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Phylogenetic analysis of the bluestain fungus Ophiostoma minus based on partial ITS rDNA and beta-tubulin gene sequences.

In an attempt to clarify the relationship between fungi classified as Ophiostoma minus, but of different geographic origins and mating systems, sequencing of the 5.8S and ITS 2 rDNA, and beta-tubulin gene was carried out. The beta-tubulin gene was highly informative, supporting the sub-division of O. minus into two groups based on geographic origin. Furthermore, isolates previously classified as O. pseudotsugae were confirmed as being clearly distinct from O. minus. However, sequencing did not reveal any polymorphisms between isolates with homothallic as compared to heterothallic mating systems. This was supported by crosses using methylbenzamidazole-2-yl carbamate nuclear markers which showed that hybridisation between isolates of different mating systems was possible. However, we propose that different mating systems may still signal a divergence of isolates of O. minus.

Ascomycota↗

Anopheles culicifacies: effects of age on the male reproductive system and mating ability of virgin adult mosquitoes.

Under controlled laboratory conditions of 28-30 degrees C and 16:8 L:D photoperiod, an attempt was made to develop an age-grading technique for Anopheles culicifacies males. Mating activity was maximal when females were 5-12 days old males were 5-7 days old. The numbers of total and mature spermatocysts declined significantly with age, and the proportion of the testes occupied by the sperm reservoir increased as virgin males grew older. Mating resulted in the loss of spermatozoa and accessory gland substance from the reproductive system. Loss of mating ability of older virgin males seemed to be age-related, because the reproductive system contained ample supplies of accessory gland substance and spermatozoa. Morphological changes of the reproductive system, due to mating and age, were used to infer the age and reproductive history of unknown males in a laboratory evaluation.

Aging↗

Is self-fertilization an evolutionary dead end? Revisiting an old hypothesis with genetic theories and a macroevolutionary approach.

G. Ledyard Stebbins suggested that self-fertilization (selfing) may be an evolutionary dead end because it may result in the loss of genetic diversity and consequently preclude adaptation to changing environments. While the basic premise of selfing as a dead end is widely accepted, there have been few rigorous evaluations of the hypothesis. We examine the foundations of the dead-end hypothesis by considering theoretical advances in the study of mating-system evolution. We discuss theories predicting the irreversibility of self-fertilization and the extinction of selfing lineages through the loss of adaptive potential and genetic degradation. In the second portion of the review, focusing on the irreversibility of selfing, we summarize the contribution of phylogenetic studies of mating-system evolution to determine if evolutionary history supports this well-established hypothesis. Most studies are in accord with the hypothesis; no single study unequivocally demonstrates the transition from highly selfing to outcrossing lineages. Finally, we discuss the problems encountered when phylogenetic studies rely on reconstruction of ancestral mating systems. To avoid some of these problems, we applied likelihood ratio tests of irreversibility of mating-system evolution to several data sets and found that current data sets are probably too small for this test.

Journal Article↗

Ecological correlates of extra-group paternity in mammals.

Extra-group paternity (EGP) can form an important part of the mating system in birds and mammals. However, our present understanding of its extent and ecology comes primarily from birds. Here, we use data from 26 species and phylogenetic comparative methods to explore interspecific variation in EGP in mammals and test prominent ecological hypotheses for this variation. We found extensive EGP (46% of species showed more than 20% EGP), indicating that EGP is likely to play an important role in the mating system and the dynamics of sexual selection in mammals. Variation in EGP was most closely correlated with the length of the mating season. As the length of the mating season increased, EGP declined, suggesting that it is increasingly difficult for males to monopolize their social mates when mating seasons are short and overlap among females in oestrus is likely to be high. EGP was secondarily correlated with the number of females in a breeding group, consistent with the idea that as female clustering increases, males are less able to monopolize individual females. Finally, EGP was not related to social mating system, suggesting that the opportunities for the extra-group fertilizations and the payoffs involved do not consistently vary with social mating system.

Animals↗

Quantitative measure of sexual selection with respect to the operational sex ratio: a comparison of selection indices.

