Behavioral Reproductive Isolation Research Articles

Asymmetrical mate preference in recently adapted White Sands and black lava populations of Sceloporus undulatus

Abstract Speciation can proceed rapidly when natural and sexual selection act in concert. For example speciation can be accelerated when traits that confer a selective advantage in a particular habitat also influence mate preference. Studying parallel but evolutionarily independent instances of ecological divergence can illuminate the interaction between natural and sexual selection during speciation. Locally adapted populations of the eastern fence lizard Sceloporus undulatus have recently evolved in three different habitats in the Chihuahuan desert: blanched color morphs occur on the gypsum dunes of White Sands, melanic color morphs occur on the Carrizozo lava flow, and brown color morphs occur in the surrounding desert scrubland. In addition to differences in cryptic dorsal coloration, populations also differ in the size and color of ventral patches used for social signaling. This system therefore provides an opportunity to investigate the interplay of natural and sexual selection during rapid ecological speci-ation. We used mate preference experiments to determine whether locally adapted populations may exhibit the early stages of behavioral reproductive isolation. We observed an asymmetrical mate preference in this system; White Sands males preferentially courted local females, while males from dark soils and black lava populations did not exhibit a preference for local mates. We also found that female behavior and ventral patch phenotype were associated with male courtship. Our results suggest that the observed preference for local mates evolved at White Sands, and we discuss the possible link between local adaptation and traits involved in mate preference in this system.

Hydrocarbon divergence and reproductive isolation in Timema stick insects

BackgroundIndividuals commonly prefer certain trait values over others when choosing their mates. If such preferences diverge between populations, they can generate behavioral reproductive isolation and thereby contribute to speciation. Reproductive isolation in insects often involves chemical communication, and cuticular hydrocarbons, in particular, serve as mate recognition signals in many species. We combined data on female cuticular hydrocarbons, interspecific mating propensity, and phylogenetics to evaluate the role of cuticular hydrocarbons in diversification of Timema walking-sticks.ResultsHydrocarbon profiles differed substantially among the nine analyzed species, as well as between partially reproductively-isolated T. cristinae populations adapted to different host plants. In no-choice trials, mating was more likely between species with similar than divergent hydrocarbon profiles, even after correcting for genetic divergences. The macroevolution of hydrocarbon profiles, along a Timema species phylogeny, fits best with a punctuated model of phenotypic change concentrated around speciation events, consistent with change driven by selection during the evolution of reproductive isolation.ConclusionAltogether, our data indicate that cuticular hydrocarbon profiles vary among Timema species and populations, and that most evolutionary change in hydrocarbon profiles occurs in association with speciation events. Similarities in hydrocarbon profiles between species are correlated with interspecific mating propensities, suggesting a role for cuticular hydrocarbon profiles in mate choice and speciation in the genus Timema.

The relationship between intraspecific assortative mating and reproductive isolation between divergent populations

Abstract The term ‘assortative mating’ has been applied to describe two very different phenomena: (1) the tendency for individuals to choose phenotypically similar mates from among conspecifics; or (2) the tendency to prefer conspecific over hete-rospecific mates (behavioral reproductive isolation). Both forms of assortative mating are widespread in nature, but the relationship between these behaviors remains unclear. Namely, it is plausible that a preference for phenotypically similar conspecifics incidentally reduces the probability of mating with phenotypically divergent heterospecifics. We present a model to calculate how the level of reproductive isolation depends on intraspecific assortative mating and the phenotypic divergence between species. For empirically reasonable levels of intraspecific assortment on a single trait axis, we show that strong reproductive isolation requires very substantial phenotypic divergence. We illustrate this point by applying our model to empirical data from threespine stickleback Gasterosteus aculeatus and Darwin’s Finches (Geospiza spp). We conclude that typical levels of intraspecific assortment cannot generally be extrapolated to explain levels of interspecific reproductive isolation. Instead, reproductive isolation between species likely arises from different mate choice behaviors, or multivariate assortative mating.

