Genetic variation in Ecballium elaterium (Cucurbitaceae): Breeding system and geographic distribution

Dioecy, the separation of sexes, has arisen independently many times in the course of angiosperm evolution. Avoidance of inbreeding is clearly involved in the evolution of dioecy, and as a consequence we predict that dioecious populations should maintain higher levels of genetic variation than closely related nondioecious populations. We tested that prediction by comparing allozymic variation in two closely related taxa, the monoecious and dioecious subspecies of the Mediterranean cucurbit, Ecballium elaterium. Thirteen polymorphic loci were screened for seeds sampled from 10 monoecious and 13 dioecious populations spanning the geographic ranges of the subspecies in Spain. The dioecious subspecies showed strikingly greater allelic diversity and heterozygosity than the monoecious subspecies. A hierarchical F-statistic analysis clearly demonstrated considerable genetic variation within populations for the dioecious populations, whereas for the monoecious populations almost all genetic variation resulted from differences among populations. The general pattern of homozygosity within monoecious populations suggests that they are highly inbred. In order to assess historical influences on current patterns of genetic variation, we conducted a genetic-distance analysis. The observed relationship between genetic distance and geographic distance between populations supports the hypothesis that the subspecies' current allopatric distributions on the Iberian Peninsula are the result of separate waves of colonization from the north (monoecious) and south (dioecious).     

Survival costs and reproductive benefits of floral display in a sexually dimorphic dioecious shrub, Leucadendron xanthoconus

The evolutionary causes of sexual dimorphism in plants have not been as widely studied as in animals and the importance of sexual selection in causing dimorphism remains controversial. Sexual selection is most obvious when it favours the evolution of a trait which enhances mating success at the expense of decreased viability. We studied the relationship between floral display (number of inflorescences), pollinator attraction and plant survival in a dioecious shrub, Leucadendron xanthoconus. Pollinator attraction, measured as the number of insect pollinators, increased linearly with floral display in males. However, males with extravagant displays had a higher probability of dying. Our data suggest that male plants are undergoing selection on floral display for increased mating success counterbalanced by selection against plants with extravagant displays. Seed set in females did not increase with floral display, except at very low inflorescence numbers. Nor was female survival correlated with floral display. Because inflorescences are terminal in the species, selection for more inflorescences in males causes increased ramification, thinner terminal branches and smaller leaves. Thus vegetative dimorphism in this species appears to be caused by selection for extravagant floral display in males, but not females. Limits to dimorphism are imposed by survival costs of elaborate display.     

Pollination by deceit in nutmeg (Myristica insipida, Myristicaceae): floral displays and beetle activity at male and female trees

Pollination by deceit in Myristica insipida, a beetle-pollinated nutmeg, was hypothesized to operate on intersexual differences in flower production and longevity, producing a daily fluctuation between floral display maxima and minima. Sticky traps were used to continuously monitor beetle activity. Flower production and naturally occurring intersexual differences in display were recorded. Male and female trees flowered in synchrony producing daily display maxima at 1800-0600 and display minima at 1400-1800. Rewarding male trees produced three times the number of flowers of female trees, but the greater longevity of female flowers reduced the intersexual difference in display maxima to a factor of two. There was no intersexual difference in display minima. Beetles were demonstrated to be sensitive to differences in both maximum and minimum displays on rewarding male trees, a necessary prerequisite for directional selection on display size. Beetle captures were significantly higher at male trees during floral display maxima, and no intersexual differences in capture rate occurred during floral display minima. However, capture rates at male trees did not decline as predicted, and the pattern of captures was consistent with crepuscular activity. Beetle captures at male and female trees were lower from 1800 to 0600 and 1000 to 1400, and higher from 0600 to 1000 and 1400 to 1800, but the differences were only significant at female trees. These data suggests that foraging errors are numerous, frequent, and the result of overall foraging activity.     

