Mating System Variation Research Articles

Postpollination discrimination between self and outcross pollen covaries with the mating system of a self-compatible flowering plant.

Variation in the mating system of hermaphroditic plant populations is determined by interactions between genetic and environmental factors operating via both pre- and postmating processes. Models predicting the maintenance of intermediate outcrossing rates in animal-pollinated plants often assume that the mating system is primarily controlled by floral morphology and pollinator availability, but rarely has the influence of postpollination processes on variation in outcrossing been examined. We investigated the influence of stylar discrimination between illegitimate and legitimate pollen-tube growth and the pollen-load capacity of stigmas on mating-system variation in the annual, tristylous species Eichhornia paniculata using controlled crosses and genetic markers. This species exhibits an exceptionally broad range of outcrossing rates in natural populations. There was significant variation among populations in the pollen-load capacity of stigmas and the ability of styles to discriminate between illegitimate vs. legitimate pollen. There was strong correspondence between stylar-discrimination ability and variation in outcrossing rate among populations and style morphs. The combination of stigmatic pollen-load capacity and stylar discrimination explained more than 80% of the variation in outcrossing rates among populations. The finding that stigmatic pollen-load capacity and stylar-discrimination ability contributed significantly to explaining the wide range of outcrossing rates in E. paniculata suggests that postpollination mechanisms play an important role in governing mating patterns in this species. The difference in levels of stylar discrimination between outcrossing and selfing populations may reflect a trade-off between selection for increased outcrossing and greater reproductive assurance.

Conflict and cooperation over sex: the consequences of social and genetic polyandry for reproductive success in dunnocks.

Conflict and cooperation within and between the sexes are among the driving forces that lead to the evolution of mating systems. Among mating strategies, female genetic polyandry and male reproductive cooperation pose challenging evolutionary questions regarding the maintenance of systems where one sex suffers from reduced fitness. Here, we investigate the consequences of social and genetic polyandry for reproductive success of females and males in a population of the dunnock, Prunella modularis. We show that female multiple mating ameliorates the negative effects of inbreeding. We, however, found little evidence that females engage in extra-group (pair) mating with less related or more heterozygous males. Breeding in socially polyandrous groups reduced the amount of paternity lost to extra-group males, such that, on average, cobreeding and monogamous males fledged a similar number of young. Importantly, c. 30% of cobreeding male dyads were related, suggesting they could gain indirect fitness benefits. Taken together, cobreeding males achieve equivalent reproductive success to monogamous counterparts under most circumstances. Our study has revealed unexpected complexities in the variable mating system of dunnocks in New Zealand. Our results differ from the well-known Cambridge dunnock study and can help our understanding of the evolution and maintenance of various breeding systems in the animal kingdom.

Transcriptome-enabled marker discovery and mapping of plastochron-related genes in Petunia spp.

BackgroundPetunia (Petunia × hybrida), derived from a hybrid between P. axillaris and P. integrifolia, is one of the most economically important bedding plant crops and Petunia spp. serve as model systems for investigating the mechanisms underlying diverse mating systems and pollination syndromes. In addition, we have previously described genetic variation and quantitative trait loci (QTL) related to petunia development rate and morphology, which represent important breeding targets for the floriculture industry to improve crop production and performance. Despite the importance of petunia as a crop, the floriculture industry has been slow to adopt marker assisted selection to facilitate breeding strategies and there remains a limited availability of sequences and molecular markers from the genus compared to other economically important members of the Solanaceae family such as tomato, potato and pepper.ResultsHere we report the de novo assembly, annotation and characterization of transcriptomes from P. axillaris, P. exserta and P. integrifolia. Each transcriptome assembly was derived from five tissue libraries (callus, 3-week old seedlings, shoot apices, flowers of mixed developmental stages, and trichomes). A total of 74,573, 54,913, and 104,739 assembled transcripts were recovered from P. axillaris, P. exserta and P. integrifolia, respectively and following removal of multiple isoforms, 32,994 P. axillaris, 30,225 P. exserta, and 33,540 P. integrifolia high quality representative transcripts were extracted for annotation and expression analysis. The transcriptome data was mined for single nucleotide polymorphisms (SNP) and simple sequence repeat (SSR) markers, yielding 89,007 high quality SNPs and 2949 SSRs, respectively. 15,701 SNPs were computationally converted into user-friendly cleaved amplified polymorphic sequence (CAPS) markers and a subset of SNP and CAPS markers were experimentally verified. CAPS markers developed from plastochron-related homologous transcripts from P. axillaris were mapped in an interspecific Petunia population and evaluated for co-localization with QTL for development rate.ConclusionsThe high quality of the three Petunia spp. transcriptomes coupled with the utility of the SNP data will serve as a resource for further exploration of genetic diversity within the genus and will facilitate efforts to develop genetic and physical maps to aid the identification of QTL associated with traits of interest.Electronic supplementary materialThe online version of this article (doi:10.1186/s12864-015-1931-4) contains supplementary material, which is available to authorized users.