Despite numerous indices proposed to predict the evolution of mating systems, a unified measure of sexual selection has remained elusive. Three previous studies have compared indices of sexual selection under laboratory conditions. Here, we use a genetic study to compare the most widely used measures of sexual selection in natural populations. We explored the mating and reproductive successes of male and female bank voles, Clethrionomys glareolus, across manipulated operational sex ratios (OSRs) by genotyping all adult and pup bank voles on 13 islands using six microsatellite loci. We used Bateman's principles (Is and I and Bateman gradients) and selection coefficients (s' and beta') to evaluate, for the first time, the genetic mating system of bank voles and compared these measures with alternative indices of sexual selection (index of monopolization and Morisita's index) across the OSRs. We found that all the sexual selection indices show significant positive intercorrelations for both males and females, suggesting that Bateman's principles are an accurate and a valid measure of the mating system. The Bateman gradient, in particular, provides information over and above that of other sexual selection indices. Male bank voles show a greater potential for sexual selection than females, and Bateman gradients indicate a polygynandrous mating system. Selection coefficients reveal strong selection gradients on male bank vole plasma testosterone level rather than body size.

Animals↗

Predominance of genetic monogamy by females in a hammerhead shark, Sphyrna tiburo: implications for shark conservation.

There is growing interest in the mating systems of sharks and their relatives (Class Chondrichthyes) because these ancient fishes occupy a key position in vertebrate phylogeny and are increasingly in need of conservation due to widespread overexploitation. Based on precious few genetic and field observational studies, current speculation is that polyandrous mating strategies and multiple paternity may be common in sharks as they are in most other vertebrates. Here, we test this hypothesis by examining the genetic mating system of the bonnethead shark, Sphyrna tiburo, using microsatellite DNA profiling of 22 litters (22 mothers, 188 embryos genotyped at four polymorphic loci) obtained from multiple locations along the west coast of Florida. Contrary to expectations based on the ability of female S. tiburo to store sperm, the social nature of this species and the 100% multiple paternity observed in two other coastal shark species, over 81% of sampled bonnethead females produced litters sired by a single male (i.e. genetic monogamy). When multiple paternity occurred in S. tiburo, there was an indication of increased incidence in larger mothers with bigger litters. Our data suggest that sharks may exhibit complex genetic mating systems with a high degree of interspecific variability, and as a result some species may be more susceptible to loss of genetic variation in the face of escalating fishing pressure. Based on these findings, we suggest that knowledge of elasmobranch mating systems should be an important component of conservation and management programmes for these heavily exploited species.

Animals↗

Regular systems of inbreeding.

Regular systems of inbreeding with discrete, nonoverlapping generations and the same number of individuals and mating pattern in every generation are studied. The matrix Q that specifies the recursion relations satisfied by the probabilities of identity is expressed in terms of the matrix M that describes the mating system. Necessary and sufficient conditions for convergence to genetic uniformity are given, and it is determined which probabilities of identity approach one. If the mating system has certain symmetries and these are imposed initially, then a matrix R, of lower dimension than Q, specifies the recursion relations. For such a mating system, for generic initial conditions, the condensed matrix R suffices for determining whether convergence to uniformity occurs and which probabilities of identity approach one. If Q is irreducible, the maximal eigenvalue of R is the same as that of Q. If Q is also aperiodic, this implies that the asymptotic rate of convergence to homogeneity of the condensed system is the same as that of the complete one. The above results apply to autosomal loci in monoecious (with or without selfing) and dioecious populations and to X-linked loci. As an example, all the eigenvalues and right and left eigenvectors of Q for circular mating are found.

Animals↗

Experimental dissection of inbreeding and its adaptive significance in a flowering plant, Aquilegia canadensis (Ranunculaceae).

