Myong Gi Chung

SPATIAL GENETIC STRUCTURE OF CLONAL AND SEXUAL REPRODUCTION IN POPULATIONS OF ADENOPHORA GRANDIFLORA (CAMPANULACEAE)

The spatial distribution of clonal versus sexual reproduction in plant populations should generally have differing effects on the levels of biparental inbreeding and the apparent selfing rate, produced via mating by proximity through limited pollen dispersal. We used allozyme loci, join‐count statistics, and Morans spatial autocorrelation statistics to separate the spatial genetic structure caused by clonal reproduction from that maintained in sexually reproduced individuals in two populations of Adenophora grandiflora, a perennial herb. Join‐count statistics showed that there were statistically significant clustering of clonal genotypes within distances less than 4 m. Both the entire populations and the sets of sexually reproduced individuals exhibited significant spatial autocorrelation at less than about 12 m, and the sexually reproduced individuals are substantially structured in an isolation‐by‐distance manner, consistent with a neighborhood size of about 50.

Genetic structure of age classes in Camellia japonica (Theaceae)

Abstract.— Camellia japonica L. (Theaceae), an insect‐ and bird‐pollinated, broad‐leaved evergreen tree, is widely distributed in Japan and the southern Korean peninsula. The species has a relatively even age distribution within populations, which may influence the spatial genetic structure of different age classes relative to species with typical L‐shaped age distributions. To determine whether the internal spatial genetic structure found in seedlings and young individuals carries over into adults, we used allozyme loci, F‐statistics, spatial autocorrelation statistics (Morans I), and coancestry measures to examine changes in genetic structure among seven age classes in a population (60‐m × 100‐m area) in southern Korea. In seedlings, weak but significant positive values of Morans I‐statistics and coancestry measures were found for distances less than 14 m, which is consistent with a mechanism of limited seed dispersal combined with overlapping seed shadows. This spatial structure, however, dissipates in older age classes, and in adults genetic variation has an essentially random spatial distribution. Morisitas index of dispersion of individuals in each age class showed that seedlings and juveniles are more highly clustered than are older individuals. These results suggest that self‐thinning changes the spatial relationships of individuals, and thus genotypes. A multilocus estimate of FST (0.008) shows a small but statistically significant difference in allele frequencies among age classes. In summary, intrapopulation genetic structure within and among age classes of C. japonica was significant but weak. Despite presumably limited seed dispersal, weak spatial genetic structure in juveniles suggests overlapping seed shadows followed by self‐thinning during recruitment. The present study also demonstrates that studies of spatial genetic structure focusing on limited numbers of generations may not be sufficient to reveal the entire picture of genetic structure in populations with overlapping generations.

Spatial genetic structure in populations of the terrestrial orchid Cephalanthera longibracteata (Orchidaceae)

Orchid seeds are unusual for being the smallest among flowering plants. These dust-like seeds are wind-borne and, thus, would seem to have the potential for long-distance dispersal (a common perception); this perception has led to a prediction of near-random spatial genetic structure within orchid populations. Mathematical models (e.g., simple ballistic model) for wind-dispersed seeds and wind-tunnel experiments, in contrast, indicate that most seeds of orchids should fall close to the maternal plant (<6 m), supporting a prediction of significant fine-scale genetic structure within populations. In reality we do not know much about seed dispersion in orchids. To determine which of these two predictions is more appropriate, Wrights F statistics and spatial autocorrelation analysis were used to examine the genetic structure within two adult populations of the terrestrial orchid Cephalanthera longibracteata (Orchidaceae) in southern Korea. In results comparable to those of other self-compatible, mixed-mating plant species, C. longibracteata populations exhibited low levels of genetic diversity (mean H(e) = 0.036) and a significant excess of homozygosity (mean F(IS) = 0.330), consistent with substantial inbreeding via selfing and/or mating among close relatives in a spatially structured population. Spatial autocorrelation analysis revealed significant positive genetic correlations among plants located <10 m, with relatedness at <3 m comparable to that expected for half sibs and first cousins. This genetic structure supports the prediction that the majority of seed dispersal occurs over distances of less than 10 m and is responsible for generating substantial overlap in seed shadows within C. longibracteata populations.

Spatial genetic structure of allozyme polymorphisms within populations of Pinus Strobus (Pinaceae).