The structure and organization of song in Southern House Wrens (Troglodytes aedon chilensis)

Studies of birdsong across very broad geographic scales, such as between the north temperate zone and the tropics, provide special opportunities to understand the role of variable ecologies, life histories and mating pressures on song structure and organization. The problem is typically studied through comparative, cross-species analyses because few species have such broad distributions to encompass both regions. The House Wren is an important exception, having the widest distribution of any native songbird in the Americas, from Canada to Tierra del Fuego. Across this range, they manifest considerable variation in life history, mating systems and migration, but there is no systematic research on corresponding song variation. Here we provide a first detailed characterization of song structure and organization for Southern House Wrens (Troglodytes aedon chilensis) in western Argentina and provide preliminary comparisons to Northern House Wrens. Songs of Southern House Wrens contained two distinct sections: an introduction of broadband noisy, or harmonic, notes followed by a louder terminal section of tonal, frequency-modulated syllables with a mean of seven syllables and three syllable types per song. The syllable repertoire was large (28), mostly shared and used to construct very large song repertoires (up to 170 song types with no evidence of a ceiling), but much smaller repertoires of commonly produced song types (24). Males tended to repeat song types many times before switching (eventual variety) but, at times, sang with immediate variety. Compared to Northern House Wrens, there were differences in the detailed form of some notes and syllables as well as in the relative emphasis of the softer introduction versus louder terminal section of songs. In broader patterns of song construction, organization, delivery, and the size of syllable and song repertoires, the two populations were very similar. These patterns are discussed in light of differences in life history, mating and migration between them.

Sexual reproduction with variable mating systems can resist asexuality in a rock-paper-scissors dynamics.

While sex can be advantageous for a lineage in the long term, we still lack an explanation for its maintenance with the twofold cost per generation. Here we model an infinite diploid population where two autosomal loci determine, respectively, the reproductive mode, sexual versus asexual and the mating system, polygynous (costly sex) versus monogamous (assuming equal contribution of parents to offspring, i.e. non-costly sex). We show that alleles for costly sex can spread when non-costly sexual modes buffer the interaction between asexual and costly sexual strategies, even without twofold benefit of recombination with respect to asexuality. The three interacting strategies have intransitive fitness relationships leading to a rock–paper–scissors dynamics, so that alleles for costly sex cannot be eliminated by asexuals in most situations throughout the parameter space. Our results indicate that sexual lineages with variable mating systems can resist the invasion of asexuals and allow for long-term effects to accumulate, thus providing a solution to the persisting theoretical question of why sex was not displaced by asexuality along evolution.

When sexual meets apomict: genome size, ploidy level and reproductive mode variation of Sorbus aria s.l. and S. austriaca (Rosaceae) in Bosnia and Herzegovina.