Inbreeding is a major component of the mating system in populations of many plants and animals, particularly hermaphroditic species. In flowering plants, inbreeding can occur through self-pollination within flowers (autogamy), self-pollination between flowers on the same plant (geitonogamy), or cross-pollination between closely related individuals (biparental inbreeding). We performed a floral emasculation experiment in 10 populations of Aquilegia canadensis (Ranunculaceae) and used allozyme markers to estimate the relative contribution of each mode of inbreeding to the mating system. We also examined how these modes of inbreeding were influenced by aspects of population structure and floral morphology and display predicted to affect the mating system. All populations engaged in substantial inbreeding. On average, only 25% of seed was produced by outcrossing (range among populations = 9-37%), which correlated positively with both population size (r = +0.61) and density (r = +0.64). Inbreeding occurred through autogamy and biparental inbreeding, and the relative contribution of each was highly variable among populations. Estimates of geitonogamy were not significantly greater than zero in any population. We detected substantial biparental inbreeding (mean = 14% of seeds, range = 4-24%) by estimating apparent selfing in emasculated plants with no opportunity for true selfing. This mode of inbreeding correlated negatively with population size (r = -0.87) and positively with canopy cover (r = +0.90), suggesting that population characteristics that increase outcross pollen transfer reduce biparental inbreeding. Autogamy was the largest component of the mating system in all populations (mean = 58%, range = 37-84%) and, as expected, was lowest in populations with the most herkogamous flowers (r = -0.59). Although autogamy provides reproductive assurance in natural populations of A. canadensis, it discounts ovules from making superior outcrossed seed. Hence, high autogamy in these populations seems disadvantageous, and therefore it is difficult to explain the extensive variation in herkogamy observed both among and especially within populations.

Adaptation, Biological↗

How are deleterious mutations purged? Drift versus nonrandom mating.

Accumulation of deleterious mutations has important consequences for the evolution of mating systems and the persistence of small populations. It is well established that consanguineous mating can purge a part of the mutation load and that lethal mutations can also be purged in small populations. However, the efficiency of purging in natural populations, due to either consanguineous mating or to reduced population size, has been questioned. Consequences of consanguineous mating systems and small population size are often equated under "inbreeding" because both increase homozygosity, and selection is though to be more efficient against homozygous deleterious alleles. I show that two processes of purging that I call "purging by drift" and "purging by nonrandom mating" have to be distinguished. Conditions under which the two ways of purging are effective are derived. Nonrandom mating can purge deleterious mutations regardless of their dominance level, whereas only highly recessive mutations can be purged by drift. Both types of purging are limited by population size, and sharp thresholds separate domains where purging is either effective or not. The limitations derived here on the efficiency of purging are compatible with some experimental studies. Implications of these results for conservation and evolution of mating systems are discussed.

Biological Evolution↗

Geographic structure of pollinator communities, reproductive assurance, and the evolution of self-pollination.

Reproductive assurance is often invoked as an explanation for the evolution of self-fertilization in plants. However, key aspects of this hypothesis have received little empirical support. In this study, I use geographic surveys of pollinator communities along with functional studies of floral trait variation to examine the role of pollination ecology in mating system differentiation among populations and subspecies of the annual plant Clarkia xantiana. A greenhouse experiment involving 30 populations from throughout the species' range indicated that variation in two floral traits, herkogamy and protandry, was closely related to levels of autofertility and that trait variation was partitioned mainly among populations. Emasculation experiments in the field showed that autonomous selfing confers reproductive assurance by elevating fruit and seed production. Surveys of pollinator communities across the geographic range of the species revealed that bee pollinator abundance and community composition differed dramatically between populations of the outcrossing subspecies xantiana and the selfing subspecies parviflora despite their close proximity. Specialist bee pollinators of Clarkia were absent from selfing populations, but they were the most frequent visitors to outcrossing populations. Moreover, within the outcrossing subspecies xantiana, there was a close correspondence between specialist abundance and population differentiation in herkogamy, a key mating system trait. This spatial covariation arose, in part, because geographically peripheral populations had reduced herkogamy, higher autofertility, and lower pollinator abundance compared to central populations of xantiana. Finally, I detected strong spatial structure to bee communities both across the range of the species and within the outcrossing subspecies. In both cases, spatial structure was stronger for specialist bees compared to generalist bees, and pollinator communities varied in parallel with population variation in herkogamy. These results provide evidence that mating system differentiation parallels spatial variation in pollinator abundance and community composition at both broad and more restricted spatial scales, consistent with the hypothesis that pollinator abundance and reproductive assurance are important drivers of plant mating system evolution.

Adaptation, Biological↗

Multiple paternity in a natural population of a salamander with long-term sperm storage.

