Michiko Masuda

Evolution of Campanula Flowers in Relation to Insect Pollinators on Islands

Since Darwin’s observations on the biota of the Galapagos, the evolution of island plants has attracted the attention of many investigators (Carlquist, 1974). Adaptive radiation of organisms on oceanic islands continues to be one of the most interesting topics in evolutionary biology. Opportunities for studying evolution are excellent on islands because of the high levels of variation. The development of the theory of island biogeography has offered a basis for the study of the ecology and evolution of island biotas in relation to neighboring continents (MacArthur and Wilson, 1967; MacArthur, 1972). The theory predicts that the number of species on an island will decrease with distance from the mainland and that species with greater colonizing ability are more likely to establish on a given island. Plant species on islands may evolve through interactions with flora and fauna that are different from those on the nearby mainland.

Differentiation of spring emerging and autumn emerging ecotypes in Galium spurium L. var. echinospermon

SummaryEcotypes of Galium spurium L. var. echinospermon with distinct germination phenologies were found to occur in two adjacent plots of a grassland nature reserve with different management histories: a spring-germinating population is present in a winter-burnt plot, and an autumn-germinating type in an unburnt plot. These ecotypes share common flowering and fruiting phenologies, and disperse their seeds in early summer. Markedly contrasting thermal dormancy/germination characteristics were demonstrated for their seeds in systematic laboratory tests performed after several types of seed storage including storage in the field. The primary dormancy of seeds of the spring germinator was removed by moist-chilling or field winter-chilling, while that of the autumn germinator was removed by moist storage at 25°C or field summer temperatures. Biseasonal seedling emergence of the species appears to be due to a local differentiation of distinct ecotypes.

Reproductive ecology of a cleistogamous annual, Impatiens noli-tangere L., occurring under different environmental conditions

Demographic and reproductive schedules were compared among five populations of a cleistogamous annual,Impatients noli-tangere L., occurring in habitats with contrasting moisture and/or light environments. In all populations, flowering extended for 2–3 months during which light environment and mortality changed. Seasonal patterns of growth and mortality were significantly different among the five populations studied. The beginning and duration of flowering and the ratio of chasmogamous flowers to cleistogamous ones were also significantly different among populations. An experiment was conducted under different light conditions (open and closed) to separate the genetic and environmental components of the variation in reproductive traits observed among populations ofI. noli-tangere. Transplanted plants showed significant among-population variation in flowering time, as is observed in natural populations, suggesting genetic differentiation among populations of this species. On the other hand, the ratio of chasmogamous flowers did not differ among plants transplanted from three populations. Based on these results, the authors suggest that facultative cleistogamy is a conditional strategy under seasonally changing environments.

Different responses of pollinating bees to size variation and sexual phases in flowers ofCampanula

To examine the response of pollinating bees to size and sexual phases of flowers, we constructed an artificial population ofCampanula having large flower variation and presented it to potentially pollinating bees in nurseries. The pollinating bee groups (halictid, megachilid and bumble bees) responded differentially to both the flower size and to the sexual phases of the flowers. Whereas visitation rate of megachilid bees increased with the flower size, those of halictid bees and bumble bees did not show particular trends; for example, bumble bees visited almost all of the flowers consistently. Visitation frequencies to male-and female-phased flowers were significantly different between megachilids at Tokyo and halictids. This study indicates that pollinator attraction could not solely explain the evolution of the flower size inCampanula, and that other factors such as pollen transfer efficiency, should be considered.

Evolution of floral dimorphism in a cleistogamous annual, Impatiens noli-tangere L. occurring under different environmental conditions

The resource allocation models for the evolution of a mixed production of cleistogamous and chasmogamous flowers were tested in four natural populations of Impatiens noli-tangere L. When we incorporated the estimated parameter values into our basic model, under the assumption that the environmental conditions did not change through the reproductive season, the model predicted the obligate chasmogamy in all four populations. However, including the fluctuation of the fertility of chasmogamous flowers in the model, the seasonal switching of chasmogamous and cleistogamous flower production becomes an evolutionary stable strategy, which explains the reproductive behavior of the individuals in two populations. Only after the relationship between the geitonogamy of chasmogamous flowers and the number of chasmogamous flower production was considered in the model did the simultaneous production of both of the dimorphic flowers become an evolutionary stable strategy. Further, the complete model clarifies the reproductive behavior of the individuals in the other two populations.