Allopolyploidy and intraspecific heteroploid crosses are associated, in certain groups, with changes in the mating system. The genus Sorbus represents an appropriate model to study the relationships between ploidy and reproductive mode variations. Diploid S. aria and tetraploid apomictic S. austriaca were screened for ploidy and mating system variations within pure and sympatric populations in order to gain insights into their putative causalities. Flow cytometry was used to assess genome size and ploidy level among 380 S. aria s.l. and S. austriaca individuals from Bosnia and Herzegovina, with 303 single-seed flow cytometric seed screenings being performed to identify their mating system. Pollen viability and seed set were also determined. Flow cytometry confirmed the presence of di-, tri- and tetraploid cytotype mixtures in mixed-ploidy populations of S. aria and S. austriaca. No ploidy variation was detected in single-species populations. Diploid S. aria mother plants always produced sexually originated seeds, whereas tetraploid S. austriaca as well as triploid S. aria were obligate apomicts. Tetraploid S. aria preserved sexuality in a low portion of plants. A tendency towards a balanced 2m : 1p parental genome contribution to the endosperm was shared by diploids and tetraploids, regardless of their sexual or asexual origin. In contrast, most triploids apparently tolerated endosperm imbalance. Coexistence of apomictic tetraploids and sexual diploids drives the production of novel polyploid cytotypes with predominantly apomictic reproductive modes. The data suggest that processes governing cytotype diversity and mating system variation in Sorbus from Bosnia and Herzegovina are probably parallel to those in other diversity hotspots of this genus. The results represent a solid contribution to knowledge of the reproduction of Sorbus and will inform future investigations of the molecular and genetic mechanisms involved in triggering and regulating cytotype diversity and alteration of reproductive modes.

Sexual antagonism in the pistil varies among populations of a hermaphroditic mixed-mating plant.

Sexual conflicts and their evolutionary outcomes may be influenced by population-specific features such as mating system and ecological context; however, very few studies have investigated the link between sexual conflict and mating system. The self-compatible, mixed-mating hermaphrodite Collinsia heterophylla (Plantaginaceae) is thought to exhibit a sexual conflict over timing of stigma receptivity. This conflict involves (i) delayed stigma receptivity, which intensifies pollen competition, and (ii) early fertilization forced by pollen, which reduces seed set. We investigated the potential for the conflict to occur under field conditions and performed glasshouse crosses within eight populations to assess its consistency across populations. Flowers were visited, and produced seeds after pollination, at all developmental stages, suggesting that the conflict can be of significance under natural conditions. In the glasshouse, early pollination imposed costs in all populations. Overall, the timing of first seed set was most strongly affected by the maternal parent, denoting stronger female than male ability to influence the onset of stigma receptivity. Crosses also revealed a negative relationship between donor- and recipient-related onset of receptivity within individuals, a novel result hinting at trade-offs in sex allocation or a history of antagonistic selection. Neither timing of stigma receptivity, timing of first seed set, nor pollen competitive ability covaried with population outcrossing rate. In conclusion, these results indicate that sexually antagonistic selection may be present in varying degrees in different populations of C. heterophylla, but this variation does not appear to be directly related to mating system variation.

Helping enhances productivity in campo flicker (Colaptes campestris) cooperative groups.

Reproductive adults in many bird species are assisted by non-breeding auxiliary helpers at the nest, yet the impact of auxiliaries on reproduction is variable and not always obvious. In this study, we tested Hamilton's rule and evaluated the effect of auxiliaries on productivity in the facultative cooperative breeder campo flicker (Colaptes campestris campestris). Campo flickers have a variable mating system, with some groups having auxiliaries and others lacking them (i.e., unassisted pairs). Most auxiliaries are closely related to the breeding pair (primary auxiliaries), but some auxiliaries (secondary auxiliaries) are unrelated females that joined established groups. We found no effect of breeder quality (body condition) or territory quality (food availability) on group productivity, but the presence of auxiliaries increased the number of fledglings produced relative to unassisted pairs. Nonetheless, the indirect benefit of helping was small and did not outweigh the costs of delayed breeding and so seemed insufficient to explain the evolution of cooperative breeding in campo flickers. We concluded that some ecological constraints must limit dispersal or independent breeding, making staying in the group a "best-of-a-bad-job" situation for auxiliaries.

