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Divergent selection and the evolution of signal traits and mating preferences.

Mating preferences are common in natural populations, and their divergence among populations is considered an important source of reproductive isolation during speciation. Although mechanisms for the divergence of mating preferences have received substantial theoretical treatment, complementary experimental tests are lacking. We conducted a laboratory evolution experiment, using the fruit fly Drosophila serrata, to explore the role of divergent selection between environments in the evolution of female mating preferences. Replicate populations of D. serrata were derived from a common ancestor and propagated in one of three resource environments: two novel environments and the ancestral laboratory environment. Adaptation to both novel environments involved changes in cuticular hydrocarbons, traits that predict mating success in these populations. Furthermore, female mating preferences for these cuticular hydrocarbons also diverged among populations. A component of this divergence occurred among treatment environments, accounting for at least 17.4% of the among-population divergence in linear mating preferences and 17.2% of the among-population divergence in nonlinear mating preferences. The divergence of mating preferences in correlation with environment is consistent with the classic by-product model of speciation in which premating isolation evolves as a side effect of divergent selection adapting populations to their different environments.

Adaptation, Biological↗

Sympatric, simultaneous, and random mating between markless trout (iwame; Oncorhynchus iwame) and red-spotted masu salmon (amago; Oncorhynchus masou ishikawae).

The markless trout (iwame, Oncorhynchus iwame) has long been suspected not to be an independent species, but rather a markless mutant form of the red-spotted masu salmon (amago, O. masou ishikawae). Nevertheless, no field study has examined this issue. Here we report a field study on the reproductive ecology of iwame and amago coexisting in the upper stream of the Inabe River. We found that iwame and amago mate sympatrically, simultaneously, and randomly, and observed five cases of heterogeneous oviposition between the two. Our results suggest that no pre-mating reproductive isolation exists between iwame and amago. Because previous studies have shown the absence of post-mating isolation, we propose that iwame and amago are the same species. Iwame should be considered as a markless form of the red-spotted masu salmon, and O. iwame as a junior synonym of O. masou ishikawae.

Animals↗

Mhc class II DNA polymorphisms within and between chromosomal species of the Spalax ehrenbergi superspecies in Israel.

Restriction fragment length polymorphisms (RFLPs) of two major histocompatibility class II genes (P alpha 1 and Q beta) were studied in 13 populations of four chromosomal species (2n = 52, 54, 58 and 60) of the mole rat, Spalax ehrenbergi superspecies in Israel. A substantial frequency of allelic fragments was found in both genes for all populations, including a desert isolate. In the P alpha 1 gene, one allelic fragment is a result of a deletion mutation which is diagnostic of the 2n = 52 chromosomal species. All other ten allelic variants are the result of point mutations. All mutations are located in a short region flanking the 3' end of the gene. Based on Mhc polymorphisms we confirm earlier evidence that gene flow does not occur between the older chromosomal species (2n = 52, 54, 58), and that reproductive isolation decreases, progressively from the oldest to the youngest species (2n = 60).

Animals↗

Surnames and genetic structure of a high-altitude Quechua community from the Ichu River Valley, Peruvian Central Andes, 1825-1914.

Changes in isolation, inbreeding, population subdivision, and isonymous relationships are examined in six Quechua communities from the upper valley of the Ichu River in the Peruvian Central Andes (3700 m). All marriages registered between 1825 and 1914 in the Parish of Santa Ana were analyzed. The data (1680 marriages) were divided into 2 periods (1825-1870 and 1871-1914) and into the 6 villages that constitute the parish. Endogamy rates are between 81% and 100%, indicating high levels of reproductive isolation. The inbreeding indicated by isonymy (Ft, Fr, and Fn) is lower than in other mountain populations studied. Isonymy values, calculated from the different surname combinations made possible by the Ibero-American Surnames System, indicate a strong rejection of consanguineous marriages, particularly between patrilineal relatives, in agreement with the parental structure typical of Andean populations. The comparison between observed and expected repeated-pair values reveals a moderate level of subdivision within populations, which could be related to cultural and socioeconomic factors. Nonmetric multidimensional scaling was used to investigate temporal changes in the isonymous relationships among the communities. The results reveal a decrease in the interpopulational variability measured by surnames, in agreement with an increase in exogamy. Surnames and data contained in historical and demographic records yield reliable information, and they can be used to reconstruct the biological history of Amerindian populations over the last few centuries.

