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PubMed2006

Multiple paternity and breeding system in the gopher tortoise, Gopherus polyphemus.

Little is known about the reproductive behaviors and the actual outcomes of mating attempts in the gopher tortoise (Gopherus polyphemus). We examined the mating system and reproductive behaviors of a population of gopher tortoises in central Florida. Using microsatellite markers, we assigned fathers to the offspring of seven clutches and determined that multiple fathers were present in two of the seven clutches examined. We found that gopher tortoises exhibited a promiscuous mating system with larger males fertilizing the majority of clutches. The advantage of larger males over smaller males in fertilizing females may be a result of larger males winning access to females in aggressive bouts with other males or larger males may be more attractive to females. Clutches produced by larger females tended to be sired by a single male, whereas clutches of smaller females tended to be sired by multiple males.

Animals↗
PubMed2002

Divergence in mate choice systems: does evolution play by rules?

Understanding the genetic bases of phenotypes associated with the earliest stages of divergence will reveal a great deal about species formation. I review a number of model systems, most involving plant-insect interactions, that have already revealed genetic aspects of incipient speciation. It is suggested that progress in understanding the causal forces driving mating signal evolution and incipient speciation will be expedited in model systems where; (1) ecological and evolutionary information is available, (2) different aspects of mating behaviors that function in mate and/or species recognition are known, (3) genetic analysis of single phenotypes is undertaken, (4) analysis of sexual selection and isolation is performed under natural conditions (or in the wild), and (5) comparative data from related species are available to assess phylogenetic trends.

Animals↗
PubMed2005

Faithful without care: the evolution of monogyny.

The traditional paradigm of male polygamy and female monogamy has been replaced by the recognition that both sexes may typically mate with several partners. As a consequence, much attention has focused on the evolution of polyandry, while the evolutionary significance of monogyny (male monogamy) remains poorly understood. Monogyny, a taxonomically widespread mating system that includes dramatic examples of male self-sacrifice, is predicted when the benefits of paternal investment exceed those of searching for additional mates. However, monogyny also occurs in animals lacking paternal investment, instead representing a form of paternity protection. It has been suggested that such mating systems are expected where the costs of mate search for males are high. However, this argument fails to recognize that if there is a low probability of a male finding a mate, then there may be a high probability that he will not need to defend his paternity. Using a mathematical model, we show that monogyny as a means of increasing paternity is favored when the sex ratio is male biased, but not necessarily by high search costs. The importance of a male-biased sex ratio for the evolution of monogyny is supported by various empirical studies.

Animals↗
PubMed2005

Selection for high gamete encounter rates explains the evolution of anisogamy using plausible assumptions about size relationships of swimming speed and duration.

A previous general model describing physical constraints on gamete encounter rates was modified to incorporate assumptions that increased size causes decreased swimming speed and increased fertile period (or other proportional enhancement to gamete fertility). The analysis indicates that with moderately strong size dependence of fertile period and a range of speed dependencies, selection for high encounter rates pressures mating systems that develop any heritable difference in size between the gametes of different mating types to exaggerate the difference and evolve from isogamy to anisogamy. The smaller gamete has an optimal size, but the larger faces continuing selection for increased size. This continues to a size that is estimated to be sufficient to make pheromone production of sperm attractants practical. This mechanism then bridges the missing link between isogametes and oogamy in a previous analysis of the effectiveness of pheromones in explaining the success of male-female mating systems. The evolution and success of anisogamy and oogamy can be explained solely on the basis of physical effects on the encounter process.

Animals↗
PubMed2007

One-male harems and female social dynamics in Guinea baboons.

Little is known about the mating system and social organization of Guinea baboons. This study investigated whether Guinea baboons have a harem-based mating system similar to that of hamadryas and gelada baboons and whether one-male mating units also correspond to social units. Ten adult females in a captive multi-male multi-female group of Guinea baboons were focally observed 2 h per week for 12 weeks, and all observed copulations within the group were recorded. Some males copulated with a single female while others had harems of 2-4 females. All females copulated with a single male except 1 female that switched harems early in the study. The focal females had higher rates of social interaction with their harem members, especially their harem male, than with individuals outside the harem. Females appeared to be subordinate to the harem male but little or no physical aggression or herding behavior from the male was observed. Variation in female social interactions within the harem was not accounted for by their sexual interactions with the male or their genetic relatedness with the females. Females, however, appeared to maintain social relationships with their female relatives in other harems. Taken together, the results of this study show that both mating and affiliative interactions in Guinea baboons are concentrated within one-male units and that the social dynamics within and between these units share some similarities as well as differences with those of hamadryas and gelada baboons.

