Responses to selection on male-phase duration in Chamerion angustifolium

Protandry (when male function precedes female) can enhance fitness by reducing selfing and increasing pollen export and outcrossed siring success. However, responses to selection on protandry may be constrained by genetic variation and correlations among floral traits. We examined these potential constraints in protandrous Chamerion angustifolium (Onagraceae) by estimating genetic variation in male-phase duration and associated floral traits using a paternal half-sib design and selection experiment. Narrow-sense heritability of male-phase duration was estimated as 0.23 (SE +/- 0.04) and was positively correlated with floral display. The selection experiment shortened male-phase duration 0.8 SD from the parental average of 17.0 h and lengthened it by 2.0 SD. Furthermore, fixed floral longevity caused a negative association between male- and female-phase durations. These results suggest that selection on male-phase duration is not limited by genetic variation. However, changes in male-phase duration may influence pollinators through correlated changes in floral display and reduced opportunities for pollen receipt during female phase.     

The effect of protandry on siring success in Chamerion angustifolium (Onagraceae) with different inflorescence sizes

Abstract Protandry, a form of temporal separation of gender within hermaphroditic flowers, may reduce the magnitude of pollen lost to selfing (pollen discounting) and also serve to enhance pollen export and outcross siring success. Because pollen discounting is strongest when selfing occurs between flowers on the same plant, the advantage of protandry may be greatest in plants with large floral displays. We tested this hypothesis with enclosed, artificial populations of Chamerion angustifolium (Onagraceae) by experimentally manipulating protandry (producing uniformly adichogamous or mixed protandrous and adichogamous populations) and inflorescence size (two-, six-, or 10-flowered inflorescences) and measuring pollinator visitation, seed set, female outcrossing rate, and outcross siring success. Bees spent more time foraging on and visited more flowers of larger inflorescences than small. Female outcrossing rates did not vary among inflorescence size treatments. However, seed set per fruit decreased with increasing inflorescence size, likely as a result of increased abortion of selfed embryos, perhaps obscuring the magnitude of geitonogamous selfing. Protandrous plants had a marginally higher female outcrossing rate than adichogamous plants, but similar seed set. More importantly, protandrous plants had, on average, a twofold siring advantage relative to adichogamous plants. However, this siring advantage did not increase linearly with inflorescence size, suggesting that protandry acts to enhance siring success, but not exclusively by reducing between-flower interference.     

非人灵长类性打搅行为研究进展

性打搅行为是指非人灵长类社群内非交配个体对正在发生性行为个体的干扰行为,该行为在非人灵长类中普遍存在,通过综述目前已知的36种非灵长类的性打搅行为的报道,探讨性打搅行为的功能和意义。性打搅行为具有更高度的种间和种内差异,种间的性打搅的方式不同、同时受到社会结构和婚配制度的影响;在种内受到性别、年龄、等级等因素的影响。结合川金丝猴性打搅的研究结果,从性打搅的类型、被打搅的反应、性打搅的功能以及相关假说等方面入手,对非人灵长类性打搅行为进行系统总结,旨在为我国相关领域的研究提供参考,为生殖行为学研究者启发思路,推动本领域的发展。     

Inferences on mating and sexual systems of two Pacific Cinetorhynchus shrimps (Decapoda,Rhynchocinetidae) based on sexual dimorphism in body size and cheliped weaponry

Sexual dimorphism in body size and weaponry was examined in two Cinetorhynchus shrimp species in order to formulate hypotheses on their sexual and mating systems. Collections of Cinetorhynchus sp. A and Cinetorhynchus sp. B were made in March, 2011 on Coconut Island, Hawaii, by hand dipnetting and minnow traps in coral rubble bottom in shallow water. Although there is overlap in male and female size, some males are much larger than females. The major (pereopod 1) chelipeds of males are significantly larger and longer than those of females. In these two Cinetorhynchus species, males and females have third maxillipeds of similar relative size, i.e., those of males are not hypertrophied and probably not used as spear-like weapons as in some other rhynchocinetid (Rhynchocinetes) species. Major chelae of males vary with size, changing from typical female-like chelae tipped with black corneous stout setae to subchelate or prehensile appendages in larger males. Puncture wounds or regenerating major chelipeds were observed in 26.1 % of males examined (N = 38 including both species). We interpret this evidence on sexual dimorphism as an indication of a temporary male mate guarding or “neighborhoods of dominance” mating system, in which larger dominant robustus males defend females and have greater mating success than smaller males. Fecundity of females increased with female size, as in most caridean species (500–800 in Cinetorhynchus sp. A; 300–3800 in Cinetorhynchus sp. B). Based on the sample examined, we conclude that these two species have a gonochoric sexual system (separate sexes) like most but not all other rhynchocinetid species in which the sexual system has been investigated.     