Variation of reproduction in some species of the tribe Aveneae (Poaceae)

Patterns of reproduction were studied in a series of inter- and intraspecific units in the genus Avena. The range of variation of the mating system is from autogamy to allogamy. The first system is expressed in cultivated types, while the second is observed in wild and weedy ones. In a wild oat of putative hybrid origin or being a mutation, the development of pollen grains deviates from that noted in grasses as normal.

Mating system variation and assortative mating of sympatric bromeliads (Pitcairnia spp.) endemic to neotropical inselbergs.

The mating system is an important component of the complex set of reproductive isolation barriers causing plant speciation. However, empirical evidence showing that the mating system may promote reproductive isolation in co-occurring species is limited. The mechanisms by which the mating system can act as a reproductive isolation barrier are also largely unknown. Here we studied progeny arrays genotyped with microsatellites and patterns of stigma-anther separation (herkogamy) to understand the role of mating system shifts in promoting reproductive isolation between two hybridizing taxa with porous genomes, Pitcairnia albiflos and P. staminea (Bromeliaceae). In P. staminea, we detected increased selfing and reduced herkogamy in one sympatric relative to two allopatric populations, consistent with mating system shifts in sympatry acting to maintain the species integrity of P. staminea when in contact with P. albiflos. Mating system variation is a result of several factors acting simultaneously in these populations. We report mating system shifts as one possible reproductive barrier between these species, acting in addition to numerous other prezygotic (i.e., flower phenology and pollination syndromes) and postzygotic barriers (Bateson-Dobzhansky-Muller genetic incompatibilities).

Different mating expenditure in response to sperm competition risk between generations in the bivoltine butterfly Pieris napi

Examining how the response to sperm competition risk varies in a population is essential in order to understand variation in reproductive success and mating system. In polyandrous butterflies, male ...

Geographical Body Size Clines in Sika Deer: Path Analysis to Discern Amongst Environmental Influences

Using Japanese sika deer from boreal and subtropical forests, we investigated the effects of environmental factors on body size variation and tested the following ecological hypotheses pertaining to Bergmann’s rule in mammals: (1) heat conservation, (2) heat dissipation, (3) starvation resistance, (4) food availability and (5) insularity. Data on body sizes and habitats of sika deer from 31 populations on the Japanese archipelago were collected. Body mass and cranial greatest length (CGL; reflecting skeletal body size) were measured and analysed separately among gender groups. Path analyses were used to clarify inter-variable relationships and estimate direct and indirect effects of environmental variables on body size. Consistent with Bergmann’s rule, a clear latitudinal cline of body size was found for sika deer. Subsequent path analyses showed that the abiotic factors, specifically mean annual temperature and annual precipitation, had significant negative effects on body size, and annual temperature had the greatest effect among tested environmental variables. Winter severity and food availability during spring were significantly associated with body mass but not with CGL. Both heat conservation and dissipation hypotheses were accepted and food availability and starvation resistance hypotheses were applicable to variation of body mass but not to CGL, indicating that phenotypic changes in fat reserves strongly influence variation in body mass. Path diagram modelling of inter-variable relationships fit well for females but not for males, and unexplained variation of male body size suggested the presence of unidentified factors. Variation in mating systems among populations may effect body size variation of the male sika deer.