Animals↗
PubMed2004

Reconstruction of parental microsatellite genotypes reveals female polyandry and philopatry in the lemon shark, Negaprion brevirostris.

Because sharks possess an unusual suite of reproductive characteristics, including internal fertilization, sperm storage, relatively low fecundity, and reproductive modes that range from oviparity to viviparity, they can provide important insight into the evolution of mating systems and sexual selection. Yet, to date, few studies have characterized behavioral and genetic mating systems in natural populations of sharks or other elasmobranchs. In this study, highly polymorphic microsatellite loci were used to examine breeding biology of a large coastal shark, the lemon shark, Negaprion brevirostris, at a tropical lagoon nursery. Over six years, 910 lemon sharks were sampled and genotyped. Young were assigned into sibling groups that were then used to reconstruct genotypes of unsampled adults. We assigned 707 of 735 young sharks to one of 45 female genotypes (96.2%), and 485 (66.0%) were assigned to a male genotype. Adult female sharks consistently returned to Bimini on a biennial cycle to give birth. Over 86% of litters had multiple sires. Such high levels of polyandry raise the possibility that polyandry evolved in viviparous sharks to reduce genetic incompatibilities between mother and embryos. We did not find a relationship between relatedness of mates and the number of offspring produced, indicating that inbreeding avoidance was probably not driving pre- or postcopulatory mate choice. Adult male sharks rarely sired more than one litter at Bimini and may mate over a broader geographic area.

Age Factors↗
PubMed2001

How cuckoldry can decrease the opportunity for sexual selection: data and theory from a genetic parentage analysis of the sand goby, Pomatoschistus minutus.

Alternative mating strategies are common in nature and are generally thought to increase the intensity of sexual selection. However, cuckoldry can theoretically decrease the opportunity for sexual selection, particularly in highly polygamous species. We address here the influence of sneaking (fertilization thievery) on the opportunity for sexual selection in the sand goby Pomatoschistus minutus, a marine fish species in which males build and defend nests. Our microsatellite-based analysis of the mating system in a natural sand goby population shows high rates of sneaking and multiple mating by males. Sneaker males had fertilized eggs in approximately 50% of the assayed nests, and multiple sneakers sometimes fertilized eggs from a single female. Successful males had received eggs from 2 to 6 females per nest (mean = 3.4). We developed a simple mathematical model showing that sneaking in this polygynous sand goby population almost certainly decreases the opportunity for sexual selection, an outcome that contrasts with the usual effects of cuckoldry in socially monogamous animals. These results highlight a more complex and interesting relationship between cuckoldry rates and the intensity of sexual selection than previously assumed in much of the literature on animal mating systems.

Animals↗
PubMed2001

Genetic consequences of polygyny and social structure in an Indian fruit bat, Cynopterus sphinx. II. Variance in male mating success and effective population size.

Variance in reproductive success is a primary determinant of genetically effective population size (Ne), and thus has important implications for the role of genetic drift in the evolutionary dynamics of animal taxa characterized by polygynous mating systems. Here we report the results of a study designed to test the hypothesis that polygynous mating results in significantly reduced Ne in an age-structured population. This hypothesis was tested in a natural population of a harem-forming fruit bat, Cynopterus sphinx (Chiroptera: Pteropodidae), in western India. The influence of the mating system on the ratio of variance Ne to adult census number (N) was assessed using a mathematical model designed for age-structured populations that incorporated demographic and genetic data. Male mating success was assessed by means of direct and indirect paternity analysis using 10-locus microsatellite genotypes of adults and progeny from two consecutive breeding periods (n = 431 individually marked bats). Combined results from both analyses were used to infer the effective number of male parents in each breeding period. The relative proportion of successfully reproducing males and the size distribution of paternal sibships comprising each offspring cohort revealed an extremely high within-season variance in male mating success (up to 9.2 times higher than Poisson expectation). The resultant estimate of Ne/N for the C. sphinx study population was 0.42. As a result of polygynous mating, the predicted rate of drift (1/2Ne per generation) was 17.6% higher than expected from a Poisson distribution of male mating success. However, the estimated Ne/N was well within the 0.25-0.75 range expected for age-structured populations under normal demographic conditions. The life-history schedule of C. sphinx is characterized by a disproportionately short sexual maturation period scaled to adult life span. Consequently, the influence of polygynous mating on Ne/N is mitigated by the extensive overlap of generations. In C. sphinx, turnover of breeding males between seasons ensures a broader sampling of the adult male gamete pool than expected from the variance in mating success within a single breeding period.