The population structure of genetic variation for four allozyme loci was investigated for two populations (one old growth, OG, and the other logged, SS) of eastern white pine (Pinus strobus). Both seedlings and reproductive adults were studied for both study populations. Spatial autocorrelation statistics were used to examine the distribution of allozyme polymorphisms. The spatial genetic structure in adults of population OG indicated that individual genotypes were distributed in a structured, isolation-by-distance manner, consistent with observed levels of pollen and seed dispersal. In contrast, adult genotypes in population SS were nearly randomly distributed, probably as a result of logging. Nonetheless, spatial structuring of genotypes of seedlings occurred at both sites, indicating the power of limited seed flow, as well as temporal Wahlund effects, to create structure. None of four loci in both seedling populations showed a significant departure from Hardy-Weinberg proportions, whereas one and two significant deviations were found for loci in the two respective adult populations. These departures may be attributed to episodic reproductive events.

Spatial genetic structure in a Neolitsea sericea population (Lauraceae)

Neolitsea sericea (Bl.) Koidz. (Lauraceae) is a dioecious, insect-pollinated, and broad-leaved evergreen tree with bird-dispersed seeds. We used allozyme loci, F-statistics, and spatial autocorrelation statistics (Moran’s I) to examine the changes in genetic structure among five age classes within a study population (60 m × 100 m area) in southern Korea. No significant differences in expected heterozygosity were found among the age classes. The mean F-values averaged over loci were similar among age classes and showed overall conformance of heterozygosities with Hardy–Weinberg proportions. Differences in allelic frequencies among age classes were small (mean GST=0.012), and statistically significant only for one locus (Pgd-2). The mean Moran’s I-values for each of five age classes indicated essentially random spatial distribution. The homogeneity of genetic structure and genetic diversity among the five age classes may reflect the occurrence of similar reproductive events, year after year. The results may reflect the attractive red drupes of N. sericea in that they cause various frugivorous birds to disperse the seed long distances and independently, which in turn may help N. sericea maintain higher levels of genetic diversity within populations.

Clonal and spatial genetic structure in Eurya emarginata (Theaceae)

Eurya emarginata (Thunb.) Makino (Theaceae) is a dioecious, insect-pollinated tree, which combines sexual reproduction and clonal spread. It is narrowly distributed in coastal areas from southern China, along southern Korea, and extending to central and southern Japan. We used allozyme loci and spatial autocorrelation statistics to examine the clonal structure and the spatial distribution of allozyme polymorphisms of sexually reproduced individuals in a study population (70 m × 120 m area) in Korea. The population maintains moderate levels of allozyme variability (mean He=0.133), and multilocus genotypic diversity is high (mean DG=0.992). The frequency of clones was 0.377 (90/239), and the mean clone-pair distance was very large (35.06 m ± 1.91 m [SE]), indicating that clones can form at long distances away from their ancestors. In addition, it was found that genetic patch width was at least 24 m. However, join-count statistics for the total number of unlike joins showed that, contrary to expectation for such species, clonal reproduction does not contribute substantially to genetic isolation by distance neither among the sexually reproduced individuals nor the whole population. In contrast, limits to seed and pollen dispersal create substantial levels of genetic structure.

Spatial genetic structure in populations of the terrestrial orchid Orchis cyclochila (Orchidaceae)

Orchid seeds are minute, dust-like, wind-borne and, thus, would seem to have the potential for long-distance dispersal. Based on this perception, one may predict near-random spatial genetic structure within orchid populations. In reality we do not know much about seed dispersal in orchids and the few empirical studies of fine-scale genetic structure have revealed significant genetic structure at short distances (< 5m), suggesting that most seeds of orchids fall close to the maternal plant. To obtain more empirical data on dispersal, Ripley’s L(d)-statistics, spatial autocorrelation analyses (coancestry, fij analyses) and Wright’s F statistics were used to examine the distribution of individuals and the genetic structure within two populations of the terrestrial orchid Orchis cyclochila in southern Korea. High levels of genetic diversity (He = 0.210) and low between-population variation were found (FST = 0.030). Ripley’s L(d)-statistics indicated significant aggregation of individuals, and patterns varied depending on populations. Spatial autocorrelation analysis revealed significant positive genetic correlations among individuals located <1 m, with mean fij values expected for half sibs. This genetic structure suggests that many seeds fall in the immediate vicinity of the maternal plant. The finding of significant fine-scale genetic structure, however, does not have to preclude the potential for the long distance dispersal of seeds. Both the existence of fine-scale genetic structure and low FST are consistent with a leptokurtic distribution of seed dispersal distances with a very flat tail.