Sexual dimorphism in the andromonoecious Euphorbia nicaeensis: effects of gender and inflorescence development

BACKGROUND AND AIMS: In andromonoecious taxa with separate floral types along the inflorescence, architectural or plastic effects can simulate floral sexual dimorphism. Both the primary and secondary sexual characteristics of the cyathia of the protogynous andromonoecious species Euphorbia nicaeensis were analysed according to their sex and arrangement on the inflorescence. METHODS: The production of male and hermaphrodite cyathia at each inflorescence level was surveyed in two natural populations. The longevity, size, pollen production and viability, and nectar secretion of both types of cyathia were checked between inflorescence levels and between sexes at the only level at which they occur together. This sampling method makes it possible to know whether differences between cyathia types are based on sex or are attributable to inflorescence development. KEY RESULTS: Male cyathia were produced predominantly at the first and second inflorescence levels, whereas at levels 3-5, the cyathia were almost exclusively hermaphrodite. Viable pollen production by male cyathia at the second inflorescence level was higher than that of hermaphrodite cyathia at the third level but, when males and hermaphrodites at the same level were compared, their pollen production was similar. Male and hermaphrodite cyathia were similar in size, irrespective of the inflorescence level, although the exclusively hermaphrodite cyathia of the last level were smaller. Both cyathium types produced similar amounts of sugar. However, male cyathia produced nectar during their whole lifespans, whereas hermaphrodites produced it exclusively during their male phase. Moreover, the nectary activity of male cyathia started earlier in the day than that of hermaphrodites. CONCLUSIONS: An apparent floral dimorphism exists in the primary sexual characteristics of Euphorbia nicaeensis because differences in pollen production between cyathium types are due to theirs positions. Similarly, differences affecting most secondary sexual characteristics are only apparent between the two cyathium types. However, E. nicaeensis shows a true but slight floral dimorphism in some of the secondary sex characters related to nectar secretion. The lack of nectar production by the female phase of the hermaphrodite cyathia of E. nicaeensis indicates that this is a deceit-pollinated species.     

Gender variation of sequential inflorescences in a monoecious plant Sagittaria trifolia (Alismataceae)

In protogynous plants, female flowers of early blooming plants are at a reproductive disadvantage because they cannot set fruit due to the lack of available pollen. To study this phenomenon, gender expression of the monoecious herb Sagittaria trifolia was investigated over the entire flowering season in two field and two cultivated populations in Hubei and Hunan Provinces, China. In racemes of S. trifolia, flowers open sequentially from bottom to top, with female flowers opening first followed by male flowers. This creates a temporal separation of sexes in the species. Under field conditions small plants are often male, with production of both male and female flowers increasing with plant size. Femaleness increased among sequential inflorescences since female flower production increased whereas male flower production did not. Seed production was greater in large inflorescences because they contain more female flowers, and the number of ovules increased in female flowers at basal positions within the raceme. A consistent pattern of high seed set was observed in flowers from both field and cultivated populations. About 1 % of unfertilized ovules resulted from no pollination and 2 % of the seeds produced were only partly developed due to resource limitation. In the first inflorescence of the six experimental populations, 6.7-40.0 % of individuals produced only male flowers, and female flowers of 1.9-6.5 % individuals were aborted. The occurrence of male flowers in early blooming inflorescences could be an adaptive strategy to conserve resources and enhance pollination of female flowers in protogynous S. trifolia.     

Sexual behaviour and evolution of sexual dimorphism in body size in Jaera (Isopoda Asellota)

Individuals of the genus Jaera do not mate at random. In the species from the Mediterranean group, J. italica and. J. nordmanni, large males and medium sized females are at an advantage and their sizes are positively assorted. These effects are attributable to sexual competition between males. In the Ponlo-caspian species J. istri, no advantage of large males exists, but sexual selection could be the cause for a long passive phase prior to copulation and for normalizing selection upon female size at pairing. In the Atlantic species, J. albifrons, no selection can be ascertained.
Differential mating success in males appears as one of the causes of the evolution of sexual dimorphism in body size, which makes males larger, of equal size, or smaller than females according to the species. The reason for this reversal in dimorphism seems to differ in the two sexes. Sexual selection provides an explanation for the evolution of male size, while the interspecific changes in female length are more likely due to ecological factors.     