Sperm competition appears to be an important aspect of any mating system in which individual female organisms mate with multiple males and store sperm. Post-copulatory sexual selection may be particularly important in species that store sperm throughout long breeding seasons, because the lengthy storage period may permit extensive interactions among rival sperm. Few studies have addressed the potential for sperm competition in species exhibiting prolonged sperm storage. We used microsatellite markers to examine offspring paternity in field-collected clutches of the Ocoee salamander (Desmognathus ocoee), a species in which female organisms store sperm for up to 9 months prior to fertilization. We found that 96% of clutches were sired by multiple males, but that the majority of females used sperm from only two or three males to fertilize their eggs. The high rate of multiple mating by females suggests that sperm competition is an important aspect of this mating system. Comparison of our data with those of other parentage studies in salamanders and newts reveals that multiple mating may be common in urodele amphibians. Nevertheless, the number of males siring offspring per clutch in D. ocoee did not differ appreciably from that in other species of urodeles with shorter storage periods.

Animals↗

Sexual size dimorphism in shorebirds, gulls, and alcids: the influence of sexual and natural selection.

Charadrii (shorebirds, gulls, and alcids) have an unusual diversity in their sexual size dimorphism, ranging from monomorphism to either male-biased or female-biased dimorphism. We use comparative analyses to investigate whether this variation relates to sexual selection through competition for mates or natural selection through different use of resources by males and females. As predicted by sexual selection theory, we found that in taxa with socially polygynous mating systems, males were relatively larger than females compared with less polygynous species. Furthermore, evolution toward socially polyandrous mating systems was correlated with decreases in relative male size. These patterns depend on the kinds of courtship displays performed by males. In taxa with acrobatic flight displays, males are relatively smaller than in taxa in which courtship involves simple flights or displays from the ground. This result remains significant when the relationship with mating system is controlled statistically, thereby explaining the enigma of why males are often smaller than females in socially monogamous species. We did not find evidence that evolutionary changes in sexual dimorphism relate to niche division on the breeding grounds. In particular, biparental species did not have greater dimorphism in bill lengths than uniparental species, contrary to the hypothesis that selection for ecological divergence on the breeding grounds has been important as a general explanation for patterns of bill dimorphism. Taken together, these results strongly suggest that sexual selection has had a major influence on sexual size dimorphism in Charadrii, whereas divergence in the use of feeding resources while breeding was not supported by our analyses.

Animals↗

The Hermaphrodite's Dilemma.

Given sexual conflict, mating encounters between simultaneous hermaphrodites will conform to a new, conditional, non-zero sum game of strategy, the Hermaphrodite's Dilemma; special cases of which include Prisoner's Dilemma and Game of Chicken. The model predicts that hermaphrodite mating systems will be based on reciprocation with cheating in a preferred sexual role. This model suggests that study of hermaphrodite mating systems will provide direct evidence for the existence of sexual conflict and suggests that sexual conflict may act to stabilize hermaphroditism through such mating systems.

Animals↗

Prediction of parental dominance combinations for planned matings, methodology, and simulation results.

Optimal use of dominance information requires a mating system and predictions of specific combining abilities for each set of prospective parents. Current evaluation procedures provide such predictions only for a limited number of parents. A procedure is described that predicts the specific combining ability for any parents. In this procedure, for each set of parents and their ancestors, the additive relationship matrix is created as a dense matrix. This matrix is then used to create a parental dominance matrix in a sparse matrix form, in which the rows of the matrix correspond to all parental combinations for which predictions are already available. Each new prediction requires a solution of the system of equations with the parental dominance matrix as the left-hand side. The efficiency of the mating system that accounts for dominance was evaluated in a simulation study. The simulated data files varied with respect to proportion of males and females selected, proportion of cattle born through embryo transfer, and additive and dominance variance. Sires and dams were preselected based on the additive merits only, but specific matings were arranged based on the combined additive plus dominance merit. The response to selection with consideration of dominance increased from 3.8 to 16.6% of the response from one generation of additive selection. The response was greater when the additive variance was smaller, the dominance variance was larger, the intensity of additive selection was lower, and the proportion of full sibs was greater. Use of dominance in the mating system is feasible and results in an additional genetic response to selection.