Animals↗
PubMed2000

The optimal number of fathers. Evolution, demography, and history in the shaping of female mate preferences.

Around the world polygynous marriage (one man, several women) is vastly more common than polyandrous marriage (one woman, several men), and women tend to be more cautious about entering into sexual relationships than men are. Such patterns are often assumed to reflect essential differences between the sexes. However, the same dichotomy between "ardent" males and "coy" females is not found in other primates. Furthermore, under a range of circumstances females enhance their reproductive success by mating with multiple partners and use polyandrous mating (soliciting copulations from several or more males) to circumvent male-imposed costs on their free choice of mates. The existence of one-male mating systems does not prove that females "naturally" gravitate to them. Typically monandrous (copulating with just one partner) mating systems are maintained by one male excluding rivals or by other circumstances that distort female options. As with many other animals, primate females (including women) can benefit reproductively from polyandrous matings. Understanding this takes us beyond narrow research programs intent on demonstrating "universal" differences between the sexes, and allows us to study females as flexible and opportunistic individuals who confront recurring reproductive dilemmas and tradeoffs within a world of shifting options.

Biological Evolution↗
PubMed2004

Reproductive success and effective population size in woodrats (Neotoma macrotis).

Discrepancies between the census size and the genetically effective size of populations (N(e)) can be caused by a number of behavioural and demographic factors operating within populations. Specifically, strong skew in male reproductive success, as would be expected in a polygynous mating system, could cause a substantial decrease in N(e) relative to census size. Because the mating system of Neotoma macrotis had previously been described as one nearing harem polygyny, I examined the distribution of reproductive success and genetic variation within a population of this species. Combining genetic data and three years of field observations, I show that variance in reproductive success does not deviate from poisson expectations within either sex and variance in success is similar between the sexes. Furthermore, both males and females had multiple partners across litters in addition to some evidence of multiple paternity within litters. Despite a lack of strong skew in reproductive success, an estimate of N(e) based on a number of demographic parameters suggests that the ratio of N(e)/N in this population is 0.48. Although the ratio of N(e)/N suggests that the population is experiencing higher rates of genetic drift than would be expected based on census size alone, the population maintains high levels of genetic diversity. Estimates of neighbourhood size and patterns of recruitment to the study site suggest that immigration plays an important role in this population and may contribute to the maintenance of high levels of genetic diversity.

Animals↗
PubMed1987

Genetic instability in the mating type system of Tetrahymena pigmentosa.

Selfing clones of Tetrahymena pigmentosa show several interesting genetic features, and provide some insight into the mechanisms of mating type (mt) determination. They differ significantly from those of Tetrahymena thermophila. They are distributed nonrandomly in crosses. Their rates of stabilization are highly variable, but most are much lower than those reported for T. thermophila. A number of subclones derived from nearly all the selfers have maintained stable mts in culture for several years. However, some subclones manifest persistent selfing, long after the calculated completion of allelic assortment for heterozygous loci. This phenomenon along with the perpetual maintenance of dominant mts in heterozygotes shows that phenotypic assortment is not involved in mt expression. In crosses, many selfers exhibit quantitative and qualitative aberrations in the transmission of alleles to the gametes; some of the micronuclear changes underlying these aberrations occur during vegetative growth. There are rare illegitimate appearances of dominant alleles in sexual progeny, and more common illegitimate appearances of the most recessive phenotype.--Various models to explain mt determination in this species are considered. One which might account for the troubling phenomena of the system consists of an active mat expression site, with "cassettes" at other sites specific for the different dominant alleles and capable of transposition to the expression site.

Animals↗
PubMed2006

Inbreeding depression and mixed mating in Leptosiphon jepsonii: a comparison of three populations.