Foraging behavior of three bee species in a natural mimicry system: female flowers which mimic male flowers in Ecballium elaterium (Cucurbitaceae)

Summary The behavior of Apis mellifera and two species of solitary bees which forage in the flowers of monoecious Ecballium elaterium (L.) A. Rich (Cucurbitaceae) were compared. The female flowers of E. elaterium resemble male flowers visually but are nectarless, and their number is relatively smaller. Apis mellifera was found to discriminate between the two genders and to pay relatively fewer visits to female flowers (mean of 30% relative to male flowers) from the beginning of their activity in the morning. The time spent by honeybees in female flowers is very short compared to that spent in male flowers. It is surmised that the bees remember the differences between the flowers where they foraged on the previous days. In contrast, the two species of solitary bees Lasioglossum politum (Morawitz) (Halictidae) and Ceratina mandibularis Fiese (Anthophoridae) visit the female flowers with nearly equal frequencies at the beginning of each foraging day and stay longer in these flowers. Over the day there is a decline in the relative frequency of visits to female flowers and also in the mean time spent in them. The study shows that bees can collect rewards at high efficiency from the flowers of Ecballium elaterium because of their partial discrimination ability and the scarcity of the mimic flowers. It is suggested that the memory pattern of some solitary bees may be different from that of Apis mellifera. It seems that the limited memory and discrimination ability of bees can lead to a high frequency of visits to the mimic flowers during a long flowering season.     

Systematics and evolution of the Disa draconis complex (Orchidaceae)

The systematic status of Disa draconis (L.f.) Sw. is revised following extensive field studies of population variation in the Western Cape, South Africa. Principal component and cluster analyses revealed clear distinctions between populations from sandplain, semi-arid and montane environments. Diagnostic characters were found in each of the population clusters indicating the existence of a species complex, rather than a single taxon as in the current taxonomy. We propose, therefore, that the name D. draconis (L.f.) Sw. be restricted to the individuals of the sandplain populations from which the type was collected. We reinstate an earlier name, Disa harveiana Lindl., to describe the montane populations which possess several autapomorphic characters. We also show that the geographical variation in spur length and flowering time within this species can be partitioned into two geographically distinct subspecies: D. harveiana subsp. harveiana and D. harveiana subsp. longicalcarata Johnson & Linder. The populations from the semi-arid Karoo region were recognized as a distinct new species, Disa karooica Johnson & Linder, on the basis of their peculiar petal structure. A cladistic analysis indicated that the three species forming the D. draconis complex form a monophyletic and relatively specialized lineage within Disa sect. Coryphaea. The revised classification also has important conservation implications as D. draconis , previously considered a common species, is now restricted to a few highly threatened populations on the west coast near Cape Town.     

Impact of plant flowering phenology on the cost/benefit balance in a nursery pollination mutualism,with honest males and cheating females

This study documents the flowering phenology and its potential consequences on a nursery pollination mutualism between a dioecious plant, in which honest male plants, but not cheating females, allow the specific pollinator to reproduce within inflorescences. Very few pollinators were found to emerge during plant anthesis, leading to a low (if any) potential benefit through pollen dispersal. This opens the question why male plants do not also cheat their pollinators. Female plants flowered late in the season, when many males had just achieved their own anthesis, which increased the efficiency of pollen transfer. Finally, some late‐flowering males reached their anthesis simultaneously with females, which open the possibility for pollinator to choose between honest males and cheating females. Nevertheless, female plants were found to produce fruits, even though fruit production was limited by pollen (and pollinator) supply, meaning that cheating was not entirely retaliated by the mutualistic partner.     

Pollinator responses to variation in floral display and flower size in dioecious Sagittaria latifolia (Alismataceae)

In animal-pollinated plants with unisexual flowers, sexual dimorphism in floral traits may be the consequence of pollinator-mediated selection. Experimental investigations of the effects of variation in flower size and floral display on pollinator visitation can provide insights into the evolution of floral dimorphism in dioecious plants. Here, we investigated pollinator responses to experimental arrays of dioecious Sagittaria latifolia in which we manipulated floral display and flower size. We also examined whether there were changes in pollinator visitation with increasing dimorphism in flower size. In S. latifolia, males have larger flowers and smaller floral displays than females. Visitation by pollinators, mainly flies and bees, was more frequent for male than for female inflorescences and increased with increasing flower size, regardless of sex. The number of insect visits per flower decreased with increasing floral display in males but remained constant in females. Greater sexual dimorphism in flower size increased visits to male inflorescences but had no influence on the number of visits to female inflorescences. These results suggest that larger flower sizes would be advantageous to both females and males, and no evidence was found that females suffer from increased flower-size dimorphism. Small daily floral displays may benefit males by allowing extended flowering periods and greater opportunities for effective pollen dispersal.     