Sexual segregation occurs in bats within fragmented remnant woodlands in an agricultural landscape

Species‐specific responses to landscape configuration and landscape composition have been studied extensively. However, little work has been done to compare intraspecific differences in habitat preferences. Bats have potential as good bioindicator taxa in woodland habitats. Therefore, studying sex differences in responses to woodland and the wider landscape can allow us to gain insight into the relative importance of these habitats for both bats and other taxa. In this study, we aimed to test the predictions that (i) habitat type and connectivity will influence the probability of recording female bats in woodlands and (ii) sex differences in response to habitat type and connectivity will be species‐specific. Bat capture data was collected in 206 woodlands over 3 years in England. The probability of detecting females relative to males was modeled in response to a range of woodland characteristics and landscape metrics for six bat species. We recorded sex differences in responses to landscape features in three species. We found a higher probability of capturing female Myotis nattereri in woodlands that were surrounded by a higher proportion of improved grasslands, whereas female Myotis mystacinus were less likely to be recorded in woodlands surrounded by semi‐natural vegetation. Female Plecotus auritus were more likely to be recorded in isolated woodlands with less connectivity to other woodlands and where agriculture dominated the surrounding landscape. Our findings indicate that sexual segregation occurs across several UK bat species in response to landscape connectivity and composition. Sexual segregation in response to landscape characteristics in bats should therefore be an important consideration in the management of fragmented agricultural landscapes.     

Sexual interference in two Chamerion species with contrasting modes of movement herkogamy

Evolution of herkogamy has been ascribed to preventing autonomous selfing within flowers, but this argument has received challenges due to the negative effects of sexual interference on reproductive success of plant species. We examined the effects of the two contrasting modes of herkogamy, detailed by the different patterns of style movement, on the foraging time of pollinators on flowers, pollen removal, and deposition in two Chamerion species. Our results suggested that the duration of foraging time of bumblebees was generally higher in C. angustifolium (L.) Holub than in C. conspersum (Hausskn.) Holub during the male-phase of flowers, but the reverse during the female-phase of flowers. As a result, stigma interfered with pollen removal more in C. conspersum than in C. angustifolium, based on examinations of pollen removal after one visit by a pollinator. However, this negative effect on total pollen removal was covered by the high pollinator diversity and the possible high visitation rate in the study sites. In contrast, we found that almost no difference existed in pollen deposition rate after one visit of a pollinator between the two Chamerion species, and emasculation did not increase the total pollen deposition in either species, indicating that anthers did not interfere with pollen deposition in the two Chamerion species. Our results suggested that, compared with C. conspersum, the derived mode of herkogamy in C. angustifolium reduced interference of stigmas in pollen removal. This might contribute to the wide distribution of C. angustifolium, to some degree, in environments of pollinator scarcity and low activity in newly colonized habitats.     

木棉黄花个体的适应意义

木棉(Bombax ceiba)花大、红色, 多为鸟媒传粉, 但也存在频率极低的黄花个体, 有蜜蜂访花。本文对比研究了海南岛木棉红花、黄花的花部特征及传粉过程, 以解释黄花个体的适应意义。木棉两种花色表型的花蜜组分(葡萄糖和果糖)及挥发性物质(烷烃类、酯类、酚类等)没有显著差异, 但黄花花瓣的反射波长范围均在鸟类与蜜蜂视觉范围内, 可同时吸引鸟和蜜蜂。木棉黄花的雌雄异熟程度较小(花粉活性时期与柱头可受期重叠较大), 可在传粉媒介访花次数有限的情况下同时输出或接受花粉。另一方面, 黄花有着更大程度的柱头探出式雌雄异位, 降低了较小程度雌雄异熟可能导致的自交与雌雄功能干扰。木棉存在明显的花粉限制, 且黄花的坐果率(1.08 ± 0.56)%显著低于红花(3.27 ± 0.93)%。我们推测, 黄花个体可能通过增加木棉访花者类群多样性、降低雌雄异熟程度而提高了木棉的繁殖成功率, 也通过吸引蜜蜂而降低了蜜蜂对红花鸟媒传粉的干扰。     