Animals↗

Inferring social behavior from sexual dimorphism in the fossil record.

Sexual dimorphism is commonly used as evidence of the behavior of extinct species. Even so, few analyses scrutinize whether extant comparative data support inferences of mating systems or behavior in extinct species. This analysis evaluates the relations between measures of dimorphism and several estimates of mating system and intrasexual competition. Dimorphism alone provides poor resolution for reconstructing behavior. Many behavioral inferences based on perceived dimorphism are not supported by extant comparative data. This reflects the large standard errors of relations between dimorphism estimates and behavioral classifications. Used with caution, dimorphism can provide a hint of the behavior of extinct species in some cases. However, in many cases inferred dimorphism allows little more than an inference of polygyny, without any indication of specific types of mating systems.

Animals↗

Number of lethal equivalents in human populations: how good are the previous estimates?

Lee et al. (1996) recently developed a method for interval estimation of the number of lethal equivalents by using a hierarchical structure of likelihood functions. This hierarchical model consists of two multinomial trials: one of the sampling process of the parents from the population of interest, and the other for the survival of the offspring of the families obtained by mating the parents. The method, initially developed for selfing and full-sib mating, is extended here to include more general mating systems as well as mixtures of mating systems. We applied it to human data sets for which confidence intervals were previously not available. Our point estimates were close to previous ones, and the standard deviations were generally quite small. Thus, even if debate over the meaning of the concept of lethal equivalents has not been entirely resolved, our results showed that the previous estimates are at least statistically meaningful.

Alleles↗

Quantifying genetic and genotypic gain gaps in Eucalyptus: the hidden cost of ignoring inbreeding and dominance.

Understanding the mating system of Eucalyptus species is necessary to accurately estimate genetic parameters and improve breeding programs. Eucalyptus species often exhibit mixed mating systems, leading to complex relationships among progenies. Traditional tree breeding programs that assume a half-sibling relationship for open-pollinated (OP) trials may overestimate genetic gains by neglecting the effects of inbreeding and dominance. This study focuses on Eucalyptus pellita, a species with a mixed mating system, to quantify the impact of selfing and dominance on breeding strategies. We simulated OP trial growth data for 100 randomly selected families in a randomized complete block design, using published estimated parameters for diameter at breast height (DBH). Our analysis indicated that marker-based models, particularly those incorporating dominance effects, provide more accurate genetic parameter estimates and larger predicted genetic gains than pedigree-based models. These results reveal pronounced genetic and genotypic gain gaps when traditional models are employed, underscoring the imperative for integrating dominance-informed genomic selection strategies. Thus, our study provides essential guidance for optimizing breeding programs to sustainably enhance productivity and genetic quality in Eucalyptus plantations.

Eucalyptus↗

Male mating patterns in wild multimale mountain gorilla groups.

Although mountain gorillas, Gorilla gorilla beringei, are classified as having a one-male mating system, approximately 40% of the social units are multimale groups. I observed two multimale groups of mountain gorillas at the Karisoke Research Center, Rwanda, Africa, for 17 months to determine male mating patterns and male-male mating harassment in relation to both male dominance rank and female reproductive status. Dominant males mated significantly more than did individual subordinate males, and dominant males mated more with cycling adult and pregnant females. The dominant males participated in 47 and 83% of observed matings in the two groups. Subordinate males were more likely than dominant males to mate with subadult females. Eleven of 14 females were observed to mate with more than one male, and multiple males mated with three of the five females observed at the probable time of conception. Mating harassment was initiated and received by both dominant and subordinate males. Mating harassment occurred infrequently (during 30 and 22% of matings in each group), usually consisted of mild aggression, and usually terminated copulations by subordinate males, but not those by dominant males. These results suggest that multimale mountain gorilla groups can be favourable environments for subordinate males to obtain mating opportunities. Dominant males may be unable or unwilling to prevent subordinate males from mating. Based on behavioural observations, mountain gorillas can have a multimale mating system but further research on the role of females in male mating success and paternity determination is needed to understand fully this species' mating system. Copyright 1999 The Association for the Study of Animal Behaviour.

Journal Article↗