Keiko Kitamura

Regional Differentiation in Genetic Components for the American Beech, Fagus grandifolia Ehrh., in Relation to Geological History and Mode of Reproduction

Fagus grandifolia, were investigated throughout its geographical range, using allozyme polymorphisms. A total of 1,131 trees from 21 populations were examined for 32 alleles of 10 polymorphic and two monomorphic loci in eight enzyme systems. The mean expected heterozygosity was 0.186, which indicates a relatively high genetic diversity within the populations. The levels of population differentiation were high, as revealed by genetic parameters, i.e., GST=0.168 and FST=0.167. The results of principal component analysis on allele frequencies clearly revealed unique regional patterns of differentiation in genetic components among populations “with” and “without” vegetative regeneration by root suckers. The American beech populations consist of two genetically distinct clusters, one from the Gulf-coastal plain, eastern coastal plain, Piedmont Plateau and Ozark Plateau; and the other from the remaining northern glaciated territories. Populations from the Blue Ridge and Great Smoky Mountains turned out to belong to the latter cluster, which is also characterized by extensive regeneration via root suckers. The consequences of regional differentiation in genetic components are discussed in relation to the postglacial spread from refugia to the current geographic distributions and the mode of reproduction.

The longevity of broadleaf deciduous trees in Northern Hemisphere temperate forests: insights from tree-ring series

Understanding the factors controlling the expression of longevity in trees is still an outstanding challenge for tree biologists and forest ecologists. We gathered tree-ring data and literature for broadleaf deciduous (BD) temperate trees growing in closed-canopy old-growth forests in the Northern Hemisphere to explore the role of geographic patterns, climate variability, and growth rates on longevity. Our pan-continental analysis, covering 32 species from 12 genera, showed that 300-400 years can be considered a baseline threshold for maximum tree lifespan in many temperate deciduous forests. Maximum age varies greatly in relation to environmental features, even within the same species. Tree longevity is generally promoted by reduced growth rates across large genetic differences and environmental gradients. We argue that slower growth rates, and the associated smaller size, provide trees with an advantage against biotic and abiotic disturbance agents, supporting the idea that size, not age, is the main constraint to tree longevity. The oldest trees were living most of their life in subordinate canopy conditions and/or within primary forests in cool temperate environments and outside major storm tracks. Very old trees are thus characterized by slow growth and often live in forests with harsh site conditions and infrequent disturbance events that kill much of the trees. Temperature inversely controls the expression of longevity in mesophilous species (Fagus spp.), but its role in Quercus spp. is more complex and warrants further research in disturbance ecology. Biological, ecological and historical drivers must be considered to understand the constraints imposed to longevity within different forest landscapes.

Estimation of outcrossing rates at small-scale flowering sites of the dwarf bamboo species, Sasa cernua

We estimated the outcrossing rates at small-scale flowering sites of an endemic dwarf bamboo species, Sasa cernua. The multi-locus estimation of the outcrossing rate of the dwarf bamboo population was 0.148 (SD 0.118). Two culms with the highest outcrossing rates had heterozygous genotypes at one locus, but other culms in the locus were homozygotes. Five culms with high outcrossing rates bore 2–17% seeds with homozygous genotypes. Due to predominant selfing, the overall inbreeding coefficient of seeds was high, although it declined in seedlings. This suggests that selection against inbred progenies began early in the establishment process in the natural habitat.

Decline in gene diversity and strong genetic drift in the northward-expanding marginal populations of Fagus crenata