Sexual selection in complex communities: Integrating interspecific reproductive interference in structured populations

The social structure of populations plays a key role in shaping variation in sexual selection. In nature, sexual selection occurs in communities of interacting species; however, heterospecifics are rarely included in characterizations of social structure. Heterospecifics can influence the reproductive outcomes of intrasexual competition by interfering with intraspecific sexual interactions (interspecific reproductive interference [IRI]). We outline the need for studies of sexual selection to incorporate heterospecifics as part of the social environment. We use simulations to show that classic predictions for the effect of social structure on sexual selection are altered by an interaction between social structure and IRI. This interaction has wide‐ranging implications for patterns of sexual conflict and kin‐selected reproductive strategies in socially structured populations. Our work bridges the gap between sexual selection research on social structure and IRI, and highlights future directions to study sexual selection in interacting communities.     

Sexual system in the tanaidacean Falsapseudes bowmani (Crustacea: Malacostraca: Peracarida)

With more than 40,000 species, Malacostraca is the most diverse crustacean class. Most malacostracans are gonochoristic, but simultaneous hermaphrodites are also known. Tanaidacea is one of two malacostracan orders that includes simultaneously hermaphroditic species; so far, simultaneous hermaphroditism has been confirmed externally and internally in only two tanaidacean species, both in the genus Apseudes (Apseudidae). Here we show, through external and internal morphological observations of fixed specimens, that the apseudid Falsapseudes bowmani is a simultaneous hermaphrodite, making Falsapseudes the second tanaidacean genus in which simultaneous hermaphroditism has been confirmed both externally and internally. In this species, the epistome (a projection on the clypeus) was thick and elongate in large specimens but was thin and spiniform in smaller specimens; the brooding of eggs or embryos was observed only in thin‐epistome individuals, although a pair of ovaries was confirmed in both thick‐ and thin‐epistome individuals. This suggests that individuals with a thick epistome may act as males while also retaining the female reproductive organs.     

Pollinator-mediated assortative mating in mixed ploidy populations of Chamerion angustifolium (Onagraceae)

Establishment of polyploid individuals within diploid populations is theoretically unlikely unless polyploids are reproductively isolated, pre-zygotically, through assortative pollination. Here, we quantify the contribution of pollinator diversity and foraging behaviour to assortative pollen deposition in three mixed-ploidy populations of Chamerion angustifolium (Onagraceae). Diploids and tetraploids were not differentiated with respect to composition of insect visitors. However, foraging patterns of the three most common insect visitors (all bees) reinforced assortative pollination. Bees visited tetraploids disproportionately often and exhibited higher constancy on tetraploids in all three populations. In total, 73% of all bee flights were between flowers of the same ploidy (2x–2x, 4x–4x); 58% of all flights to diploids and 83% to tetraploids originated from diploid and tetraploid plants, respectively. Patterns of pollen deposition on stigmas mirrored pollinator foraging behaviour; 73% of all pollen on stigmas (70 and 75% of pollen on diploid and tetraploid stigmas, respectively) came from within-ploidy pollinations. These results indicate that pollinators contribute to high rates of pre-zygotic reproductive isolation. If patterns of fertilization track pollen deposition, pollinator–plant interactions may help explain the persistence and spread of tetraploids in mixed-ploidy populations.     