Analysis of Variance↗

Molecular, morphological and behavioural data reveal the presence of a cryptic species in the widely studied Drosophila serrata species complex.

The Drosophila serrata species complex from Australia and New Guinea has been widely used in evolutionary studies of speciation and climatic adaptation. It is believed to consist of D. serrata, D. birchii and D. dominicana, although knowledge of the latter is limited. Here we present evidence for a previously undescribed cryptic member of the D. serrata species complex. This new cryptic species is widespread in far north Queensland, Australia and is likely to have been previously mistaken for D. serrata. It shows complete reproductive isolation when crossed with both D. serrata and D. birchii. The cryptic species can be easily distinguished from D. serrata and D. birchii using either microsatellite loci or visual techniques. Although it occurs sympatrically with both D. serrata and D. birchii, it differs from these species in development time, viability, wing size and wing morphology. Its discovery explains patterns of recently described mitochondrial DNA divergence within D. serrata, and may also help to clarify some ambiguities evident in early evolutionary literature on reproductive incompatibility within the D. serrata species complex.

Analysis of Variance↗

Sympatric speciation by sexual conflict.

It is well established that sexual conflict can drive an endless coevolutionary chase between the sexes potentially leading to genetic divergence of isolated populations and allopatric speciation. We present a simple mathematical model that shows that sexual conflict over mating rate can result in two other general regimes. First, rather than "running away" from males, females can diversify genetically into separate groups, effectively "trapping" the males in the middle at a state characterized by reduced mating success. Female diversification brings coevolutionary chase to the end. Second, under certain conditions, males respond to female diversification by diversifying themselves. This response results in the formation of reproductively isolated clusters of genotypes that emerge sympatrically.

Adaptation, Physiological↗

Isolation of porcine reproductive and respiratory syndrome (PRRS) virus from Heko-Heko disease of pigs.

Porcine reproductive and respiratory syndrome (PRRS) virus was isolated at high frequency from the sera and lungs of pigs affected with Heko-Heko disease. In addition, a considerable amount of Mycoplasma hyorhinis (Mhr) was also isolated from the lungs. Inoculation of gnotobiote pigs with the first isolate of PRRS virus resulted in the reproduction of proliferative and interstitial pneumonia. The virus was recovered from the inoculated pigs for long periods. Superinfection with PRRS virus and Mhr appeared to produce more serious pneumonia than inoculation with PRRS virus alone. In this study, the presence of PRRS virus was confirmed in Japan, and the PRRS virus was considered as being the most important pathogen for Heko-Heko disease.

Animals↗

The convoluted evolution of snail chirality.

The direction that a snail (Mollusca: Gastropoda) coils, whether dextral (right-handed) or sinistral (left-handed), originates in early development but is most easily observed in the shell form of the adult. Here, we review recent progress in understanding snail chirality from genetic, developmental and ecological perspectives. In the few species that have been characterized, chirality is determined by a single genetic locus with delayed inheritance, which means that the genotype is expressed in the mother's offspring. Although research lags behind the studies of asymmetry in the mouse and nematode, attempts to isolate the loci involved in snail chirality have begun, with the final aim of understanding how the axis of left-right asymmetry is established. In nature, most snail taxa (>90%) are dextral, but sinistrality is known from mutant individuals, populations within dextral species, entirely sinistral species, genera and even families. Ordinarily, it is expected that strong frequency-dependent selection should act against the establishment of new chiral types because the chiral minority have difficulty finding a suitable mating partner (their genitalia are on the 'wrong' side). Mixed populations should therefore not persist. Intriguingly, however, a very few land snail species, notably the subgenus Amphidromus sensu stricto, not only appear to mate randomly between different chiral types, but also have a stable, within-population chiral dimorphism, which suggests the involvement of a balancing factor. At the other end of the spectrum, in many species, different chiral types are unable to mate and so could be reproductively isolated from one another. However, while empirical data, models and simulations have indicated that chiral reversal must sometimes occur, it is rarely likely to lead to so-called 'single-gene' speciation. Nevertheless, chiral reversal could still be a contributing factor to speciation (or to divergence after speciation) when reproductive character displacement is involved. Understanding the establishment of chirality, the preponderance of dextral species and the rare instances of stable dimorphism is an important target for future research. Since the genetics of chirality have been studied in only a few pulmonate species, we also urge that more taxa, especially those from the sea, should be investigated.