Limited pollen flow and high selfing rates toward geographic range limit in an Atlantic forest bromeliad

Bromeliaceae is a Neotropical family that evolved ecological key innovations in association with extensive adaptive radiation. Its species present a variety of different mating system strategies varying within and among species, within genera and subfamilies. Also, species with a wide geographical range can display large variation in mating system, reproductive success and genetic diversity. Here we combined data from hand pollinations and genetic analysis to assess outcomes of contemporary gene flow and mating system variation at the range edge of Vriesea gigantea. Results from pollen germination rates showed that this species is cryptically self-incompatible. Hand-pollination experiments and genetic analysis of progeny arrays revealed that V. gigantea has a mixed mating system, with high selfing rates (s=0.612), and high inbreeding coefficient (F=0.372). Inbreeding in V. gigantea at southern edge of its distribution range was caused by high levels of selfing rather than by mating among relatives. Moreover, strong pollen pool genetic structure was observed (Φ’FT=0.671), with an increase from north to south. The parameters observed help us to understand historical and ecological conditions under which V. gigantea has experienced moderate to high levels of selfing in the face of reduced pollen flow from central to peripheral populations due to recent southward range expansion.

Studying fish social behavior and cognition: implications for fish welfare and conservation

Event Abstract Back to Event Studying fish social behavior and cognition: implications for fish welfare and conservation Rui F. Oliveira1, 2* 1 Instituto Superior de Psicologia Aplicada, Portugal 2 Instituto Gulbenkian de Ciência, Portugal Within vertebrates teleost fish are the most diverse and plastic taxa in terms of social behavior. With over 29,000 species described so far, one can find all different types of social organization, mating systems and parental care types. Moreover, it is relatively common to find variation of these characters within closely related species, which makes them suitable for comparative studies on the evolution of social behavior (e.g. variation in mating systems and parental care type in African cichlids). Fish are also champions of social plasticity, as can be illustrated by the flexible patterns of sexual expression, as in the case of protrandrous and protogynous sex-change, simultaneous hermaphroditism and intra-sexual variation in the form of discrete alternative male phenotypes. Complex cognitive abilities used in social interactions have also evolved in fish, such as individual recognition, transitive inference and social learning. Therefore, teleosts offer unique opportunities to study both the evolution and the function of social behavior and cognition. In this talk I will summarize the work that our lab has been doing to establish zebrafish as a model organism for the study of social behavior and cognition and I will illustrate how knowledge on this are can be applied to fish welfare and to conservation issues. Keywords: Social Behaviour, Cognition, conservation, Animal Welfare, Teleost fishes Conference: XV European Congress of Ichthyology, Porto, Portugal, 7 Sep - 11 Sep, 2015. Presentation Type: Plenary Invited Presentation Topic: Physiology, Behavior and Toxicology Citation: Oliveira RF (2015). Studying fish social behavior and cognition: implications for fish welfare and conservation. Front. Mar. Sci. Conference Abstract: XV European Congress of Ichthyology. doi: 10.3389/conf.FMARS.2015.03.00226 Copyright: The abstracts in this collection have not been subject to any Frontiers peer review or checks, and are not endorsed by Frontiers. They are made available through the Frontiers publishing platform as a service to conference organizers and presenters. The copyright in the individual abstracts is owned by the author of each abstract or his/her employer unless otherwise stated. Each abstract, as well as the collection of abstracts, are published under a Creative Commons CC-BY 4.0 (attribution) licence (https://creativecommons.org/licenses/by/4.0/) and may thus be reproduced, translated, adapted and be the subject of derivative works provided the authors and Frontiers are attributed. For Frontiers’ terms and conditions please see https://www.frontiersin.org/legal/terms-and-conditions. Received: 16 Dec 2015; Published Online: 16 Dec 2015. * Correspondence: PhD. Rui F Oliveira, Instituto Superior de Psicologia Aplicada, Lisboa, Portugal, ruiol@ispa.pt Login Required This action requires you to be registered with Frontiers and logged in. To register or login click here. Abstract Info Abstract The Authors in Frontiers Rui F Oliveira Google Rui F Oliveira Google Scholar Rui F Oliveira PubMed Rui F Oliveira Related Article in Frontiers Google Scholar PubMed Abstract Close Back to top Javascript is disabled. Please enable Javascript in your browser settings in order to see all the content on this page.