Patterns of hybridization and population genetic structure in the terrestrial orchids Liparis kumokiri and Liparis makinoana (Orchidaceae) in sympatric populations

We investigated the potential for gene flow and genetic assimilation via hybridization between common and rare species of the terrestrial orchid genus Liparis, focusing specifically on sympatric and allopatric populations of the common Liparis kumokiri and the rare Liparis makinoana. We utilized analyses of genetic diversity, morphology, and the spatial distributions of individuals and genotypes to quantify the dynamics of interspecific gene flow at within‐ and among‐population scales. High levels of allozyme genetic diversity (HE) were found in populations of the rare L. makinoana (0.317), whereas the common L. kumokiri (N = 1744 from 14 populations) revealed a complete lack of variation. This contrast may reflect different breeding systems and associated rates of genetic drift (L. makinoana is self‐incompatible, whereas L. kumokiri is self‐compatible). At the two known sympatric sites, individuals were found that recombined parental phenotypes, possessing floral characteristics of L. kumokiri and vegetative characteristics of L. makinoana. These putative hybrids were the only individuals found segregating alleles diagnostic of both parental species. Analysis of these individuals indicated that hybrid genotypes were skewed towards L. kumokiri and later generation recombinants of L. kumokiri at both sympatric sites. Furthermore, Ripleys bivariate L(r) statistics revealed that at one site these hybrids are strongly spatially clustered with L. kumokiri. Nonetheless, the relatively low frequency of hybrids, absence of ongoing hybridization (no F1s or first generation backcrossess), and strong genetic differentiation between morphologically ‘pure’ parental populations at sympatric sites (FST = 0.708–0.816) indicates that hybridization was not an important bridge for gene flow. The results from these two species suggest that natural hybridization has not played an important role in the diversification of Liparis, but instead support the view that genetic drift and limited gene flow are primarily responsible for speciation in Liparis. Based on genetic data and current status of the species, implications of the research for conservation are considered to provide guidelines for appropriate conservation and management strategies.

Landscape-level spatial genetic structure in Quercus acutissima (Fagaceae)

Quercus acutissima (Fagaceae), a deciduous broad-leaved tree, is an important forest element in hillsides of South Korea. We used allozyme loci, Wrights F statistics, and multilocus spatial autocorrelation statistics to examine the distribution of genetic diversity within and among three local populations and the spatial genetic structure at a landscape scale (15 ha, 250 × 600 m) on Oenaro Island, South Korea. Levels of genetic diversity in Q. acutissima populations were comparable to mean values for other oak species. A moderate but significant deficit of heterozygotes (mean F(IS) = 0.069) was detected within local populations and low but significant differentiation was observed among populations (F(ST) = 0.010). Spatial autocorrelation analyses revealed little evidence of significant genetic structure at spatial scales of 100-120 m. The failure to detect genetic structure within populations may be due to intraspecific competition or random mortality among saplings, resulting in extensive thinning within maternal half-sib groups. Alternatively, low genetic differentiation at the landscape scale indicates substantial gene flow among local populations. Although wind-borne pollen may be the primary source of gene flow in Q. acutissima, these results suggest that acorn movement by animals may be more extensive than previously anticipated. Comparison of these genetic data for Oenaro Island with a disturbed isolated inland population suggests that population-to-population differences in internal genetic structure may be influenced by local variation in regeneration environment (e.g., disturbance).

Temporal aspects of the fine-scale genetic structure in a population of Cinnamomum insularimontanum (Lauraceae)

Cinnamomum insularimontanum Hayata (Lauraceae) is an insect-pollinated, broad-leaved evergreen tree with bird-dispersed seeds. We used allozyme loci, Wrights fixation index, spatial autocorrelation statistics (Morans I), and coancestry measures to examine changes in genetic structure among four age-classes within a recently founded study population (60×100 m area) in southern Korea. There were no significant differences in expected heterozygosity among age classes. However, significant genetic differentiation among age classes was detected (P<0.0001). Fixation indices within age classes showed significant deficits of observed heterozygosity, which may be caused by partial selfing. The homogeneity of genetic structure among four age-classes may reflect similar spatial patterns of seed immigration from surrounding populations occurring year after year. Finally, the average Morans I and coancestry estimates indicated essentially random spatial distributions of alleles for each of the four age-classes and between seedlings and 2–4 year juveniles vs adult trees. These findings are very similar to those observed in the same study area for another member of the Lauraceae, Neolitsea sericea, which has a very similar life history and ecological characteristics (ie, bird-dispersed fruits, insect pollination, and a similar age structure). Together, these results suggest that the fleshy drupes of lauraceous species represent an adaptation to aid in the independent dispersal of seed by birds, which in turn may increase the genetic diversity of founders colonizing new habitats.