The mutualism between the dioecious Ficus tinctoria and its pollinators was studied at Hsitzewan, Kaohsiung. I investigated the phenology of F. tinctoria from December 2004 to May 2006. Wasps trapped in sticky boards were recorded from April 2005 to May 2006. The average diameter of receptive figs (B phase) of female F. tinctoria was significantly different from that of functional male trees (P < 0.01), but both with similar coloration. The durations of pre-receptive figs (A phase) through wasp- releasing figs (D phase) or mature figs (E phase) in F. tinctoria were slight longer in winter and spring (8.3-8.8 weeks) than in summer and autumn (6.8-7.0 weeks). The receptive figs were asynchronous both within-tree and among-trees. The receptive figs occurred in every month of 2005 and without seasonality. There were considerable overlaps (82.6%) in the receptive figs between functional male and female trees. The frequency of tender leaf phase of F. tinctoria was positively correlated with local rainfall and temperature, but the frequency of leaf falling phase was negatively correlated with them. The frequency of receptive figs of female trees was only positively correlated with rainfall, and the frequency of their interforal figs (C phase) were both negatively correlated with rainfall and temperature. Fig production of functional male F. tinctoria was not correlated with rainfall and temperature. There are five species of fig wasps, including one species of pollinator (Liporrh opalumgibbosae), one species of Sycoscapter, two species of Philotrypesis, and one species of Neosycophila. No significant difference was found in the average number of 4 species of fig wasps from D phase figs (6.3 pollinators, 8.8 Philotrypesis sp.1, 5.7 P. sp.2, and 5.7 Sycoscapter sp.1 per fig). In addition, one species of Sycophila (Eurytomidae) was also found inside figs of F. tinctoria. The coordination between D phase figs of male F. tinctoria to its B phase was iii 26.2%, which was similar to the coordination of it to B phase of female trees (29.5%). However, more pollinators arrived on male trees than that on female trees. The coordination between pollinators and B phase figs of male trees (43.0%) was higher than that of female trees (14.6%). Fig-pollinator mutualism in F. tinctoria is probably maintained by its asynchronous phenology, and aseasonal changes of the receptive figs. Seed production at female trees can be maintained by a small size of pollinator populations.

This paper presents an analysis of tree-ring growth patterns of male and female Fraxinus mandshurica trees from 1931 to 2007. The specific object was to study the response of radial growth to climate variables separately for male and female trees. The results show that the growth patterns in the two genders were similar during the mid-1950s to 1970s but different in the periods 1931–1940s and 1980–2007. In the period 1980–2007, the mean sensitivity and mean widths of the tree rings were significantly different between the genders (p < 0.05). The climate-growth response in female and male trees was also different. Female trees are sensitive to precipitation in November of the previous year, whereas male trees respond to mean temperature in November of the previous year. The results confirm that climatic sensitivity in male and female trees of dioecious species is different, yet this difference is not stable through time.

Female and male plants often differ in reproductive effort and habitat requirements. The aim of this study was to analyze these differences between the sexes and the effect of climate on tree-ring width in European yew (Taxus baccata). The study was conducted in five yew populations in western Poland. Wood samples were taken from 196 trees (98 females and 98 males) and subjected to the standard procedure of dendrochronological dating. Mean tree-ring width was significantly higher in males since about the beginning of sexual maturity. No such relationship was observed in the youngest population, which is the most distant from the current geographic limit for this species. In most of the analyzed populations, width of tree rings in female individuals, in contrast to males, was negatively correlated with high temperatures in August and September in the year prior to the formation of the tree ring, and correlated positively with precipitation in June and July in the current year. The differentiation of tree-ring width between males and females likely began when the yew trees reached sexual maturity, probably because of the assumed greater reproductive effort of females in comparison with males. The lack of difference in the youngest population may result from a short time since the beginning of sexual maturity or from a milder climate in that region. Different reactions of the two sexes to climate indicate that this may affect the range and viability of populations at the limits of the range.