Development and characterization of microsatellite markers for Vriesea simplex (Bromeliaceae) and cross-amplification in other species of Bromeliaceae

Ten polymorphic microsatellite loci were isolated and characterized for Vriesea simplex epiphytic and endemic species from the Brazilian Atlantic Rainforest. The number of alleles per locus ranged from 2 to 16. The observed and expected heterozygosity ranged from 0.000 to 0.414 and 0.068 to 0.795, respectively. All loci departed significantly from Hardy–Weinberg equilibrium. Linkage disequilibrium was not detected in any pair of loci. Transferability of 10 loci was positive across ten other Bromeliaceae species. The loci will be used for studying population genetic structure, mating system variation, and the origin and maintenance of reproductive barriers to gene exchange among sympatric Vriesea species.

Length polymorphism at the avpr1a locus is correlated with male reproductive behavior in a natural population of prairie voles (Microtus ochrogaster)

Laboratory studies have shown that vasopressin can influence sociosexual behavior through its action on the vasopressin 1a receptor (V1aR). There is substantial evidence that the length of a microsatellite in the gene (avpr1a) encoding for the V1aR can affect social attachment to females and paternal behavior in male prairie voles under laboratory conditions. However, previous field studies of prairie voles have failed to detect a strong effect of the length of a male’s avpr1a allele on their sociosexual behavior, but these studies are typically much shorter than the average prairie vole breeding lifespan. We examined the relationship between male avpr1a microsatellite allele length and sociosexual behavior in a natural population of prairie voles for 15 weeks, closer to the lifespan of prairie voles in nature. Contrary to predictions, we found that males with the longest avpr1a microsatellite alleles were significantly more likely to sire offspring with more than one female and to sire offspring that survived until trappable age than males with the shortest avpr1a microsatellite allele lengths. This relationship was the strongest for males with the longest tenure on the study site. As in previous field studies, we did not find evidence of a relationship between a male’s avpr1a genotype and any index of social behavior including male residency status or the number of females with which males associate. This is the first study to support the hypothesis that a male’s avpr1a genotype is a factor underlying variation in the genetic mating system of prairie voles under natural conditions.

Mating system and environmental variation drive patterns of adaptation in Boechera spatifolia (Brassicaceae).

Determining the relative contribution of population genetic processes to the distribution of natural variation is a major goal of evolutionary biology. Here, we take advantage of variation in mating system to test the hypothesis that local adaptation is constrained by asexual reproduction. We explored patterns of variation in ecological traits and genome-wide molecular markers in Boechera spatifolia (Brassicaceae), a species that contains both apomictic (asexual) and sexual individuals. Using a combination of quantitative genetics, neutral genetic (SSR) and genome-wide single nucleotide polymorphism, we assessed the hypothesis that asexual lineages should have reduced signatures of adaptation relative to sexual conspecifics. All three measures (traits, SSRs, SNPs) demonstrated that apomicts are genetically distinct from sexuals, regardless of population location. Additionally, phylogenetic clustering revealed that the apomictic group shared a single common ancestor. Across the landscape, sexual genome-wide SNP variation was strongly associated with latitude (r(2) >0.9), indicating that sexual populations have differentiated across an environmental gradient. Furthermore, flowering time and growth rate, as assessed in a common garden, strongly covary with the elevation and latitude of the source population. Despite a wide geographic distribution that largely overlaps with sexual populations, there was little evidence for differentiation in molecular markers or quantitative characters among apomictic populations. Combined, these data indicated that, in contrast to asexual populations, sexual populations show evidence of local adaptation.

Extensive long-distance pollen dispersal and highly outcrossed mating in historically small and disjunct populations of Acacia woodmaniorum (Fabaceae), a rare banded iron formation endemic.