The species distribution of Fagus crenata, or Japanese beech, in the Japanese archipelago shifted northward during phytogeographical changes that occurred during the Pleistocene and Holocene epochs. Presently, the continuous natural distribution of beech reaches north to the Kuromatsunai Depression of Hokkaido Island, Japan. In addition, dozens of marginal patches and isolated individuals north of the continuous distribution have been observed. F. crenata grows remarkably well among these small-scattered northern marginal populations, which must have originated from seeds dispersed beyond the northern limit of the continuous beech forest. It is conceivable that the distribution of F. crenata is still in the process of expanding northward. We investigated the genetic structure of 33 beech populations to evaluate the population gene diversity at the leading northern edge of the range expansion. We analyzed 12 nuclear microsatellite loci in each of the 1,693 individuals. Genetic diversity parameters such as expected heterozygosity and allelic richness were clearly lower in the northernmost populations. We found genetic differentiation in the northernmost distribution range (FST = 0.045, G′ST = 0.242). STRUCTURE analysis revealed that the southwestern continuous populations consisted of homogeneous ancestral clusters. However, northeastern marginal populations consisted of mixtures of highly differentiated clusters with higher levels of genetic drift than found in the continuous populations.

Genetic characteristics reflecting the population size and disturbance regime of Siebold’s beech (Fagus crenata Blume) populations at the northernmost distribution

Abstract To understand the population dynamics of tree populations at the range limit of a species’ range, it is important to determine which population size and disturbance regime are critical to genetic diversity. Siebolds beech (Fagus crenata Blume) is a major canopy tree species of cool-temperate forests in Japan, with the northernmost distribution reaching the Kuromatsunai Depression in southwestern Hokkaido, Japan. We examined the genetic variation and dynamics of three beech forests, with different disturbance history and population attributions in the northernmost population. The Tsubamenosawa (TSU) and Sannosuke (SAN), both natural forest, have historically experienced little artificial disturbance, whereas the Soibetsu (SOI), a secondary forest, was intensively logged, and thus strongly disturbed in the past. In total, 35 alleles were detected among 12 loci, encoding 9 enzyme systems. At TSU, He and RS were 0.141 and 2.280, respectively. At SAN, He and RS were 0.142 and 2.604, respectively. At SOI were 0.182 and 2.628, respectively. Parameters of genetic diversity changed with population size, small isolated population indicated low values. Gene flow distance for low density mature trees in the natural forests was greater than that for high-density secondary forest. However, effective population sizes (Ne) were 34.7, 64.3 and 60.3 in TSU, SAN and SOI, respectively, reflecting differences in the density of mature individuals. The results suggested that the population with the low density of mature trees kept genetic diversity through long distance gene flow. The mature tree density affected the effective population size in the northernmost beech populations.

Distinctions in Fine-Scale Spatial Genetic Structure Between Growth Stages of Picea jezoensis Carr.

Conifers in northern forests, such as fir and spruce, preferably regenerate on coarse woody debris, including fallen logs, stumps, and snags. In northern Japan, the sub-boreal conifer species Picea jezoensis is completely dependent on coarse woody debris for seedling establishment. To understand the fine-scale spatial genetic structure (FSGS) of this species, a 5-ha plot was established in central Hokkaido, and 531 individual trees were categorized into four life-stages (seedling, sapling, juvenile, and mature) on the basis of age and size. The FSGS of the established seedlings and later growth stages was investigated using 11 nuclear simple sequence repeat loci. A STRUCTURE analysis of seedlings and saplings established on fallen logs revealed that genetically related individuals were spatially localized between adjacent logs. We also found a significant FSGS in early life-stages based on a decline in the kinship coefficient calculated between individuals over shorter to longer spatial distances. Furthermore, the estimation of dispersal kernels indicated the frequent occurrence of short-distance seed dispersal. These results indicated that genetically related seedlings and saplings regenerated on the same or nearby fallen logs. In contrast to the results for the early stages, mature-stage trees showed no significant FSGS. We ran a simulation to examine the hypothesis that the FSGS could be eliminated by demographic thinning during life history processes. We calculated values for simulated offspring generated under three sets of conditions; i.e., by removing (i) inbred individuals, (ii) randomly chosen individuals, and (iii) all individuals on the specific fallen logs. However, the results for the FSGS were significant for all simulated data sets. This indicated that inbreeding depression, stochastic loss, or eradication of establishment sites by local disturbances alone could not explain the lack of FSGS among mature-stage trees. Therefore, it is possible that the colonization history of mature trees present on the study site might differ from that of the current offspring.