BACKGROUND AND AIMS: Inbreeding depression is thought to play a central role in the evolution and maintenance of cross-fertilization. Theory indicates that inbreeding depression can be purged with self-fertilization, resulting in positive feedback for the selection of selfing. Variation among populations of Leptosiphon jepsonii in the timing and rate of self-fertilization provides an opportunity to study the evolution of inbreeding depression and mating systems. In addition, the hypothesis that differences in inbreeding depression for male and female fitness can stabilize mixed mating in L. jepsonii is tested. METHODS: In a growth room experiment, inbreeding depression was measured in three populations with mean outcrossing rates ranging from 0.06 to 0.69. The performance of selfed and outcrossed progeny is compared at five life history stages. To distinguish between self-incompatibility and early inbreeding depression, aborted seeds and unfertilized ovules were counted in selfed and outcrossed fruits. In one population, pollen and ovule production was quantified to estimate inbreeding depression for male and female fitness. KEY RESULTS: Both prezygotic barriers and inbreeding depression limited self seed set in the most outcrossing population. Cumulative inbreeding depression ranged from 0.297 to 0.501, with the lowest value found in the most selfing population. Significant inbreeding depression for early life stages was found only in the more outcrossing populations. Inbreeding depression was not significant for pollen or ovule production. CONCLUSIONS: The results provide modest support for the hypothesized relationship between inbreeding depression and mating systems. The absence of early inbreeding depression in the more selfing populations is consistent with theory on purging. Differences in male and female expression of inbreeding depression do not appear to stabilize mixed mating in L. jepsonii. The current estimates of inbreeding depression for L. jepsonii differ from those of previous studies, underscoring the effects of environmental variation on its expression.

Biomass↗
PubMed1994

Unpredictable food and sexual size dimorphism in insects.

The evolution of sexual size dimorphism is likely to be affected by the amount of resources each sex invests in offspring. Male nuptial gifts, occurring in many insect species, might reduce the value of large size in females and increase the value for males. For large nuptial gifts and an accompanying shift in dimorphism to evolve, males with larger gifts should be rewarded, in effect trading the larger gift for more offspring. We suggest that food variability, causing some males to have much to provide and some females to be in great need, would be conducive to the evolution of such a mating system, and we present comparative data on butterflies supporting the suggestion. In a gift-giving mating system, growing male and female juveniles should react differently to food shortage. A female maturing at small size can to some extent buffer her disadvantage through nuptial gifts, whereas a male maturing at small size will suffer from his inability to provide substantial gifts. Thus, males benefit more than females from continued growth in the face of food shortage, leading to a shift in size dimorphism. Here we confirm this prediction in a butterfly, Pieris napi, with large nuptial gifts.

Animals↗
PubMed1999

Determining the reproductive behaviour of individual brown-headed cowbirds using microsatellite DNA markers.

The reproductive behaviour of brood-parasitic birds has been of long-standing interest to evolutionary biologists but key features such as the mating system, patterns of host use and degree of overlap of female egg-laying ranges are largely unknown for particular species. Here we use microsatellite DNA markers to obtain such information for one of the best-studied brood parasites, the brown-headed cowbird, Molothrus ater. Parentage analyses of young and adults from a marked population at Delta, Manitoba for 1 year revealed the following key features of the reproductive behaviour of this species. (1) Monogamy is the predominant genetic mating system in this population, although matings by a single male with more than one female occur infrequently. (2) There is substantial variance in reproductive success among individuals, with only 23 out of 34 (68%) of all resident females and 21 out of 54 (39%) of all resident males in the study population identified as producing offspring. (3) Nonbreeding adults are seen less frequently and have shorter periods of residency on the study site suggesting they may form a subpopulation of 'floaters'. (4) Roughly half the females on site laid their eggs in the nests of a single host; most strikingly, the most fecund female laid all 13 of her eggs in red-winged blackbird, Agelaius phoeniceus, nests. This raises the possibility that cowbird populations may consist of combinations of females some of whom are host generalists and others host specialists. (5) Female egg-laying areas rarely overlap. (6) Realized annual fecundity of females may be lower than previously suggested. These results provide important information that can be used to clarify the mechanism and the selection pressures that have led to the evolution of different features of the reproductive behaviour of these birds. Copyright 1999 The Association for the Study of Animal Behaviour.

Journal Article↗
PubMed2000

Sociality in the gray mouse lemur (Microcebus murinus) in northwestern Madagascar.