Animal Structures↗

Reproductive processes in two oak (Quercus) contact zones with different levels of hybridization.

Patterns of reproductive isolation between two sympatric species of oaks, Quercus gambelii and Q. grisea, that exhibit strong ecological differentiation were examined. A full diallel cross using four trees of each species (i.e. all possible pollinations among eight trees) was performed. This design was repeated at two sites that represent different outcomes of sympatry: (1) a xeric mountain ridge where many hybrids are established (HZ); and (2) a mesic valley bottom where virtually no hybrids are established (MOCYN). By measuring fruit survival at several developmental stages, both the timing and strength of reproductive barriers within and between sites, species, cross types, and pollen dosage levels were examined. In three of four cases, heterospecific fruit set was significantly reduced compared to conspecific fruit set. This reduction occurred after the time of fertilization, but before the onset of embryo growth. Increasing the dose of pollen from an average of 9-194 grains/stigma did not affect this result. Thus, early postfertilization processes play a strong role in species fidelity in these oaks. Quercus gambelii experienced a five-fold decrease in conspecific fruit set at HZ relative to MOCYN. In contrast, heterospecific fruit set of Q. gambelii was the same at both sites. Poor Q. gambelii pollen performance is implicated as playing the major role in this result. One Q. gambelii individual at HZ was highly fecund, and had higher heterospecific than conspecific fruit set; slight introgression in this tree was detected uisng RAPD markers. The Environmental Emasculation Hypothesis that posits that environmental stress can increase the probability of hybrid formation by reducing the competitive ability of male gametes of one species is proposed.

Crosses, Genetic↗

Why do species exist? Insights from sexuals and asexuals.

Why does life diversify into the more or less discrete entities we recognise as species? Two main explanations have been proposed: i) species are a consequence of adaptation to different ecological niches, ii) species are a consequence of sexual reproduction and reproductive isolation. Phylogenetic studies of case-study groups can provide insights into the relative importance of divergent selection and isolation for speciation, but it can be difficult to infer causes of speciation unambiguously. The example of North American tiger beetles from the genus Cicindela is discussed. An alternative approach is to compare diversification between related sexual and asexual taxa to infer the relative importance of the two explanations. We outline expected patterns of diversification in sexual and asexual lineages under different scenarios using coalescent theory. Whether sexuals or asexuals diversify to a greater extent depends on the balance among various stages of diversification, particularly on the effects of sexual reproduction on rates of adaptive evolution. Rotifers offer a unique system to test these ideas, allowing comparison of patterns of genetic and functional morphological diversification in sexual (bdelloid) and asexual (monogonont) clades.

Journal Article↗

Genetic identification of two sibling species of Lutzomyia longipalpis (Diptera: Psychodidae) that produce distinct male sex pheromones in Sobral, Ceará State, Brazil.

Lutzomyia longipalpis, the main sandfly vector for New World visceral leishmaniasis is a complex of an as yet undefined number of sibling species. At present, there is no consensus on the status (single species vs. species complex) of Brazilian populations. We applied five microsatellite loci to test the hypothesis that L. longipalpis occurs as two sympatric cryptic species in Sobral, Ceará State, Brazil as predicted by male sex pheromone chemotypes described previously for field specimens from this site [S-9-methyl-germacrene-B (9MGB) and a cembrene compound]. Abdominal spot morphology corresponds with pheromone type at this locality (9MGB in '1 spot' males and cembrene in '2 spot' males). Genotype data from 190 wild-caught L. longipalpis specimens collected in October 1999 and April 2001 were used to estimate genetic differentiation between the two sex pheromone populations and sampling dates. No significant (P > 0.05) genetic differences were found between the 1999 and 2001 9MGB samples (theta = 0.018; RST = -0.005), and genetic differentiation was low between the cembrene collections (theta = 0.037, P < 0.05; RST = -0.043, P > 0.05). By contrast, highly divergent allelic frequencies (largely at two microsatellite loci) corresponded to significant (P > 0.05) genetic differentiation (theta = 0.221; RST = 0.215) for all comparisons between samples with different pheromones. When pheromone samples were pooled across sample date, genetic differentiation was high (theta = 0.229; P < 0.001; Nem = 0.84). The allele frequency distribution at each of the five microsatellite loci was similar for males and females from the two collection years. Two of these loci showed highly divergent allele frequencies in the two sex pheromone populations. This was reflected in the highly significant genetic differentiation obtained from the male genotypes, between populations producing different pheromones (theta = 0.229-0.268; P < 0.0001 for the 2001 and theta = 0.254-0.558; P < 0.0001 for the 1999 collections, respectively). Similar results were obtained when the females, assigned to a pheromone type, were included in the analysis. Both a Bayesian analysis of the data set and a population assignment test provided strong evidence for two distinct populations corresponding to pheromone type. Given its genotype, the probability of assigning a 9MGB male to the original 9MGB population was 100% once the two years' collections were pooled. For cembrene-producing '2 spot' males this probability although still high, was lower than for 9MGB males, at 86%. This microsatellite data together with previously reported reproductive isolation between the two Sobral populations confirm that premating barriers are important in speciation of L. longipalpis.