Mate choice and the genetic basis for colour variation in a polymorphic dart frog: inferences from a wild pedigree

Understanding how reproductive barriers evolve during speciation remains an important question in evolution. Divergence in mating preferences may be a common first step in this process. The striking colour pattern diversity of strawberry dart frog (Dendrobates pumilio) populations has likely been shaped by sexual selection. Previous laboratory studies have shown that females attend to male coloration and prefer to court with males of their own colour, suggesting that divergent morphs may be reproductively isolated. To test this hypothesis, we used molecular data to estimate pedigree relationships from a polymorphic population. Whereas in the laboratory both red and yellow females preferred to court with males of their own phenotype, our pedigree shows a pattern of assortative mating only for red females. In the wild, yellow females appear to be less choosy about their mates, perhaps because they incur higher costs associated with searching than females of the more common red phenotype. We also used our pedigree to investigate the genetic basis for colour-pattern variation. The phenotype frequencies we observed were consistent with those expected if dorsal background coloration is controlled by a single locus, with complete dominance of red over yellow. Our results not only help clarify the role of sexual selection in reducing gene flow, but also shed light on the mechanisms underlying colour-pattern variation among sympatric colour morphs. The difference we observed between mating preferences measured under laboratory conditions and the pattern of mate choice observed in the wild highlight the importance of field studies for understanding behavioural reproductive isolation.

Asymmetric Mating Isolation between Two Caridean Shrimp:Lysmata wurdemanniandLysmata boggessi

ABSTRACT In many animals, sex pheromones are involved in behavioral reproductive isolation, which is thought to be more important than other isolation barriers in causing rapid speciation. Lysmata species living in the western Atlantic waters have been recently redefined taxonomically, with some of the species having overlapping distribution. It has been found that prezygotic mating isolation is not complete between two Lysmata species—Lysmata wurdemanni and Lysmata boggessi—and there is asymmetry in mate recognition between the two species. Male-role L. wurdemanni can mate with female-role L. boggessi, but not vice versa. This study shows that the asymmetric mating isolation is caused by the difference in sex pheromone detection (both distance and contact sex pheromones). Male L. wurdemanni can detect and respond to distance sex pheromones secreted by female L. boggessi, even at low concentrations, but male L. boggessi did not respond to distance sex pheromones of L. wurdemanni at any concentration, whic...

Behavioural reproductive isolation and speciation in Drosophila

The origin of premating reproductive isolation continues to help elucidate the process of speciation and is the central event in the evolution of biological species. Therefore, during the process of species formation the diverging populations must acquire some means of reproductive isolation so that the genes from one gene pool are prevented from dispersing freely into a foreign gene pool. In the genus Drosophila, the phenomenon of behavioural reproductive isolation, which is an important type of premating (prezygotic) reproductive isolating mechanisms, has been extensively studied and interesting data have been documented. In many cases incomplete sexual isolation has been observed and the pattern and degree of isolation within and between the species have often been used to elucidate the phylogenetic relationships. The present review documents an overview of speciation mediated through behavioural incompatibility in different species groups of Drosophila with particular reference to the models proposed on the basis of one-sided ethological isolation to predict the direction of evolution. This study is crucial for understanding the mechanism of speciation through behavioural incompatibility and also for an understanding of speciation genetics in future prospects.

Repeated Unidirectional Introgression of Nuclear and Mitochondrial DNA Between Four Congeneric Tanganyikan Cichlids

With an increasing number of reported cases of hybridization and introgression, interspecific gene flow between animals has recently become a widely accepted and broadly studied phenomenon. In this study, we examine patterns of hybridization and introgression in Ophthalmotilapia spp., a genus of cichlid fish from Lake Tanganyika, using mitochondrial and nuclear DNA from all four species in the genus and including specimens from over 800 km of shoreline. These four species have very different, partially overlapping distribution ranges, thus allowing us to study in detail patterns of gene flow between sympatric and allopatric populations of the different species. We show that a significant proportion of individuals of the lake-wide distributed O. nasuta carry mitochondrial and/or nuclear DNA typical of other Ophthalmotilapia species. Strikingly, all such individuals were found in populations living in sympatry with each of the other Ophthalmotilapia species, strongly suggesting that this pattern originated by repeated and independent episodes of genetic exchange in different parts of the lake, with unidirectional introgression occurring into O. nasuta. Our analysis rejects the hypotheses that unidirectional introgression is caused by natural selection favoring heterospecific DNA, by skewed abundances of Ophthalmotilapia species or by hybridization events occurring during a putative spatial expansion in O. nasuta. Instead, cytonuclear incompatibilities or asymmetric behavioral reproductive isolation seem to have driven repeated, unidirectional introgression of nuclear and mitochondrial DNA into O. nasuta in different parts of the lake.

