Sexual polymorphism,growth, and reproductive effort in Egyptian Thymelaea hirsuta (Thymelaeaceae)

Gender phenotypes of Thymelaea hirsuta (L.) Endl. were surveyed in six major habitats in the western Mediterranean region of Egypt. Five gender phenotypes were observed at all sites: subandroecious (male), subgynoecious (female), protogynous, protandrous, and gender-labile individuals. Patterns of vegetative growth, reproductive effort, and sex-size relationships were also determined. Females and males were comparable in overall abundance (35.5% and 30.2%, respectively). However, the frequency of sex forms varied significantly among habitats. Plant size (canopy volume) was used as a measure of environmental quality for the species: the coastal dunes were by far the most favorable habitat, followed by the nonsaline depressions, inland plateau, and inland siliceous deposits. The smallest plants were associated with the inland ridges and saline depressions. Protandrous individuals, and female shrubs at the more favorable habitats (e.g., the coastal dunes and nonsaline depressions), had greater canopy volume than males. In contrast, males were larger than females at the less favorable habitats (e.g., the inland plateau and saline depressions). Advantageous growth conditions at the coastal dunes are demonstrated by the greater rate of increase in crown diameter and crown volume noted there, for each size class and sex form. The greatest rates of increase were present in smaller-sized individuals. Size differences between the sex forms were considered in terms of trade-offs between individual growth rate (GR) and reproductive effort (RE); evidence of a trade-off was mixed. Indeed, in a comparison of GR and RE at two extremes of habitat (coastal dune vs. inland plateau), females had greater GR and greater RE at the relatively benign coastal dunes, while at the more harsh, inland plateau site males showed greater GR and RE than females. Peak reproductive effort varied significantly with gender phenotype and habitat type.     

Sexual tetramorphism in Thymelaea hirsuta (Thymelaeaceae): morph ratios in open-pollinated progeny

Natural populations of Thymelaea hirsuta have previously been shown to comprise four distinct sexual morphs: males, females, protogynous individuals, i.e., first female then male, and protandrous individuals, i.e., first male then female. The objective of the present study has been to confirm the genetic basis of this sexual tetramorphism by quantifying morph ratios in the open-pollinated progeny of the four sexual phenotypes growing in a natural population. All four phenotypes were recovered in the progeny of each morph. All observed plants displayed a single sexual phenotype, thus confirming the genetic basis of the tetramorphism. The progeny sex ratios indicate that the genetic determination of sex in this species may be influenced by cytoplasmic factors, while the observed levels of functional female fertility suggest a near-dioecious system. The evolutionary significance of this tetramorphism as a transitional stage in the evolution of dioecy is discussed.     

Sex expression in EgyptianThymelaea hirsuta (Thymelaeaceae) populations

The sex expression inThymelaea hirsuta was assessed in five habitats in the western Mediterranean desert of Egypt. The survey over two main flowering periods indicated that the sexual expression of this plant is complex and labile. Beside the dioecious male and female states, five monoecious states were detected. The dioecious states are more abundant and less labile. There is a clear shift from monoecious towards dioecious states and in general favour of males from the first (late autumn) to the second (late winter) flowering period. These results are discussed in view of the adaptive theories and evolution.     

Breeding system and spatial variation in the pollination biology of the heterocarpic Thymelaea velutina (Thymelaeaceae)

Thymelaea velutina (Thymelaeaceae) is a unique dioecious and heterocarpic shrub, each female producing both dry and fleshy fruits. It is endemic to the Balearic Islands (western Mediterranean) and has suffered an important regression in recent decades, especially due to habitat loss. It lives in coastal areas, mainly in dunes, but a few populations occur up to 1300 m in the mountains of Mallorca. In the present study, we examined its breeding system and pollen vectors, determined whether seed production is pollen-limited, and quantified the relative importance of different insect visitors at coastal and mountain sites. Selective exclusions showed no differences between populations in the relative importance of insects and wind as pollen vectors. Pollen was limiting at both populations, though not consistently in time. Flowers of T. velutina were visited mostly by generalist insects, and species composition and abundance of flower visitors varied both in space and time. The flowering period is about a month later in the mountain than in the dune, and this results in a higher frequency of insect visits to plants in the mountain due to the higher insect abundance; insects were also more diverse in the mountain, although they visited a proportionally lower number of flowers than in the dune. Despite this spatio-temporal variation in the frequency of insect visits and in the importance of a particular insect group, the ambophilous system in this species (previously described as anemophilous) ensures a relatively high fruit set at the two different sites.     

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.     