Animals↗

Fruit odor discrimination and sympatric host race formation in Rhagoletis.

Rhagoletis pomonella is a model for incipient sympatric speciation (divergence without geographic isolation) by host-plant shifts. Here, we show that historically derived apple- and ancestral hawthorn-infesting host races of the fly use fruit odor as a key olfactory cue to help distinguish between their respective plants. In flight-tunnel assays and field tests, apple and hawthorn flies preferentially oriented to, and were captured with, chemical blends of their natal fruit volatiles. Because R. pomonella rendezvous on or near the unabscised fruit of their hosts to mate, the behavioral preference for apple vs. hawthorn fruit odor translates directly into premating reproductive isolation between the fly races. We have therefore identified a key and recently evolved (<150 years) mechanism responsible for host choice in R. pomonella bearing directly on sympatric host race formation and speciation.

Animals↗

Water-borne sperm trigger vitellogenic egg growth in two sessile marine invertebrates.

A diverse array of sessile marine invertebrates mate by passive dispersal of sperm which fertilize the brooded eggs of neighbours. In two such species, a sea-mat (phylum Bryozoa) and an ascidian (phylum Chordata), vitellogenic egg growth is absent in reproductively isolated specimens, but is triggered by a water-borne factor released by conspecifics. In both of these colonial, hermaphroditic species, the active factor can be removed from water by filtration. The effect involves self-/non-self-recognition: water conditioned by a separate subcolony of the same genetic individual does not prompt oocyte growth. In each species, allosperm move from the surrounding water to the ovary and are then stored in close association with the growing oocytes. We concluded that sperm themselves are the water-borne factor that triggers the major phase of female reproductive investment. This mechanism is, to our knowledge, previously undescribed in animals, but has parallels with the initiation of maternal investment in flowering plants following the receipt of compatible pollen. The species studied may be representative of many other aquatic invertebrates which mate in a similar way. The stimulation of egg growth by allosperm could lead to intersexual conflict during oogenesis.

Animals↗

Genetic structure of a Japanese allotetraploid loach of the genus Cobitis (Osteichthyes, Cobitidae).

The Japanese allotetraploid spined loach of the genus Cobitis "yamato complex" sensu SAITOH et al. (2000), distributed in Western Japan, originated from hybridization between C. biwae on the maternal side and C. striata (Kyushu form) on the paternal side. Mitochondrial (mt) and nuclear DNA were analyzed in order to determine the genetic relationships among 15 populations spanning the entire range of the yamato complex. PCR-RFLP analysis of the ND1 mtDNA gene indicated that the yamato complex contains two divergent types of mtDNA: type A, consisting of one haplotype observed only in the Fukagawa River and type B consisting of 12 haplotypes found in the entire area. Phylogenetic analysis based on the cytochrome b mtDNA gene corroborated RFLP analysis, and indicated that type A was closely related to a different species, C. biwae (Kochi group) and C. striata (large race), rather than type B. The results of RAPD analysis on the Fukagawa River individuals, where types A and B sympatrically existed suggested that no reproductive isolation occurs between them. The existence of two distinct mtDNA types within the yamato complex suggest either multiple maternal origin at the speciation (tetraploidization) time or mtDNA introgression from other species afterwards.

Animals↗

DNA fingerprints of Trichinella as revealed by restriction fragment length polymorphism and single-strand conformational polymorphism (RFLP-SSCP).