Genetic differentiation, behavioural reproductive isolation and mixis cues in three sibling species of Monogonont rotifers

Many aquatic species usually considered to be 'cosmopolitan' have been identified as cryptic species complexes, based on deep genetic differentiation. However, reproductive isolation among sibling cryptic species has rarely been studied, and interspecific hybridization is common in some taxa.We investigated isolation mechanisms and possible introgression among three cyclical parthenogenetic rotifer species in the Epiphanes senta complex that are found in very different freshwater habitats: temperate floodplains, subtropical desert rock pools and a tropical alpine lake. Whereas Epiphanes ukera is reproductively isolated from E. chihuahuaensis and E. hawaiiensis, the latter hybridize under laboratory conditions.While reproductive isolation is incomplete, RAPD profiles indicated unique genetic signatures and showed no evidence for introgression, indicating that these three species are diverging and have independent evolutionary trajectories.Testing cues for sexual reproduction in these cyclic parthenogens demonstrated that mixis in E. chihuahuaensis and E. ukera is influenced by population density, whereas E. hawaiiensis females rarely produce mictic offspring regardless of density. Different mixis cues are likely to separate sexual periods and effectively cause reproductive isolation between the species. Epiphanes ukera and E. chihuahuaensis males display mate guarding behaviour, and E. ukera males distinguish between conspecific and heterospecific females in mate choice experiments. Geographic isolation, along with different cues for mixis induction and mate recognition, act as reproductive barriers among these sibling species.

Allelic variation in a fatty-acyl reductase gene causes divergence in moth sex pheromones

Pheromone-based behaviours are crucial in animals from insects to mammals, and reproductive isolation is often based on pheromone differences. However, the genetic mechanisms by which pheromone signals change during the evolution of new species are largely unknown. In the sexual communication system of moths (Insecta: Lepidoptera), females emit a species-specific pheromone blend that attracts males over long distances. The European corn borer, Ostrinia nubilalis, consists of two sex pheromone races, Z and E, that use different ratios of the cis and trans isomers of acetate pheromone components. This subtle difference leads to strong reproductive isolation in the field between the two races, which could represent a first step in speciation. Female sex pheromone production and male behavioural response are under the control of different major genes, but the identity of these genes is unknown. Here we show that allelic variation in a fatty-acyl reductase gene essential for pheromone biosynthesis accounts for the phenotypic variation in female pheromone production, leading to race-specific signals. Both the cis and trans isomers of the pheromone precursors are produced by both races, but the precursors are differentially reduced to yield opposite ratios in the final pheromone blend as a result of the substrate specificity of the enzymes encoded by the Z and E alleles. This is the first functional characterization of a gene contributing to intraspecific behavioural reproductive isolation in moths, highlighting the importance of evolutionary diversification in a lepidopteran-specific family of reductases. Accumulation of substitutions in the coding region of a single biosynthetic enzyme can produce pheromone differences resulting in reproductive isolation, with speciation as a potential end result.

From wild animals to domestic pets, an evolutionary view of domestication

Artificial selection is the selection of advantageous natural variation for human ends and is the mechanism by which most domestic species evolved. Most domesticates have their origin in one of a few historic centers of domestication as farm animals. Two notable exceptions are cats and dogs. Wolf domestication was initiated late in the Mesolithic when humans were nomadic hunter-gatherers. Those wolves less afraid of humans scavenged nomadic hunting camps and over time developed utility, initially as guards warning of approaching animals or other nomadic bands and soon thereafter as hunters, an attribute tuned by artificial selection. The first domestic cats had limited utility and initiated their domestication among the earliest agricultural Neolithic settlements in the Near East. Wildcat domestication occurred through a self-selective process in which behavioral reproductive isolation evolved as a correlated character of assortative mating coupled to habitat choice for urban environments. Eurasian wildcats initiated domestication and their evolution to companion animals was initially a process of natural, rather than artificial, selection over time driven during their sympatry with forbear wildcats.

Assortative Mating between European Corn Borer Pheromone Races: Beyond Assortative Meeting

BackgroundSex pheromone communication systems may be a major force driving moth speciation by causing behavioral reproductive isolation via assortative meeting of conspecific individuals. The ‘E’ and ‘Z’ pheromone races of the European corn borer (ECB) are a textbook example in this respect. ‘Z’ females produce and ‘Z’ males preferentially respond to a ‘Z’ pheromone blend, while the ‘E’ race communicates via an ‘E’ blend. Both races do not freely hybridize in nature and their populations are genetically differentiated. A straightforward explanation would be that their reproductive isolation is a mere consequence of “assortative meeting” resulting from their different pheromones specifically attracting males towards same-race females at long range. However, previous laboratory experiments and those performed here show that even when moths are paired in a small box – i.e., when the meeting between sexual partners is forced – inter-race couples still have a lower mating success than intra-race ones. Hence, either the difference in attractivity of E vs. Z pheromones for males of either race still holds at short distance or the reproductive isolation between E and Z moths may not only be favoured by assortative meeting, but must also result from an additional mechanism ensuring significant assortative mating at close range. Here, we test whether this close-range mechanism is linked to the E/Z female sex pheromone communication system.Methodology/Principal FindingsUsing crosses and backcrosses of E and Z strains, we found no difference in mating success between full-sisters emitting different sex pheromones. Conversely, the mating success of females with identical pheromone types but different coefficients of relatedness to the two parental strains was significantly different, and was higher when their genetic background was closer to that of their male partner's pheromone race.Conclusions/SignificanceWe conclude that the close-range mechanism ensuring assortative mating between the E and Z ECB pheromone races is unrelated to the difference in female sex pheromone. Although the nature of this mechanism remains elusive, our results show that it is expressed in females, acts at close range, segregates independently of the autosome carrying Pher and of both sex chromosomes, and is widely distributed since it occurs both in France and in the USA.