The mating system and social organization of the gray mouse lemur (Microcebus murinus) was investigated in two three-month field studies (covering a period before, during, and after the first mating season) in Ampijoroa, northwestern Madagascar. The spatial and temporal distribution of the sexes within a population was studied using mark/recapture techniques and radiotelemetry to assess possible contest or scramble competition between the males. Sociality was inferred from the occurrence and probability of nocturnal social encounters, the temporal stability of daily sleeping groups, and nocturnal ranging patterns of co-sleepers. Males and females were evenly distributed in the study area within a network of highly overlapping home ranges. No indications were found for the spatial monopolization of the females by certain dominant males. Males and females had spatial access to several potential mates; the mating system is therefore characterized as a multi-male/multi-female system. Male home range sizes increased during the first mating season, which was interpreted as an indicator for scramble competition between the males. Competitive mate searching, sperm competition, and temporary mate guarding as well as female mate choice are suggested as the most probable reproductive strategies. Over the course of the study the animals lived continually within the study area, and most females formed stable individualized sleeping associations. Females that slept together shared a higher percentage of their home range than did females that slept at different sites. It is suggested that this network of social relationships should be described as a dispersed but individualized neighborhood.

Animals↗
PubMed2003

[Mating types in the ciliate Dileptus anser. Genetical instability in the mating type system].

In F1 and F2 from a cross between two clones of Dileptus anser isolated from natural sources (MT 1 x x MT III; MT = mating type), along with "normal" clones, many clones were observed demonstrating abnormal phenotype with respect to the MT-character. Irregular features of the latter were as follows: a) a delay in maturation; b) temporary reversion to immature or adolescent state, which means instability of maturity state; c) expression of MT I and MT III, rather than MT II as in properly matured clones; d) changes in MT (i.e., MT instability); e) appearance of totally unexpected MTs in terms of the scheme of genetic control of MTs in D. anser previously suggested by Afon'kin and Yudin (1987)--e.g., of all three MTs in F1 from the initial (analysing!) cross. Amazingly, these abnormal D. anser clones closely resembled some selfer-clones of Tetrahymena pigmentosa, previously reported as an example of genetic instability in the ciliate MT system (Simon, 1980; Simon, Orias, 1987).

Animals↗
PubMed2003

Directional changes in sexual size dimorphism in shorebirds, gulls and alcids.

The Charadrii (shorebirds, gulls and alcids) are one of the most diverse avian groups from the point of view of sexual size dimorphism, exhibiting extremes in both male-biased and female-biased dimorphism, as well as monomorphism. In this study we use phylogenetic comparative analyses to investigate how size dimorphism has changed over evolutionary time, distinguishing between changes that have occurred in females and in males. Independent contrasts analyses show that both body mass and wing length have been more variable in males than in females. Directional analyses show that male-biased dimorphism has increased after inferred transitions towards more polygynous mating systems. There have been analogous increases in female-biased dimorphism after transitions towards more socially polyandrous mating systems. Changes in dimorphism in both directions are attributable to male body size changing more than female body size. We suggest that this might be because females are under stronger natural selection constraints related to fecundity. Taken together, our results suggest that the observed variation in dimorphism of Charadrii can be best explained by male body size responding more sensitively to variable sexual selection than female body size.

Animals↗
PubMed2001

Surprising similarity of sneaking rates and genetic mating patterns in two populations of sand goby experiencing disparate sexual selection regimes.

Molecular markers have proved extremely useful in resolving mating patterns within individual populations of a number of species, but little is known about how genetic mating systems might vary geographically within a species. Here we use microsatellite markers to compare patterns of sneaked fertilization and mating success in two populations of sand goby (Pomatoschistus minutus) that differ dramatically with respect to nest-site density and the documented nature and intensity of sexual selection. At the Tvärminne site in the Baltic Sea, the microsatellite genotypes of 17 nest-tending males and mean samples of more than 50 progeny per nest indicated that approximately 35% of the nests contained eggs that had been fertilized by sneaker males. Successful nest holders mated with an average of 3.0 females, and the distribution of mate numbers for these males did not differ significantly from the Poisson expectation. These genetically deduced mating-system parameters in the Tvärminne population are remarkably similar to those in sand gobies at a distant site adjoining the North Sea. Thus, pronounced differences in the ecological setting and sexual selection regimes in these two populations have not translated into evident differences in cuckoldry rates or other monitored patterns of male mating success. In this case, the ecological setting appears not to be predictive of alternative male mating strategies, a finding of relevance to sexual selection theory.

Animals↗