• Journal of Ecology and Environment
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• v.41 no.8
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• pp.223-234
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• 2017

Background: Spatial structure of plants in a population reflects complex interactions of ecological and evolutionary processes. For dioecious plants, differences in reproduction cost between sexes and sizes might affect their spatial distribution. Abiotic heterogeneity may also affect adaptation activities, and result in a unique spatial structure of the population. Thus, we examined sex- and size-related spatial distributions of old-growth forest of dioecious tree Torreya nucifera in extremely heterogeneous Gotjawal terrain of Jeju Island, South Korea. Methods: We generated a database of location, sex, and size (DBH) of T. nucifera trees for each quadrat ($160{\times}300m$) in each of the three sites previously defined (quadrat A, B, C in Site I, II, and III, respectively). T. nucifera trees were categorized into eight groups based on sex (males vs. females), size (small vs. large trees), and sex by size (small vs. large males, and small vs. large females) for spatial point pattern analysis. Univariate and bivariate spatial analyses were conducted. Results: Univariate spatial analysis showed that spatial patterns of T. nucifera trees differed among the three quadrats. In quadrat A, individual trees showed random distribution at all scales regardless of sex and size groups. When assessing univariate patterns for sex by size groups in quadrat B, small males and small females were distributed randomly at all scales whereas large males and large females were clumped. All groups in quadrat C were clustered at short distances but the pattern changed as distance was increased. Bivariate spatial analyses testing the association between sex and size groups showed that spatial segregation occurred only in quadrat C. Males and females were spatially independent at all scales. However, after controlling for size, males and females were spatially separated. Conclusions: Diverse spatial patterns of T. nucifera trees across the three sites within the Torreya Forest imply that adaptive explanations are not sufficient for understanding spatial structure in this old-growth forest. If so, the role of Gotjawal terrain in terms of creating extremely diverse microhabitats and subsequently stochastic processes of survival and mortality of trees, both of which ultimately determine spatial patterns, needs to be further examined.

• Journal of Ecology and Environment
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• v.35 no.2
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• pp.111-122
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• 2012

The sex ratio and spatial structure of different sexes are major components that affect the reproductive success and population persistence of dioecious plants. The differential reproductive costs between male and female plants are often believed to cause a biased sex ratio and spatial segregation of the sexes through slower growth and/or lower female survivorship. In this study, we examined the sex ratio and spatial structure of one population of $Torreya$ $nucifera$ trees in Jeju Island, Korea. We also tested the effects of the current tending actions in relation to tree vitality. At the population level, the sex ratio of the 2,861 trees was significantly biased toward males; however, it also showed considerable variation among different diameter at breast height classes and across habitats according to terrain level (from upper to lower). In 1999, before tree management (tending) began, among the ecological traits examined, only climber coverage correlated with tree vitality. Intensive tending such as climber removal since 1999 clearly enhanced the vitality of the majority of trees, but its effects were more conspicuous in medium-sized trees than in small ones, in upper terrain trees than those in other terrains, and in females than in males. Both male tree domination in small and large trees and tending effects on females are likely to reflect the effects of female reproductive costs regarding growth and/or survivorship. Spatial segregation between males and females was not observed in $T.$ $nucifera$. Habitat heterogeneity created by the forest's rocky ground and its implications regarding sex ratios and spatial structure require further studies.

• Journal of Ecology and Environment
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• v.43 no.3
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• pp.320-331
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• 2019

Background: Dioecious plant species having both male and female plants have been investigated regarding sex-related characteristics such as sex ratio, sex-differential resource requirements, and spatial segregation of the sexes. Habitat loss and fragmentation are major threats to the survival of plant populations, but dioecious species are particularly more prone to such habitat degradation than non-dioecious species because of their dimorphic sexual system. We examined the sex-related demographics of two Ilex cornuta populations being different regarding land use history. Methods: During 2016-2017, we examined I. cornuta trees with a basal diameter ${\geq}1.5cm$ in the Yongsu-ri population (YS population) and the Gotjawal Provincial Park population (GP population). Plant sex (male, female, or unsexed) was identified. The tree size (basal diameter and height of the main stem), clonal production (the ramet numbers per genet), and vitality for each clone were measured. The associations between population, sex, tree size, clonal production, and vitality were examined using ANOVAs and contingency table analyses. Finally, point pattern analyses using O-ring statistics were conducted to assess spatial patterns. Results: Upon excluding unsexed trees, the YS population with 74 trees was significantly male-biased (0.66), while the GP population with only 26 trees had a 1:1 sex ratio. In both populations, males and females did not differ in tree size. Although the mean number of ramets differed significantly between populations, females tended to produce more ramets than males. The proportion of weak trees was significantly higher in the YS than in the GP population. Neither population showed evidence of spatial segregation of the sexes. Conclusions: The two populations of dioecious I. cornuta are characterized by the small number of trees and relatively high frequencies of non-reproductive trees. Both indicate that these populations are quite susceptible to environmental and genetic stochasticity. On the other hand, the differences between populations in sex ratio, clonal production, and vitality suggest that conservation efforts for I. cornuta need to be population-specific. In order to help recover and enable this vulnerable species to persist, it is necessary to find ways to enhance their sexual reproduction and simultaneously reduce habitat disturbances due to anthropogenic activities.