Sex segregation ratio and gender expression in the genus Actinidia

The sex segregation ratio was checked in bi-parental families of Actinidia deliciosa (2n=6x=174) obtained by crossing four females (A12, Mo3, Br4, Hw1) with two males (T2, M1) and one fruiting male (M3h, subandroecious) according to a factorial mating design. The M3h fruiting male was also selfed. The sex ratio was checked in maternal families of A. kolomikta (2n=2x) and A. chinensis (2n=2x) as well as in A. deliciosa. Seedlings of both diploid species took 3–4 years to progress beyond juvenility, whereas a noticeable number of seedlings from biparental crosses of A. deliciosa involving A12 and Hw1 as seed parents were still non-flowering after seven growing seasons. Open-pollinated families of both diploid and hexaploid species as well as most families from biparental crosses showed a sex segregation ratio approaching 11. Subandroecious lines with different degrees of ovary and pistil development appeared in proportions of 0–4.2%, depending on the cross, but only 6 of the 2567 male vines checked were capable of setting fruit. No case of self-fertility or apomixis was detected among 1866 bagged female vines. Selfed M3h progenies gave only female and male phenotypes in a ratio of 1 female to 3 males. No off-type vines were found among these progenies. The same disomic sex segregation ratio seems to be operating at different ploidy levels in the genus Actinidia. Since selfed fruiting males produced both female and male individuals, the male sex appears to be the heterogametic one. Such evidence indicates that a monofactorial system based on one or more linked genes or on an X/Y chromosome set must be controlling sex expression. How a monofactorial sex-determining mechanism could operate in polyploids to give a 11 female: male ratio is discussed. Minor modifying gene(s) seem to be responsible for the feminization of males, and their expression appears enhanced by environmental conditions. Masculinizing gene(s) seem to be lacking in female genotypes.     

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 segregation in ungulates: a comparative test of three hypotheses

In most social ungulate species, males are larger than females and the sexes live in separate groups outside the breeding season. It is important for our understanding of the evolution of sociality to find out why sexual segregation is so widespread not only in ungulates but also in other mammals. Sexual body size dimorphism was proposed as a central factor in the evolution of sexual segregation in ungulates. We tested three hypotheses put forward to explain sexual segregation: the predation-risk, the forage-selection, and the activity budget hypothesis. We included in our analyses ungulate species ranging from non-dimorphic to extremely dimorphic in body size. We observed oryx, zebra, bighorn sheep and ibex in the field and relied on literature data for 31 additional species. The predation-risk hypothesis predicts that females will use relatively predator-safe habitats, while males are predicted to use habitats with higher predation risk but better food quality. Out of 24 studies on different species of ungulates, females and their offspring chose poorer quality but safer habitat in only eight cases. The forage-selection hypothesis predicts that females would select habitat based on food quality, while males should prefer high forage biomass. In fact, females selected higher quality food in only six out of 18 studies where males and females segregated, in eight studies there was no difference in forage quality and in four studies males were in better quality habitat. The activity budget hypothesis predicts that with increasing dimorphism in body size males and females will increasingly differ in the time spent in different activities. Differences in activity budgets would make it difficult for males and females to stay in mixed-sex groups due to increased costs of synchrony to maintain group cohesion. The predictions of the activity budget hypothesis were confirmed in most cases (22 out of 23 studies). The heavier males were compared to females, the more time females spent foraging compared to males. The bigger the dimorphism in body mass, the more males spent time walking compared to females. Lactating females spent more time foraging than did non-lactating females or males. Whether species were mainly bulk or intermediate feeders did not affect sexual differences in time spent foraging. We conclude that sexual differences in activity budgets are most likely driving sexual segregation and that sexual differences in predation risk or forage selection are additive factors.     

Sexual segregation in Tibetan gazelle: a test of the activity budget hypothesis

Tibetan gazelle Procapra picticaudata , a threatened endemic species on the Qinghai-Tibet Plateau, is a sexually dimorphic ungulate. Males are larger than females and both sexes live in different groups, except during the rutting season. In order to test the activity budget hypothesis, which was proposed to explain social segregation in ungulates, we studied the activity budget of male and female Tibetan gazelles during the summers of 2005 and 2006. The activity budget hypothesis predicts that females spend more time feeding and group members synchronize their activities more in the same-sex groups than in the mixed-sex groups. We found that females and males of Tibetan gazelle had different activity budgets; females spent significantly longer time feeding and comparably less time on other activities. Activity synchronization indexes between female groups and male groups were similar (female groups: 0.81±0.17, male groups: 0.80±0.19), whereas both of them were significantly higher than that of mixed-sex groups (0.58±0.29). These results suggest that although sexual segregation might be caused by multiple mechanisms, the activity budget hypothesis about sexual segregation is supported by our studies of Tibetan gazelle on the Qinghai-Tibet Plateau.     