Background and AimsUnderstanding patterns of pollen dispersal and variation in mating systems provides insights into the evolutionary potential of plant species and how historically rare species with small disjunct populations persist over long time frames. This study aims to quantify the role of pollen dispersal and the mating system in maintaining contemporary levels of connectivity and facilitating persistence of small populations of the historically rare Acacia woodmaniorum.MethodsProgeny arrays of A. woodmaniorum were genotyped with nine polymorphic microsatellite markers. A low number of fathers contributed to seed within single pods; therefore, sampling to remove bias of correlated paternity was implemented for further analysis. Pollen immigration and mating system parameters were then assessed in eight populations of varying size and degree of isolation.Key ResultsPollen immigration into small disjunct populations was extensive (mean minimum estimate 40 % and mean maximum estimate 57 % of progeny) and dispersal occurred over large distances (≤1870m). Pollen immigration resulted in large effective population sizes and was sufficient to ensure adaptive and inbreeding connectivity in small disjunct populations. High outcrossing (mean tm = 0·975) and a lack of apparent inbreeding suggested that a self-incompatibility mechanism is operating. Population parameters, including size and degree of geographic disjunction, were not useful predictors of pollen dispersal or components of the mating system.ConclusionsExtensive long-distance pollen dispersal and a highly outcrossed mating system are likely to play a key role in maintaining genetic diversity and limiting negative genetic effects of inbreeding and drift in small disjunct populations of A. woodmaniorum. It is proposed that maintenance of genetic connectivity through habitat and pollinator conservation will be a key factor in the persistence of this and other historically rare species with similar extensive long-distance pollen dispersal and highly outcrossed mating systems.

Extreme cost of rivalry in a monandrous species: male-male interactions result in failure to acquire mates and reduced longevity.

Mating system variation is profound in animals. In insects, female willingness to remate varies from mating with hundreds of males (extreme polyandry) to never remating (monandry). This variation in female behaviour is predicted to affect the pattern of selection on males, with intense pre-copulatory sexual selection under monandry compared to a mix of pre- and post-copulatory forces affecting fitness under polyandry. We tested the hypothesis that differences in female mating biology would be reflected in different costs of pre-copulatory competition between males. We observed that exposure to rival males early in life was highly costly for males of a monandrous species, but had lower costs in the polyandrous species. Males from the monandrous species housed with competitors showed reduced ability to obtain a mate and decreased longevity. These effects were specific to exposure to rivals compared with other types of social interactions (heterospecific male and mated female) and were either absent or weaker in males of the polyandrous species. We conclude that males in monandrous species suffer severe physiological costs from interactions with rivals and note the significance of male-male interactions as a source of stress in laboratory culture.

High but variable outcrossing rates in the invasive Geranium carolinianum (Geraniaceae).

• Premise of the study: Mating system plays an important role in population establishment and persistence, maintenance of genetic variation, and adaptive ability, especially for invasive species that colonize new environments to which they may be poorly adapted. In mixed-mating species, population differences in self-fertilization rates often arise due to variation in local ecological conditions (e.g., pollinator or mate availability) or genetic variation in traits promoting selfing or outcrossing. Knowledge of how and why selfing rates vary can help us understand how populations respond to different environments, how this affects patterns of genetic variation, and the role of mating systems in biological invasions.• Methods: We determined outcrossing rates in invasive (China) and native (US) populations of the weedy annual Geranium carolinianum in natural populations and an open-pollinated common garden to answer the following questions: To what extent do populations vary in mating system? Do invasive populations differ from native populations? Is interpopulation variation in mating system under genetic control?• Key results: Despite having many characteristics of selfing species, we found high variation in outcrossing rates (tm from 0.1 to 1.0) resulting from both environmental and genetic variation. Outcrossing rates were generally high and plastic in the Chinese populations.• Conclusions: A delayed selfing mechanism provides reproductive assurance while allowing facultative outcrossing when mates are not limiting. In invasive populations, high outcrossing rates were facilitated by large admixed founding populations, promoting local adaptation and the maintenance of genetic diversity.