Behavioural Reproductive Isolation in a Rotifer Hybrid Zone

Behavioural Reproductive Isolation in a Rotifer Hybrid Zone

Intraspecific sexual selection on a speciation trait, male coloration, in the Lake Victoria cichlid Pundamilia nyererei

The haplochromine cichlids of Lake Victoria constitute a classical example of explosive speciation. Extensive intra- and interspecific variation in male nuptial coloration and female mating preferences, in the absence of postzygotic isolation between species, has inspired the hypothesis that sexual selection has been a driving force in the origin of this species flock. This hypothesis rests on the premise that the phenotypic traits that underlie behavioural reproductive isolation between sister species diverged under sexual selection within a species. We test this premise in a Lake Victoria cichlid, by using laboratory experiments and field observations. We report that a male colour trait, which has previously been shown to be important for behavioural reproductive isolation between this species and a close relative, is under directional sexual selection by female mate choice within this species. This is consistent with the hypothesis that female choice has driven the divergence in male coloration between the two species. We also find that male territoriality is vital for male reproductive success and that multiple mating by females is common.

Behavioral Isolation, Test Designs, and Kaneshiro's Hypothesis

Studies of the planitibia group of the Hawaiian Drosophila inspired the controversial hypothesis that the direction of evolution can be inferred by examining the direction of asymmetrical behavioral reproductive isolation. One criticism of this hypothesis is based on the methods that were used to develop it: the sign and magnitude of the indexes of behavioral reproductive isolation may vary with the procedures employed to measure them. We evaluated this criticism by applying three common methods used for measuring behavioral reproductive isolation to the species D. heteroneura and D. silvestris. These are the species that were used in the original development of the behavioral isolation hypothesis. The methods were female choice, male choice, and multiple choice. We found a similar direction of asymmetry for all three testing methods in the pairing of D. silvestris and D. heteroneura. In contrast, the pairing between two populations of D. silvestris did not give a consistent direction of asymmetry. Thus, different methods for testing the direction of behavioral reproductive isolation give similar results at the interspecific level but are inconsistent at the intraspecific level.

Host plant influences on sex pheromone behavior of phytophagous insects.

The sexual behavior of phytophagous insects is often integrated in a variety of ways with their host plants. This integration may be manifested as effects or influences of host plants on insect physiology and behavior, including sex pheromone communication, that reflect strategies by insects to optimize mating and reproduction. Certain insects sequester or otherwise acquire host plant compounds and use them as sex pheromones or sex pheromone precursors. Other insects produce or release sex pheromones in response to particular host plant cues. Chemicals from host plants often synergize or otherwise enhance insect responses to sex pheromones. By these means, host plants may be used by insects to regulate or mediate sexual communication. For many species of insects, host plant influences on insect sex pheromone communication may be important aspects of the formation of feeding and mating aggregations, of insect strategies to locate both hosts and mates, of behavioral reproductive isolation among sibling species, and of the regulation of reproduction to coincide with the availability of food and oviposition sites. Knowledge of these relationships is critical to understanding many different areas of the behavioral ecology of plant-feeding insects.

Behavioral reproductive isolation among sympatric strains of Brachionus plicatilis M�ller 1786: insights into the status of this taxonomic species

Behavioral reproductive isolation among sympatric strains of Brachionus plicatilis M�ller 1786: insights into the status of this taxonomic species

Small Genetic Distances among Populations of Green Lacewings of the Genus Chrysoperla (Neuroptera: Chrysopidae)

Seven populations of four species of green lacewings of the genus Chrysoperla were examined for variation at 17 electrophoretically detectable loci. Objectives were (1) to determine the degree of genetic differentiation, if any, that has occurred between two cryptic sibling species, C. plorabunda (Fitch) and C. johnsoni (Henry et al.), which can be distinguished only by differences in their courtship songs and their behavioral reproductive isolation; and (2) to compare those two species with the closely related C. downesi (Smith) and the more distantly related C. harrisii (Fitch). Most of the genetic differentiation in these populations was caused by differences between the cryptic species C. plorabunda and C.johnsoni on the one hand and each of the other two species. Nei's and Roger's genetic distances among all populations were not significantly different. The only significant node was that separating C. harrisii from the other populations. Phenograms from unweighted pair-group method using arithmetic averages (UPGMA) did not resolve the cluster of populations of C. plorabunda and C.johnsoni. These cryptic sibling species are genetically very similar, suggesting recent speciation. Examination of more variable electrophoretic loci, or use of an alternative method such as mtDNA restriction site analysis, might provide enough differences to separate these otherwise good biological species.