A method was developed for gene fingerprinting, combining the principles of restriction fragment length polymorphism and single-strand conformational polymorphism (PCR-RFLP-SSCP). Taking advantage of this method, we analysed the genotypes of 20 isolates from five species of Trichinella (Trichinella spiralis, Trichinella britovi, Trichinella nativa, Trichinella murrelli and Trichinella pseudospiralis) and two uncertain genotypes (Trichinella T6 and Trichinella T8). Target genes for the analysis included three kinds of random amplified polymorphic DNA (RAPD), the gene encoding 43 kDa excretory-secretory protein, the gene encoding mitochondrial cytochrome c -oxidase subunit I and the gene of 18 S rRNA. The genotype revealed by this method was in good concordance with the taxonomy of Trichinella (six species and two uncertain genotypes which is currently accepted based on morphology, isozyme pattern and reproductive isolation). More important, this method revealed intraspecies polymorphism among geographical isolates of T. spiralis and T. britovi.

Animals↗

Loss of genetic diversity in the endemic Hector's dolphin due to fisheries-related mortality.

The endemic New Zealand Hector's dolphin is considered the rarest species of marine dolphin with a total abundance of less than 4000. The species is listed as vulnerable because of fisheries-related mortality due to entanglement in set nets. The vulnerability of this species is further increased by its fidelity to local natal ranges and the genetic isolation of regional populations. Here we present evidence, based on 108 contemporary samples and 55 historical samples dating back to 1870, of a significant loss of mitochondrial DNA (mtDNA) diversity in two regional populations of Hector's dolphin. The haplotype diversity (h) was calculated from sequences of a 206 bp fragment in the mtDNA control region, designed to identify 13 out of the 14 known maternal lineages. Over the last 20 years, the North Island population has been reduced from at least three lineages (h = 0.41) to a single lineage (h = 0; p < 0.05). Given its small size, reproductive isolation and reduced genetic diversity, this population is likely to become extinct. The diversity of the East Coast South Island population has declined significantly from h = 0.65 to h = 0.35 (p < 0.05). Based on trend analysis of the mtDNA diversity, we predict that the East Coast population will lose all mtDNA diversity within the next 20 years. This time-series of reduction in genetic variation provides independent evidence of the severity of population decline and habitat contraction resulting from fisheries and perhaps other human activities.

Animals↗

Evidence for sympatric speciation by host shift in the sea.

The genetic divergence and evolution of new species within the geographic range of a single population (sympatric speciation) contrasts with the well-established doctrine that speciation occurs when populations become geographically isolated (allopatric speciation). Although there is considerable theoretical support for sympatric speciation, this mode of diversification remains controversial, at least in part because there are few well-supported examples. We use a combination of molecular, ecological, and biogeographical data to build a case for sympatric speciation by host shift in a new species of coral-dwelling fish (genus Gobiodon). We propose that competition for preferred coral habitats drives host shifts in Gobiodon and that the high diversity of corals provides the source of novel, unoccupied habitats. Disruptive selection in conjunction with strong host fidelity could promote rapid reproductive isolation and ultimately lead to species divergence. Our hypothesis is analogous to sympatric speciation by host shift in phytophagous insects except that we propose a primary role for intraspecific competition in the process of speciation. The fundamental similarity between these fishes and insects is a specialized and intimate relationship with their hosts that makes them ideal candidates for speciation by host shift.

Animals↗

Functional incompatibility between the fertilization systems of two allopatric populations of Callosobruchus maculatus (Coleoptera: Bruchidae).

Recent studies indicate that postcopulatory sexual selection may represent an important component of the speciation process by initiating reproductive isolation via the evolutionary divergence of fertilization systems. Using two geographically isolated populations of the polyandrous beetle Callosobruchus maculatus, we investigated divergence in fertilization systems by determining the extent of postcopulatory functional incompatibility. Through reciprocal, cross-population matings we were able to separately estimate the effects of male and female population origin and their interaction on the extent of last-male sperm precedence, female receptivity to further copulation and female oviposition. Our results indicate partial incompatibility between the fertilization systems of the two populations at all three functional levels. Males derived from the same population as females outcompete rival, allopatric males with respect to sperm preemption, sperm protection, and ability to stimulate female oviposition. This pattern is reciprocated in both populations indicating that postcopulatory, prezygotic events represent important mechanisms by which between-population gene flow is reduced. We suggest the partial gametic isolation observed is a by-product of the coevolution of male and female fertilization systems by a process of cryptic female choice. Our results are consistent with a mechanism akin to conventional mate choice models although they do not allow us to reject antagonistic sexual coevolution as the mechanism of cryptic female choice.

Animals↗