Sexual differences in life-history traits in the butterfly Lycaena tityrus: a comparison between direct and diapause development

During direct development the butterfly Lycaena tityrus was previously found to display sex-related reaction norms in response to temperature. Based on selection for protandry in males and fecundity selection for larger females, males favoured early emergence over large size, leading to a dramatic weight loss at higher temperatures, whereas females maintained similar weights throughout. Because males were able to avoid a weight reduction relative to females in spite of their shorter development at lower temperatures, sexual size dimorphism existed at higher temperatures only. In the present paper we compare sexual differences in life-history traits in L. tityrus between direct and diapause development at 25 °C. We demonstrate that, regardless of developmental pathway, protandry persisted and relative sexual size dimorphism, with females being larger, remained unchanged. Although diapausing individuals were less time-constrained, allowing them to grow to considerably higher final weights in both sexes, males were not able to reduce their weight loss relative to females. This is explained by the pressure to gain a developmental advantage solely during post-diapause development, whereas direct developing males may spread the burden over the whole larval period. Our results highlight the importance of considering sexual differences in selective pressures, which may influence central life-history traits in manifold ways.     

Influence of pollen limitation and inbreeding depression in the maintenance of incomplete dichogamy in Salvia elegans

The widespread presence of incomplete dichogamy (i.e., partial separation in time between male and female phases) in flowering plants is a long‐standing question in floral evolution. In this study, we proposed four scenarios in which depending on the particular combination of pollen limitation and inbreeding depression, the presence of complete dichogamy, incomplete dichogamy, or adichogamy may be favored. Moreover, we evaluated the role of pollen limitation and inbreeding depression in a natural population of Salvia elegans to test the validity of our predicted scenarios. Our results indicate that S. elegans is partially protandrous as pollen viability and stigma receptivity overlap in the last days of life of the flower. Furthermore, through pollination treatments, we found no evidence of pollen limitation or inbreeding depression in any of the evaluated fitness components. As expected by one of the proposed scenarios, incomplete dichogamy seems to be favored in plants with absence of inbreeding depression and pollen limitation as a way to diminish interference between male and female functions.     

Sexual interference in the Golden snub-nosed monkey (Rhinopithecus roxellana): a test of the sexual competition hypothesis in a polygynous species

Sexual interference (SI), which is defined as any disturbance directed to a mating pair by other individuals, has been reported in several primate species. It is widely suggested that successful harassers experience improved mating success by increasing their access to reproductive partners as well as by reducing the mating success of rivals. Although theories of primate sexual conflict highlight male intra-sexual mating competition, females also are reported to actively disrupt copulations between mating partners. In this study, we investigated SI in a multilevel troop of Golden snub-nosed monkeys (Rhinopithecus roxellana) inhabiting the Qinling Mountains, China. Behavioral observations of 11 one-male units (OMU) that comprised the multilevel troop were conducted from September 2007 to May 2008. During this period 17.1% of 652 documented intra-OMU sexual encounters were characterized either by mild or aggressive forms of harassment. Sexual harassment was typically performed by a single individual (91.9%), and in 75.7% of cases the harasser was an adult or sub-adult female. The frequency of female harassment was positively correlated with the number of adult and sub-adult females residing in an OMU, and resulted in a significant decrease in matings ending in ejaculation. We found that the amount of SI a female received was not a significant predictor of her reproductive success. However, females who conceived during the mating season directed higher levels of harassment at other females than females who did not conceive. We evaluate the strength of the sexual competition hypothesis and the hormonally modulated aggression hypothesis in explaining patterns of SI in female Golden snub-nosed monkeys.     

Effects of soil nitrogen on diploid advantage in fireweed,Chamerion angustifolium (Onagraceae)

