Eizi Suzuki

An estimate of the number of tropical tree species

Significance People are fascinated by the amazing diversity of tropical forests and will be surprised to learn that robust estimates of the number of tropical tree species are lacking. We show that there are at least 40,000, but possibly more than 53,000, tree species in the tropics, in contrast to only 124 across temperate Europe. Almost all tropical tree species are restricted to their respective continents, and the Indo-Pacific region appears to be as species-rich as tropical America, with each of these two regions being almost five times as rich in tree species as African tropical forests. Our study shows that most tree species are extremely rare, meaning that they may be under serious risk of extinction at current deforestation rates. The high species richness of tropical forests has long been recognized, yet there remains substantial uncertainty regarding the actual number of tropical tree species. Using a pantropical tree inventory database from closed canopy forests, consisting of 657,630 trees belonging to 11,371 species, we use a fitted value of Fisher’s alpha and an approximate pantropical stem total to estimate the minimum number of tropical forest tree species to fall between ∼40,000 and ∼53,000, i.e., at the high end of previous estimates. Contrary to common assumption, the Indo-Pacific region was found to be as species-rich as the Neotropics, with both regions having a minimum of ∼19,000–25,000 tree species. Continental Africa is relatively depauperate with a minimum of ∼4,500–6,000 tree species. Very few species are shared among the African, American, and the Indo-Pacific regions. We provide a methodological framework for estimating species richness in trees that may help refine species richness estimates of tree-dependent taxa.

Diversity in specific gravity and water content of wood among Bornean tropical rainforest trees

Wood properties were measured for trees in lowland dipterocarp forests in West Kalimantan. In 1993 and 1994, 353 samples of 286 species were collected from trunk base of trees of approximately 5 cm in diameter, and the specific gravities (SG: oven dry weight/fresh volume) and water contents of wood including bark were measured. The SG of each species ranged from 0.21 to 0.84, and the mean ± SD was 0.53 ± 0.13. The wide range of SG suggests that the forest had a high diversity in wood properties. The most dominant and diversified genus in this area was Shorea, and the SG of 15 species varied from 0.21 to 0.71. The range covered SG of pioneer (six Macaranga, 0.29–0.43) and small trees in primary forests (nine Eugenia and 10 Xanthophyllum, 0.55–0.77). The SG average for tree species of secondary forests of 2–6 years old was 0.31. It was significantly smaller than that of primary forests (0.58). In a primary dipterocarp forest plot, light-wood species grew faster in diameter than heavy-wood species. Water content ranged from 0.26 to 0.76. Heavy wood had low water content. Among light-wood species, some (Shorea, Artocarpus) had low water contents and others (Ficus) had high water contents. Some riverine trees also had high water contents. These wood properties appear strongly related to the life history of trees and successional stage.

Habitat differentiation among tree species with small-scale variation of humus depth and topography in a tropical heath forest of Central Kalimantan, Indonesia

Small-scale spatial association of the distribution for 55 abundant tree species with two environmental factors (humus depth and surface microtopography) was examined in two 1-ha plots of a heath (kerangas) forest in Central Kalimantan, Indonesia. More than 80% of the 55 species showed a significant habitat preference in humus depth and/ or relative elevation in at least one plot. In particular, ten species occurring in both plots showed a consistent significant preference for humus depth or relative elevation in the two plots. Using randomization tests, however, only five species significantly associated with humus depth and no species with relative elevation. These results suggest that edaphic and topographic factors, especially humus depth, contribute to determining local spatial distribution and floristic composition of abundant tree species in the forest.

Regeneration process of coniferous forests in northern Hokkaido I.Abies sachalinensis forest andPicea glehnii forest

Regeneration of natural forests was studied in the Nakagawa Experiment Forest of Hokkaido University using age distribution surveys made by the clear felling method. In Plot 1 (30 m × 65 m),Abies sachalinensis dominated the canopy layer but there were also a fewBetula ermanii trees.Sasa senanensis densely covered the forest floor. Most of the canopy trees were from 122 to 195 years old. Seedlings younger than 50 years old ofA. sachalinensis were found on fallen logs and root bases. There were, however, few trees from 50 to 120 years old. The present canopy trees seemed to have regenerated after competitive pressure from old canopy andSasa disappeared 180 years ago. Plot 2 (50 m × 100 m) on serpentinite soil was dominated byPicea glehnii. Sasa kulirensis covered the floor but not as densely asS. senanensis in Plot 1. The ages ofP. glehnii ranged from 1 to 586 years old, and the age distribution ofA. sachalinensis was L-shaped. A small gap in the canopy formed about 290 years ago, and it gradually extended. Conifers regenerated continuously in the extending gap butB. ermanii did not. One hundred thirty years ago, part of Plot 2 was again destroyed andA. sachalinensis andB. ermanii regenerated. Thus, two types of regeneration were found. One regenerated both conifers andBetula after a sudden disturbance of canopy layer or death ofSasa, and the other, under an extending gap, regenerated only conifers.