Chemical characterization and species specificity of sex pheromones of plusiinae moths in Israel

AbstractThe female sex pheromones of four sympatric Plusiinae species, Autographa gamma, Chrysodeixis chalcites, Cornutiplusia circumflexa, and Trichoplusia ni were investigated in Israel. The research concentrated on the behavioral reproductive isolation regulated by their sex pheromones. The chemical composition of the four sex pheromones was determined by analysis of pheromone gland extracts, and in addition entrained volatiles released from virgin female C. chalcites and T. ni were also analyzed. The pheromones of these latter two species were found to contain several components, among which Z7‐12:Ac was the main one. The pheromone glands of A. gamma and C. circumflexa were found to contain relatively small amounts of material and only Z7–12:Ac and Z7–12:OH could be positively identified. These two pheromone components were present in the two species in roughly inverse rations. Whereas A. gamma utilizes Z7–12:Ac as the main or sole pheromone component, C. circumflexa requires both Z7–12:OH and Z7–12:Ac, with the alcohol being predominant. This is the first reported Plusiinae species using Z7–12:OH as the main pheromone component. Behavioral studies in the wind tunnel and field tests indicate that females of each of the four Plusiinae species utilizes a species‐specific blend of chemicals, consisting of attractive and antagonistic components, to attract conspecific males. Such a complementary strategy provides improved behavioral reproductive isolation among the sympatric species. For example, Z5–12:Ac, which is a specific component of the T. ni sex pheromone, may be redundant as regards the attraction of T. ni males. However, addition of small amounts of this compound to pheromone blends of A. gamma and C. chalcites resulted in complete inhibition of the males' flight behavior and trap catch; thus contributing significantly to the specificity of the T. ni sex pheromone. The relatively high proportion of Z7–12:OH in the sex pheromone of C. circumflexa, essential to the latter, is highly inhinitory to the three other Plusiinae species. The only case of cross‐attraction occurred when T. ni males were attracted to a certain extent to A. gamma lures and females. © 1993 Wiley‐Liss, Inc.

Biological comparisons between arrhenotokous and thelytokous biotypes ofMesochorus nigripes [Hym.: Ichneumonidae

The host preferences and sex ratios ofMesochorus biotypes from Sweden and Colorado (U.S.A.) were studied in the field and laboratory. TheseMesochorus are secondary parasites ofBathyplectes spp., parasites ofHypera postica (Gyllenhal). The Colorado biotype is 100% thelytokous, and clearly prefersBathyplectes stenostigma (Thomson) as a host. Only an arrhenotokous biotype was positively detected in Sweden, and it appeared to preferB. curculionis (Thomson). However, some of the data suggest that the thelytokous biotype was probably also present in Sweden, at least inB. stenostigma. The behavioral differences and reproductive isolation of the thelytokous and arrhenotokous biotypes indicate that they are separate sibling species, even though they cannot yet be separated morphologically, and isozyme analyses were inconclusive (possibly due to a mixture of biotypes in Sweden).

Role of female-produced sex pheromone in behavioral reproductive isolation betweenTrichoplusia ni (H�bner) andPseudoplusia includens (Walker) (Lepidoptera: Noctuidae, plusiinae)

In laboratory flight tunnel bioassays, response rates of male cabbage looper,Trichoplusia ni (Hübner), to female soybean looper,Pseudoplusia includens (Walker), were similar to response rates of maleT. ni to conspecific females for plume tracking and source contact. Male soybean loopers, however, exhibited a greatly reduced response to female cabbage loopers compared to conspecific females. Similar differences were observed in male responses to extracts of female abdominal tips. Studies of flight tunnel responses of male soybean loopers to the different chemicals known to be components of the female cabbage looper sex pheromone indicated that the reduction in response was due to inhibitory effects of (Z)-5-dodecen-1-ol acetate and (Z)-9-tetradecen-1-ol acetate, when added singly to (Z)-7-dodecen-1-ol acetate (major component of both species) at release rates and at ratios close to those observed in female cabbage loopers.