In many ecosystems, plant growth and reproduction are nitrogen limited. Current and predicted increases of global reactive nitrogen could alter the ecological and evolutionary trajectories of plant populations. Nitrogen is a major component of nucleic acids and cell structures, and it has been predicted that organisms with larger genomes should require more nitrogen for growth and reproduction and be more negatively affected by nitrogen scarcities than organisms with smaller genomes. In a greenhouse experiment, we tested this hypothesis by examining whether the amount of soil nitrogen supplied differentially influenced the performance (fitness, growth, and resource allocation strategies) of diploid and autotetraploid fireweed (Chamerion angustifolium). We found that soil nitrogen levels differentially impacted cytotype performance, and in general, diploids were favored under low nitrogen conditions, but this diploid advantage disappeared under nitrogen enrichment. Specifically, when nitrogen was scarce, diploids produced more seeds and allocated more biomass toward seed production relative to investment in plant biomass or total plant nitrogen than did tetraploids. As nitrogen supplied increased, such discrepancies between cytotypes disappeared. We also found that cytotype resource allocation strategies were differentially dependent on soil nitrogen, and that whereas diploids adopted resource allocation strategies that favored current season reproduction when nitrogen was limiting and future reproduction when nitrogen was more plentiful, tetraploids adopted resource allocation strategies that favored current season reproduction under nitrogen enrichment. Together these results suggest nitrogen enrichment could differentially affect cytotype performance, which could have implications for cytotypes’ ecological and evolutionary dynamics under a globally changing climate.     

Sexual segregation in habitat use by elephants in Chobe National Park, Botswana

We report on a study conducted on free‐ranging African elephants in the woodlands of northern Botswana. We compared bull groups and family units with regard to (1) their patterns of habitat use and (2) their ranging distances from perennial water sources. During the dry season, adult males frequented more habitat types than family units, whereas family units used a wider diversity of habitats than bulls during the wet season. Bulls roamed widely (>10 km) from perennial drinking water in the dry season, when family units congregated within 3.5 km of the rivers. During the wet season, when ephemeral pans were abundant, all elephant groups were found at intermediate distances (5 km) from the rivers. The spacing of elephants in the dry season is consistent with sexual segregation but we reject the hypothesis that this is an outcome of indirect competition for food, because our concurrent studies on elephant feeding ecology found no evidence for intraspecific competition. Instead, we propose that most adult male elephants space themselves to avoid conflict with musth bulls and roam widely in the dry season between discretely distributed feeding ‘hotspots’. The small proportion of males that are in musth remain close to family units to maximize mating opportunities, and family units are unable to range far from water in the dry season. This is due to (1) comparatively high rates of water turn‐over among juveniles and lactating cows and (2) the reduced mobility of neonates.     

A model of the evolution of dichogamy incorporating sex-ratio selection, anther-stigma interference, and inbreeding depression

Historically, explanations for the evolution of floral traits that reduce self-fertilization have tended to focus on selection to avoid inbreeding depression. However, there is growing support for the hypothesis that such traits also play a role in promoting efficient pollen dispersal by reducing anther-stigma interference. The relative importance of these two selective pressures is currently a popular topic of investigation. To date, there has been no theoretical exploration of the relative contributions of selection to avoid the genetic costs of self-fertilization and selection to promote efficient pollen dispersal on the evolution of floral traits. We developed a population genetic model to examine the influence of these factors on the evolution of dichogamy: the temporal separation of anther maturation and stigma receptivity. Our analysis indicates that anther-stigma interference can favor dichogamy even in the absence of in-breeding depression. Although anther-stigma interference and inbreeding depression are the key forces driving the initial evolution of dichogamy, selection to match the timing of pollen dispersal to the availability of ovules at the population level becomes a more potent force opposing the further evolution of dichogamy as the extent of temporal separation increases. This result may help to explain otherwise puzzling phenomena such as why dichogamy is rarely complete in nature and why dichogamy tends to be associated with asynchronous flower presentation.     

Sexual polymorphism and spatial segregation of Thymelaea hirsuta in Liguria (north-west Italy)

Abstract

Flowering phenology, sexual polymorphism and spatial segregation were studied over two consecutive years in a population of Thymelaea hirsuta (L.) Endl. (Thymelaeaceae) growing on coastal cliffs in north-western Italy. The study population was geographically segregated and, together with Croatian sites, it constituted the northernmost margin of the species' distribution range. Very frequent observations highlighted a sexual tendency towards dioecy already shown by T. hirsuta in other parts of the Mediterranean coast. The flowering period lasted 10?–?11 months, more or less. A high proportion of female flowers and a low number of hermaphrodite flowers were recorded, in contrast with the abundance borne by the species elsewhere; gender segregation into two different subgroups was noted, one consisting mainly of male and protogynous plants, the second consisting basically of female and protandrous specimens.