Effects of Salinity and Temperature on Seed Germination in a Japanese Endangered Halophyte Triglochin maritimum (Juncaginaceae)

Triglochin maritimum (Juncaginaceae). Germination tests were carried out at three salinity levels (0,200,400 mM NaCl in which seeds were exposed to increasing- or decreasing temperatures. Effects of moist-chilling pretreatment (stratification) in 0,200 and 400 mM NaCl on seed germination at 0,200 and 400 mM NaCl, respectively, were also examined. Under the highest salinity condition (400 mM NaCl), no germination was observed. The seeds germinated very well (88%) in fresh-water after 5-month moist-chilling pretreatment. Longer moist-chilling pretreatment resulted in higher germination percentages. Moist chilling pretreatment in 0,200 and 400 mM NaCl for 5 month enhanced germination percentages to 10% (in 400 mM NaCl) –88% (in 0 mM NaCl).

Spatial Genetic Structure within Two Populations of a Self-Incompatible Perennial, Chionographis japonica var. japonica (Liliaceae)

Intrapopulational spatial genetic structure was examined in two populations ofChionographis japonica var.japonica, a self-incompatible perennial, by spatial autocorrelation analysis of enzyme polymorphism. Although most spatial autocorrelation indices (MoransI) in the shortes distance class were significantly positive, most in the other distance classes did not significantly deviate from the values expected from random distributions of genotypes in both populations. This contrasts with a spatial genetic pattern previously reported for a population of the predominantly selfing congener,C. japonica var.kurohimensis, indicating that pollen-mediated gene flow highly impedes genetic substructuring within populations of outcrossingC. japonica var.japonica. Genetic similarity in very proximate distance found in outcrossingC. japonica var.japonica is probably due to restricted dispersal of seeds.

Dispersal of chasmogamous and cleistogamous seeds inViola hondoensis W. Backer et H. Boiss

To test generality of the near-and-far dispersal model proposed by Schoen and Lloyd (1984), we compared dispersal distances of cleistgamous and chasmogamous seeds ofViola hondoensis. Dispersal distance was not significantly different between cleistogamous and chasmogamous seeds, implying that the near-and-far dispersal model cannot apply toV. hondoensis. Alternatively, little pollinator availability in summer and autumn seems to be a major factor favoring facultative cleistogamy in this species.

Identification of conservation measures to protect the Japanese endangered plant species Aster kantoensis

To identify the factors responsible for degrading the habitat of the endangered plant species Aster kantoensis, as well as the vulnerable life stage where this occurs, we carried out sowing experiments. Two natural habitats were simulated, being situated along the floodplains of the Tama River in central Japan. Seeds collected from a natural habitat were sown in two apparently suitable locations (Tomoda and Ishida sites). Germination, survival, growth, and seed production were subsequently monitored from 1993 through to 1997. The Tomoda site was a gravel bar in floodplains formed by flooding in 1991, while the Ishida site (two plots) was one gravel bar where several plants were growing sparsely and another where a population had become extinct in 1992. Seed cohorts completed their life cycle within 3 years at the Ishida site and within 5 years at the Tomoda site. Monitored parameters at Ishida were substantially lower than those at Tomoda. In addition, estimates of population growth indicated an increase at Tomoda and a rapid decrease at Ishida. However, degradation of habitats seemed to occur at Tomoda over the monitored periods. In view of our results, we conclude that natural germination of about 0.13% is needed for increasing population size. The major factors for decreasing population size are believed to be the lack of safe sites for germination and seedling establishment in old habitats (>10years). Conservation measures are suggested based on these findings.