Unequal sex ratios are widespread in dioecious plants and understanding their cause is important to understanding fundamental aspects of their population dynamics, and yet what causes biased sex ratios in plants is still poorly understood. Competition experiments have been used in plants to predict the outcome of species interactions, but they have rarely been used to help explain sex ratio bias. This study used a response surface competition design to measure the relative competitive abilities of the sexes of the bryophyte Marchantia inflexa (a thallus liverwort of Marchantiaceae) to predict the outcome of competition before the onset of sexual reproductive structures. In bryophytes, dioecy and sex ratio bias is especially common, making them effective organisms for studying sex ratio bias. Given the frequency of female bias in bryophytes, the hypothesis was that females will show a higher competitive ability relative to males. The experiment was conducted in greenhouse conditions at several densities and proportions over the course of seven months. As individuals grew and formed clumps, identities were tracked, and growth measurements were made using photographs and computer imaging software. Both sexes grew on average 41% more with the opposite sex relative to their single-sex cultures. A model predicting future sex ratios showed coexistence between the sexes and predicted a male biased sex ratio of 3.2 males to 1 female. A trade-off was observed for males where single-sex cultures contained more asexual structures than mixed-sex cultures and the reverse for growth rate. Higher levels of asexual reproduction in males in single-sex cultures might be selected for to increase male dispersal for a higher probability of encountering females. This pattern was not found for females. The overyielding results suggest an interaction effect may exist due to niche differentiation between the sexes. In addition, the results suggest that in some dioecious plants a change in sex ratio can occur before differences in their allocation to sexual reproduction.

Flowering plants display diverse reproductive systems, including a variety of gender strategies and mechanisms of clonal propagation. Here, I investigate gender strategies, sex-ratio evolution, and sexual dimorphism in the North American clonal aquatic, Sagittaria latifolia (Alismataceae), which exhibits three sex phenotypes (hermaphrodites, females, males) and two modal sexual systems (monoecy, dioecy). This provides an outstanding opportunity to examine the costs and benefits of combined versus separate sexes. My research focused on the northern range limit in eastern N. America, and on disjunct populations in western N. America. I developed microsatellite (SSR) markers to investigate population genetic structure at several spatial scales, including the clonal structure of local populations to continental patterns. These analyses provided insights on the roles of historical, ecological and reproductive factors in the evolution and maintenance of sexual system diversity. Phenotypic sex ratios varied near continuously from monoecy through subdioecy (three sex phenotypes) to dioecy. A comparison of phenotypic and genotypic sex ratios in dioecious populations demonstrated close correspondence. The northern range limit was characterized by a decline in female frequency and an increased incidence of subdioecy. I evaluated two hypotheses to explain this pattern: (1) increased sex inconstancy in dioecious populations; (2) hybridization between monoecious and dioecious populations. I found support for both hypotheses, although hybridization appears to be the more common pathway to subdioecy. I parameterized a model predicting female frequency and hermaphrodite sex allocation; observed and predicted values were correlated suggesting that subdioecious populations are closer to equilibrium than expected for a clonal perennial. A comparison of eastern and western populations indicated genetic differentiation between monoecy and dioecy in the east, but in the west, due to habitat isolation, geography plays a more important role in genetic differentiation. Evidence from cpDNA haplotype variation indicated that the western range was established following long-distance colonization from the east involving a genetic bottleneck. The discovery of gynodioecious populations of S. latifolia in the west, and the absence of ecological and genetic differentiation between monoecious and dioecious populations, raise the possibility that dioecy may have evolved autochthonously in the west, and more recently than in the eastern range.

Flowering plants display diverse reproductive systems, including a variety of gender strategies and mechanisms of clonal propagation. Here, I investigate gender strategies, sex-ratio evolution, and sexual dimorphism in the North American clonal aquatic, Sagittaria latifolia (Alismataceae), which exhibits three sex phenotypes (hermaphrodites, females, males) and two modal sexual systems (monoecy, dioecy). This provides an outstanding opportunity to examine the costs and benefits of combined versus separate sexes. My research focused on the northern range limit in eastern N. America, and on disjunct populations in western N. America. I developed microsatellite (SSR) markers to investigate population genetic structure at several spatial scales, including the clonal structure of local populations to continental patterns. These analyses provided insights on the roles of historical, ecological and reproductive factors in the evolution and maintenance of sexual system diversity. Phenotypic sex ratios varied near continuously from monoecy through subdioecy (three sex phenotypes) to dioecy. A comparison of phenotypic and genotypic sex ratios in dioecious populations demonstrated close correspondence. The northern range limit was characterized by a decline in female frequency and an increased incidence of subdioecy. I evaluated two hypotheses to explain this pattern: (1) increased sex inconstancy in dioecious populations; (2) hybridization between monoecious and dioecious populations. I found support for both hypotheses, although hybridization appears to be the more common pathway to subdioecy. I parameterized a model predicting female frequency and hermaphrodite sex allocation; observed and predicted values were correlated suggesting that subdioecious populations are closer to equilibrium than expected for a clonal perennial. A comparison of eastern and western populations indicated genetic differentiation between monoecy and dioecy in the east, but in the west, due to habitat isolation, geography plays a more important role in genetic differentiation. Evidence from cpDNA haplotype variation indicated that the western range was established following long-distance colonization from the east involving a genetic bottleneck. The discovery of gynodioecious populations of S. latifolia in the west, and the absence of ecological and genetic differentiation between monoecious and dioecious populations, raise the possibility that dioecy may have evolved autochthonously in the west, and more recently than in the eastern range.

