evidence that two co-occurring groups with small sunken eyes and externally visible eyes, respectively, differentiated within the partly lightened Caballo Moro karst window cave and might represent an example for incipient sympatric speciation.     

Incipient allochronic speciation by climatic disruption of the reproductive period

Disruptive selection of life-cycle timing may cause temporal isolation directly and, ultimately, allochronic speciation. Despite the fact that segregation of the reproductive period among related species has been broadly observed across taxa, it remains controversial whether temporal isolation can function as the primary process of speciation. In the Japanese winter geometrid moth Inurois punctigera, allochronic divergence has resulted from climatic disruption of the reproductive period. In habitats with severe midwinter, two sympatric groups of moth reproduce allochronically in early and late winter. These groups are genetically diverging sister lineages and now co-occur allochronically throughout Japan. By contrast, in habitats with milder midwinter these lineages form a continuous adult period and gene flow has been facilitated between the lineages. These results, together with the fact that there is no difference in larval host use, indicate that temporal isolation has been the sole mechanism for allochronic isolation in colder habitats and that allochrony is not a by-product of other adaptations. Thus, the allochronic divergence of sympatric I. punctigera populations represents an incipient speciation process driven by midwinter disruption of the reproductive period.     

A role for ecology in male mate discrimination of immigrant females in Calopteryx damselflies?

Sexual selection against immigrants is a mechanism that can regulate premating isolation between populations but, so far, few field studies have examined whether males can discriminate between immigrant and resident females. Males of the damselfly Calopteryx splendens show mate preferences and are able to force pre‐copulatory tandems. We related male mate responses to the ecological characteristics of female origin, geographic distances between populations, and morphological traits of females to identify factors influencing male mate discrimination. Significant heterogeneity between populations in male mate responses towards females was found. In some populations, males discriminated strongly against immigrant females, whereas the pattern was reversed or nonsignificant in other populations. Immigrant females were particularly attractive to males when they came from populations with similar predation pressures and densities of conspecifics. By contrast, immigrant females from populations with strongly dissimilar predation pressures and conspecific densities were not attractive to males. Differences in the abiotic environment appeared to affect mating success to a lesser degree. This suggests that male mate discrimination is context‐dependent and influenced by ecological differences between populations, a key prediction of ecological speciation theory. The results obtained in the present study suggest that gene‐flow is facilitated between ecologically similar populations. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 506–518.     

The offspring size/fecundity trade-off and female fitness in the Atlantic molly (<Emphasis Type="Italic">Poecilia mexicana</Emphasis>, Poeciliidae)

Across a variety of taxa, large offspring have been demonstrated to have a fitness advantage over smaller offspring of the same species. However, producing large offspring often comes at the cost of having to produce fewer young, and the payoff (and thus, evolutionary outcome) of this trade-off is expected to vary between environments. Atlantic mollies (Poecilia mexicana: Poeciliidae, Teleostei), inhabiting a sulfidic cave and various non-sulfidic surface habitats in Tabasco (Mexico), are reproductively isolated and evolved divergent female life-history traits: females of the cave ecotype produce considerably fewer, but larger offspring. Stressful (sulfidic) environments may favor the production of larger offspring, as they are better able to cope with chemical stressors. It remains to be determined though if increased offspring survival outweighs the fitness cost of producing fewer but larger offspring even under benign laboratory conditions. We tested 30-day newborn survival of offspring from wild-caught P. mexicana females from diverging populations in a low-density, no predation, no cannibalism, and ad-libitum-food, benign laboratory environment. Survival rates were highly skewed towards larger cave molly offspring; however, surface molly females still had a higher fitness than cave molly females in terms of higher total numbers of surviving offspring. Our study provides evidence for an innate fitness advantage of larger cave molly offspring. Furthermore, the observed differences in life-history strategies could promote further divergence and reproductive isolation among these ecotypes of P. mexicana, because cave molly females immigrating into the adjacent surface habitats would most likely be selected against.     

Evidence of incipient speciation in the Neotropical mangrove Pelliciera rhizophorae (Tetrameristaceae) as revealed by molecular,morphological, physiological and climatic characteristics

Two variants of the Neotropical mangrove species Pelliciera rhizophorae distributed along both sides of the Isthmus of Panama were detected by different colouration of the floral bracts and the size of the floral and vegetative structures. These findings raised questions concerning a possible speciation event in P. rhizophorae, for which a series of macro‐ and microscopic morphological traits (reproductive and vegetative structures), molecular markers from plastid DNA and climatic profiles were analyzed. Samples of P. rhizophorae were collected in three localities from the Panamanian Caribbean and Pacific coasts. The data obtained from molecular markers and morphological traits showed significant differences between the variants. The climatic profiles showed contrasting characteristics of rainfall and temperature in their habitats: variant A is found in wetter zones and variant B occupies drier zones. Evidence suggesting that a process of incipient speciation has occurred in P. rhizophorae in response to ecogeographical isolation due to climatic factors is presented. The presence of two geographically separate genetic‐morphological groups, adapted to contrasting climatic conditions, will be the basis for suggesting the existence of incipient lineages in Pelliciera. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 179 , 499–510.     

On the use of genetic divergence for identifying species

Degree of genetic divergence is frequently used to infer that two populations belong to separate species, or that several populations belong to a single species. I explore the logical framework of this approach, including the following assumptions: (i) speciation takes place over very long periods of time; (ii) reproductive isolation is based on the slow accumulation many genetic differences throughout the genome; (iii) genetic divergence automatically leads to reproductive isolation between species; and (iv) pre-mating and post-mating reproductive isolation have a similar genetic basis. I argue that so many exceptions to these assumptions have been demonstrated that they cannot be used with any reliability to distinguish different species. In addition, genetic distance as a species criterion is mostly used within the framework of Mayr's Biological Species Concept and is not free of assumptions about the nature of species or of speciation. The use of genetic distance to infer separate species (or the lack of these) is not parsimonious, its theoretical foundations are not well understood, and it cannot be applied over a wide range of plants and animals. I explore alternative approaches towards solving the species problems normally solved using genetic distance. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 75 , 509–516.