Spatial distribution patterns of old-growth forest of dioecious tree Torreya nucifera in rocky Gotjawal terrain of Jeju Island, South Korea

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. We generated a database of location, sex, and size (DBH) of T. nucifera trees for each quadrat (160 × 300 m) 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. 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. 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.