京都大学 / 新制・課程博士 / 博士(理学) / 甲第23743号 / 理博第4833号 / 新制||理||1691(附属図書館) / 京都大学大学院理学研究科生物科学専攻 / (主査)教授 酒井 章子, 准教授 本庄 三恵, 教授 松下 智直 / 学位規則第4条第1項該当 / Doctor of Science / Kyoto University / DGAM

No description available.

Aciphylla species (wild spaniard/speargrass) are an iconic component of the Australasian high country flora, but their reproductive system is enigmatic. They are insect-pollinated dioecious mast seeders (synchronous highly variable seed production), which seems maladaptive. The resource supply to pollinators is highly variable, yet dioecious plants are dependent on pollinators, and dioecious masting requires male and female plants to flower synchronously. Floral display in Aciphylla is relatively large, with tall inflorescences bearing thousands of flowers, suggesting that plants would not have the resources to produce such large stalks every year. But why do they have such huge inflorescences in the first place? I tested whether pollinator attraction is providing an economy of scale which favours intermittent production of very large inflorescences, by manipulating floral display size during a high-flowering year and measuring insect visitation rates and seed set (female reproductive success). Using space-for-time substitution and selective removal of male inflorescences, I also tested whether female seed set was affected by distance to flowering male plants (i.e. changes in local pollen availability) to see if flowering asynchrony would reduce pollination success. Bags were used to exclude pollination by insects and test for wind pollination, and hand pollination was done to test for pollen limitation. Insect surveys suggest that Aciphylla has a generalist pollination system (to avoid satiating a specialist pollinator during 'mast' years'). Male inflorescences received significantly more visits than females, and some seeds were set inside bags (although only 20-30%), suggesting wind pollination may occur at low levels. Seed set rate was higher for taller inflorescences with greater flowering length in A. aurea but tall inflorescences with excess flowers led to a decrease in seed set rates in A. scott-thomsonii. Hand pollination significantly increased seed set rates although these effects were not as large as expected (e.g. 10% increases from natural to hand-pollinated inflorescences were typical). There was no evidence for resource limitation in any species. Female plants in dense flowering populations had higher seed set rates, and individual floral display size in females was particularly important when females were 'isolated' from males. Insect visitation rates were generally higher on inflorescences with a larger floral display, suggesting that display size is important for pollinator attraction. Overall, these results suggest that the pollinator-attraction benefits of such a large floral display (at both the plant and population level) are possibly providing an economy of scale, although the relative effects are small.

Monoecy, the production of distinct male and female flowers on the same plant, is an important, though little studied, sexual strategy in the rainforest understory. This study of a monoecious plant discovered a cue to induce flowering, explored the interplay of gender constraint vs. plasticity in a natural population, and tested possible causes of gender in two laboratory experiments. An experiment in the lab found that reduced photoperiod for three weeks is an unambiguous cue for flowering. The remarkably long inductive period is followed by a long and variable period of floral initiation. This results in only partial synchronization of flowering among plants in a patch, which enhances mating opportunities in this protandrous plant. Inflorescence architecture is highly constrained, and ideally produces a phenotypic gender (proportion female) of about 0.5. However, in the forest at Las Cruces, Costa Rica, most plants were less female than predicted, mostly through abortion of female buds. Plants showed gender plasticity between and within years. Large plants produced more flowers and were more female in gender, and less variable in gender, than small plants. Reproduction was poorly correlated with environmental resource availability, measured as canopy openness, soil moisture, pH, and soil phosphorus, ammonium and nitrate. Phenotypic selection analysis on seed production suggests an optimal gender of 50-60% female, yet plasticity to be less female than this optimum, and in particular to express only male function, has been maintained. In a factorial experiment in the lab, high light or high nitrogen caused plants to produce more flowers and to be proportionally more female, and larger in weight, than low light or nitrogen. The effects of light and nitrogen on reproduction, plant size, and leaf greenness suggest an energy based determination of gender. Gender may be mostly influenced by plant size, but sometimes also opportunistically by environment. Inoculation with mycorrhizas caused plants to be less female in gender, and smaller in weight, than plants that were not inoculated. This suggests a net cost of mycorrhizas under experimental conditions, and supports the emerging view of the mycorrhizal symbiosis as not necessarily mutualistic under all circumstances.