Dioecy and the evolution of pollination systems inSchiedea and Alsinidendron (Caryophyllaceae:Alsinoideae) in the Hawaiian Islands

The transition from biotic to wind pollination and the consequencesof this transition for the evolution of dioecious breeding systems wereinvestigated in Schiedea and Alsinidendron(Caryophyllaceae: Alsinoideae), genera endemic to the Hawaiian Islands. The potential for wind pollination was studied for five species ofSchiedea using a wind tunnel. Morphological correlates of windpollination for these species were then used to infer the presence orabsence of wind pollination in the remaining Schiedea species. Hermaphroditic Alsinidendron and Schiedea species,which occur in mesic to wet forests, or less commonly in dry habitats,show little or no indication of wind pollination. These species had lowpollen:ovule ratios, large relative pollen size, diffuse inflorescences,substantial nectar production in several cases, and appear to bebiotically pollinated or autogamous. Sexually dimorphic species, whichall occur in dry habitats, are wind pollinated, based on wind tunnelresults or morphological adaptations indicating the potential for windpollination. These adaptations include high pollen:ovule ratios, smallpollen size, moderately to highly condensed inflorescences, and reducednectaries and nectar production. Shifts to wind pollination anddimorphism are strongly correlated in Schiedea, suggesting theclose functional relationship of the twophenomena.     

Breeding biology, resource partitioning and reproductive effort of a dioecious shrub, Clutia pulchella L. (Euphorbiaceae)

 The flowering of Clutia pulchella was studied in a coastal scarp forest community in southern KwaZulu-Natal, South Africa. Sex dependent differences in the frequency and magnitude of floral output were recorded, with male individuals producing 4.32 times more flowers per leaf axis than females. Increased node production, as a manifestation of significantly increased branching, allowed for prolific flower production in males. Energy investment per flower was marginally greater in females (0.045 J/mg), despite copious pollen production and glandular secretion in males (0.035 J/mg). Differential flower production, reproductive organ dry mass and an unbalanced branching ratio revealed that “per plant” expenditure towards flowering was more pronounced in males (♀ flowering × 5.04). The elevated cost of fruit production (♀ flowering × 5.5), inclusive of imperfect fruit set, resulted in overall reproductive expense being slightly higher for females. Such sex-dependent reproductive investment was shown to have minimal influence on the population sex ratio, with sex frequencies and nearest neighbor distances yielding a non-significant male bias of 1.16. No evidence of spatial asymmetry was found. Received December 1, 1999 Accepted September 27, 2000     

Maintenance of gynodioecy in Wurmbea biglandulosa (Colchicaceae): gender differences in seed production and progeny success

 In gynodioecious species, females contribute genes to future generations only through ovules, and to persist in populations they must have a compensatory advantage compared with hermaphrodites that reproduce via ovules and pollen. This compensation can result from greater fecundity and/or superior success of progeny from females. We examined differences in seed production and progeny success between females and hermaphrodites in the geophyte Wurmbea biglandulosa to explain the maintenance of females. Females produced more ovuliferous flowers and had more ovules per flower than did hermaphrodites but this did not necessarily result in greater fecundity, in part because seed production of females was pollen-limited. Over four years in one population, open-pollinated females produced 1.32 more seeds than open-pollinated hermaphrodites (range 1.09–1.63). In two other populations examined for one year only females produced 1.07 and 0.79 as many seeds as hermaphrodites. Seed production of open-pollinated females and hermaphrodites was only 55% and 73% that of cross-pollinated plants, respectively, indicating that both genders were pollen-limited but females more so than hermaphrodites. Open-pollinated seeds from females were 1.18–1.27 times more likely to germinate than seeds from hermaphrodites. No gender differences existed in seedling growth or survival. Hermaphrodites were self-compatible, but selfed seed set was only 80% that of crossed seed set. Crossed seed set of females and hermaphrodites did not differ. Assuming nuclear control of male sterility, relative female fitness is insufficient to maintain females at their current frequencies of 17%, and substantial female fitness advantages at later life-cycle stages are required. Received May 4, 2001 Accepted February 25, 2002     

Deceit pollination and the effect of deforestation on reproduction in dioecious Schisandra sphenanthera (Schisandraceae) in central China

The fruits of Schisandra sphenanthera (dioecious) are used to produce ‘Nan-Wuweizi',an important Chinese medicine.However,long-term exploitation and habitat destruction have threatened these plants in ...     

