Management and micro-scale landform determine the ground flora of secondary woodlands and their verges in the Tama Hills of Tokyo, Japan
We investigated the influence of management and micro-scale landform on the species composition and richness of ground flora in secondary woodlands and their verges next to paddy fields in the Tama Hills, Tokyo, Japan. Sites representing various micro-scale landforms and different management regimes were sampled. The results of Detrended Correspondence Analysis and Indicator Species Analysis showed that there was no significant difference between management regimes on species composition in head hollow sites. However, species composition was unique in other landform types having the same management regimes. Micro-scale landform units had less effect on species composition than management regime on crest slopes and side slopes of secondary woodlands. A large variation in species composition of verges, all on the lowermost side slopes, was found within sites, but the composition was different when compared with other sites. We recognized five habitat types associated with species composition that resulted from the combined effects of landform and management regimes. Among habitat types, the verges and the well-managed woodlands had high species richness. Management practices within woodlands have enhanced the species richness on crest slopes and side slopes. High beta diversity of ground flora could be due to the variation in micro-scale landform along soil moisture regimes. Intensive management involving clear-cutting on the lower side slopes (an ecotone of mesic and wet environments) accompanied by paddy cultivation on valley bottoms enhanced the diversity of herbaceous vegetation in the verges.