关于兰科Eulophia hirsuta T. P. Lin的归属问题

根据花的构造将短毛美冠兰Eulophia hirsuta 移入粉口兰属Pachystoma。由于粉口兰属已经有Pachystoma hirsutum(1987),故另起新名,即绿岛粉口兰Pachystoma ludaoense。     

Origin and diversification of Thymelaea (Thymelaeaceae): inferences from a phylogenetic study based on ITS (rDNA) sequences

Thymelaea is a Mediterranean genus belonging to a primarily tropical and subtropical family. This genus is here presented as a particular case on which the hypothesis of an in situ evolution of the Mediterranean flora from a Tertiary subtropical stock can be phylogenetically tested. To better understand the evolutionary history of Thymelaea, molecular phylogenies based on ITS (rDNA) sequences are estimated. Phylogenetic analyses strongly support the monophyly of Thymelaea and the sister position of Daphne. Unexpected relationships between sequences of T. granatensis, T. pubescens and T. sanamunda are found and explained as a case of cryptic nuclear introgression. Phylogenetic analyses also reveal the polyphyletic character of Diarthron, sensu Tan, suggesting that Dendrostellera and Stelleropsis should be reinstated as independent genera. Based on the resulting phylogenetic hypothesis and using statistical tests it is suggested that Thymelaea experienced an initial rapid diversification. Time estimates for the origin and radiation of Thymelaea based on independent sources of evidence—paleontological, paleoclimatic and paleogeographic data, on one hand, and ITS substitution rates reported for other angiosperm groups, on the other—give strikingly similar results. The early radiation of Thymelaea is inferred to have occurred in the Upper Miocene, after this lineage diverged from that of Daphne in the Lower or Middle Miocene. Both evolutionary events are related to the overall climatic deterioration (declining temperature and increasing dryness) registered throughout the Miocene in Eurasia. The rapid and extensive diversification of Thymelaea is placed in the Iberian Peninsula or here and in the neighbouring North African territory.     

Sexual competition and N supply interactively affect the dimorphism and competiveness of opposite sexes in Populus cathayana

Several important dioecious species show sexual spatial segregation (SSS) along environmental gradients that have significant ecological effect on terrestrial ecosystem. However, little attention has been paid to understanding of how males and females respond to environmental gradients and sexual competition. We compared eco‐physiological parameters of males and females of Populus cathayana under different sexual competition patterns and nitrogen (N) supply levels. We found that males and females interacting with the same or opposite sex showed significant differences in biomass partition, photosynthetic capacity, carbon (C) and N metabolism, and leaf ultrastructure, and that the sexual differences to competition were importantly driven by N supply. The intersexual competition was enhanced under high N, while the intrasexual competition among females was amplified under low N. Under high N, the intersexual competition stimulated the growth of the females and negatively affected the males. In contrast, under low N, the males exposed to intrasexual competition had the highest tolerance, whereas females exposed to intrasexual competition showed the lowest adaptation among all competition patterns. Sexual competition patterns and N supply levels significantly affected the sexual dimorphism and competitiveness, which may play an important role in spatial segregation of P. cathayana populations.     

毛钩藤的小孢子发生和雄配子体发育

在光学显微镜和透射电镜下观察了毛钩藤(Uncaria hirsuta Havil.)的小孢子发生和雄配子体发育过程.结果表明,毛钩藤花两性,具5枚雄蕊,花药4室,花药壁由表皮、药室内壁、中层和绒毡层组成,花药开裂时,药室内壁高度纤维化带状加厚.花药壁的发育方式属于双子叶型,小孢子母细胞减数分裂的胞质分裂为同时型.小孢子在四分体时期开始沉积花粉外壁,小孢子大液泡化时期开始沉积花粉内壁.成熟花粉为2-细胞型.毛钩藤的花粉发育特征和茜草科植物基本一致.毛钩藤绒毡层属于分泌型,双重起源,分别起源于次生周缘层和药隔细胞.小孢子发育早期绒毡层开始降解并分泌形成大量乌氏体,花药开裂时绒毡层完全消失,剩下少量乌氏体.小孢子早期内壁加厚突出形成,小孢子细胞核分裂以后内壁加厚开始脱落,花药开裂时,只剩下少量的内壁加厚突出.初步推测,内壁加厚突出与乌氏体共同作用为雄配子体的发育提供营养物质.