Function and evolution of the frill of the frillneck lizard, Chlamydosaurus kingii (Sauria: Agamidae)

Relative to body size, the frill of the Australian agamid lizard Chlamydosaurus kingii is one of the largest and most spectacular display structures seen in any animal species. More than 300 hours observation of free-ranging lizards, combined with data on museum specimens, revealed that the frill is used primarily for intraspecific communication and predator deterrence. Earlier hypotheses on alternative uses for the frill (gliding, food storage, thermoregulation or auditory enhancement) are not supported. The folded frill may also enhance camouflage, but this is probably a fortuitous effect rather than an adaptation.
Male frillnecks frequently display and fight during the mating season. Male displays are highly stereotyped, and involve repeated partial erection of the frill, head-bobbing, tail-lashing, and waving of forelimbs. Both males and females erect the frill during social encounters, and in response to potential predators. Males grow larger than females and have larger heads than do females at the same body size, but no dimorphism is apparent in the relative size of the frill. The extreme development of the display structure in this species may be due to general allometric relationships, as well as to ecological features that have intensified the action of sexual selection in Chlamydosaurus.     

三室算盘子寄主上弯头细蛾的生物学特性及花气味成分分析

【目的】为了从生物学和化学生态学角度探讨弯头细蛾Epicephala ancylopa和寄主三室算盘子Glochidion sp.间专性传粉的互利共生关系稳定性。【方法】本研究在野外观察和室内实验的基础上,对专性传粉育幼互利共生体系中三室算盘子、弯头细蛾生物学特性进行详细研究,探究互利共生双方利益得失;用动态顶空吸附法分别收集三室算盘子雄花和雌花气味物质,运用气相色谱-质谱联用技术(GC-MS)分析并鉴定其有效成分,用峰面积归一化与内标法定性定量;最后通过主成分分析法比较雄花和雌花之间气味化学成分的差异性。【结果】弯头细蛾在云南省普洱太阳河国家森林公园每年有1个世代,成虫和幼虫的活动时间分别在3-4月和8-10月。三室算盘子结实率为44.20%,被蛀食率为69.94%,平均每头幼虫消耗2.55枚种子来满足自身生长发育,寄主植物留有83.06%完好的种子,以维持互利共生关系的稳定。三室算盘子雌雄花气味中共鉴定出24种挥发物,主要以单萜类和倍半萜类物质为主,其中(Z)-罗勒烯和β-榄香烯两种萜类物质含量最高(分别为47.11%和22.72%),推测其是吸引弯头细蛾传粉的主要气味成分;雄花和雌花之间气味化学成分存在明显的差异,具有两性异型性。【结论】弯头细蛾通过以卵越夏和以蛹越冬对策,实现成虫发生期与三室算盘子花期的精准匹配。弯头细蛾成虫白天静伏,傍晚开始活动,三室算盘子花的气味物质也只在晚上才明显释放,且雌雄花气味化学成分的两性异型性有利于弯头细蛾辨别雌雄花,以完成采集花粉与传粉行为。该研究结果为头细蛾属昆虫与算盘子属植物专性传粉互利共生关系稳定性的维持机制提供了新的依据,也为深入开展通过触角电生理检测和生物行为实验来筛选吸引传粉头细蛾的活性物质提供了理论依据。     

Sex,reproductive cycle and variations in the fecundity of the family schilbeidae (Osteichthyes : Siluriformes) in Lake Kainji,Nigeria.

The sex, reproductive cycle and variations in the fecundity of the family Schilbeidae (Osteichthyes : Siluriformes) in Lake Kainji, were investigated. Sexual differences in size and morphology were found to exist in all species. Sex ratio varied with species and season. There were more females than males in the population. Size at maturity varied from species to species with the males attaining maturity at a slightly lower length than the females. Breeding condition was attained mainly during the rains in Eutropius niloticus, Schilbe mystus and Siluranodon auritus and in the dry and rainy seasons in Physailia pellucida. Spawning occurred once during the breeding season in E. niloticus and S. mystus; and more than once during the prolonged breeding season in P. pellucida. Condition factors did not show any correlation with maturation of gonads. Fecundity was found to vary from species to species.