Vegetation and succession on the Krakatau Islands, Indonesia

An analysis of the vegetation of four islands of the Krakatau group (Rakata Besar, Rakata Kecil, Sertung and Anak Krakatau), Indonesia, was conducted based on our two expeditions of 1982 in centennial commemoration of the great explosion in 1883. Pioneer communities on lava flow, ash fields, beaches, cliffs and on a scoriaceous slope were quite different from each other. Forest communities were grouped into nine types using Morisitas index of similarity, Cλ(W). Two new forest types which have never been described from Krakatau were identified: Timonius compressicaulis forest and Dysoxylum caulostachyum forest. Successional relations among these community types and the Neonauclea calycina forest were discussed from the viewpoint of seed morphology, dispersal, and the development of juvenile generations in the three types of forest. Dysoxylum forest is the most advanced forest community on the Krakatau Islands. The climax forest is supposed to be a tropical monsoon forest similar to that of the volcanic island Panaitan. A successional scheme is proposed. The invasion times of three forest dominants on the three old islands of Krakatau are compared and it is concluded that the difference in vegetation between these islands, the two Rakatas and Sertung, is determined by the earlier invasion of species which could then extend their cover and build up a forest thus preventing later invaders to establish. We are greatly indebted to Dr Fred Hehuwat, Director of the National Institute of Geology and Mining in Bandung, Mr Peter E. Hehanussa, Chief manager for soil and water resource development in the National Institute of Geology and Mining, Dr Soenartono Adisoemarto, Head of the Museum Zoologicum Bogoriense, Dr Kusuwata Kartawinata, Head of the Herbarium Bogoriense, National Institute of Biology, for their kind cooper-ation and use of their facilities. We also thank the staff of the Indonesian Institute of Sciences for their invaluable help in the field and Dr P. J. Grubb who kindly commented on the manu script. The identification of flowering plants was carried out by T. Partomihardjo, who compared samples with specimens in the Herbarium Bogoriense. The identification was counterchecked by Dr M. Hotta, Associate Professor at the Yoshida College, Kyoto University. Dr C. G. G. J. van Steenis taught us a correct name of a species of Bignoniaceae. Pteridophytes were identified by Dr S. Mitsuda, Dept. of Botany, Faculty of Science, Kyoto University, and bryophytes by Dr T. Seki, Dept. of Botany, Faculty of Science, Hiroshima University.

Soils on exposed Sunda Shelf shaped biogeographic patterns in the equatorial forests of Southeast Asia

The marked biogeographic difference between western (Malay Peninsula and Sumatra) and eastern (Borneo) Sundaland is surprising given the long time that these areas have formed a single landmass. A dispersal barrier in the form of a dry savanna corridor during glacial maxima has been proposed to explain this disparity. However, the short duration of these dry savanna conditions make it an unlikely sole cause for the biogeographic pattern. An additional explanation might be related to the coarse sandy soils of central Sundaland. To test these two nonexclusive hypotheses, we performed a floristic cluster analysis based on 111 tree inventories from Peninsular Malaysia, Sumatra, and Borneo. We then identified the indicator genera for clusters that crossed the central Sundaland biogeographic boundary and those that did not cross and tested whether drought and coarse-soil tolerance of the indicator genera differed between them. We found 11 terminal floristic clusters, 10 occurring in Borneo, 5 in Sumatra, and 3 in Peninsular Malaysia. Indicator taxa of clusters that occurred across Sundaland had significantly higher coarse-soil tolerance than did those from clusters that occurred east or west of central Sundaland. For drought tolerance, no such pattern was detected. These results strongly suggest that exposed sandy sea-bed soils acted as a dispersal barrier in central Sundaland. However, we could not confirm the presence of a savanna corridor. This finding makes it clear that proposed biogeographic explanations for plant and animal distributions within Sundaland, including possible migration routes for early humans, need to be reevaluated.

Age structure and regeneration of old growthCryptomeria japonica forests on Yakushima Island

The regeneration process ofCryptomeria japonica forests was studied from surveys of age and initial growth of the coniferous stumps in plots of 2.66 ha in total on Yakushima Island, south Japan. The conifers germinated during particular regeneration periods each of which was shorter than 100 years. The periods repeated themselves several times in each plot, and conifers of each period formed patches. Both tree cutting and natural gap formation of the canopies initiated the regenerations. The saplings that were thought to have germinated before the initiation of the regeneration grew slowly. After the initiation, the saplings that germinated earlier grew more quickly. For the three codominant conifers:Cryptomeria japonica, Tsuga sieboldii, andAbies firma, the shade tolerance was inversely correlated with maximum age. Even the most shade-tolerantA. firma was a gap-dependent species in regeneration periods, and there were no species differences in the colonizing times during the regeneration period. Shade-intolerant long-lived species and shade-tolerant short-lived species coexisted as climax species, not as alternate species in the sere of succession.

Ontogenic change in leaf shape and crown form of a tropical tree,Scaphium macropodum (Sterculiaceae) in Borneo

Crown architecture was analyzed forScaphium macropodum (Sterculiaceae), a common shade-tolerant emergent tree of a tropical rain forest in West Kalimantan, Indonesia. Saplings and poles shorter than 12 m in height had no branches, and gathered their leaves at the ends of the stem. The leaves changed from entire to palmately-parted with increasing tree size. The parted leaves increased the light penetration through the clustered foliage. The size of leaves including the blade and petiole ranged from 22 cm to 147 cm. Because the weight of petiole per blade increased with leaf size, the leaf could not be enlarged infinitely. Taller trees with lateral branches bore small (about 40 cm in length) entire leaves. The light intensity in the forest increased from the ground to about 12 m tall and was nearly constant from 12 m to 18 m. Crown architecture ofS. macropodum adapted to this light environment. The monoaxial trees lower than 12 m could thus increase the amount of light with vertical elongation, and the branched trees higher than 12 m could increase it by means of lateral extension of crown area.

Dynamic steady state of patch-mosaic tree size structure of a mixed dipterocarp forest regulated by local crowding

A patch age- and tree size-structured simulator was applied to demonstrate the landscape dynamics of a lowland mixed dipterocarp forest, using census data over a 3 year interval from two 1 ha plots in northern West Kalimantan, Indonesia (Western Borneo). Tree growth rate and recruitment rate were estimated as functions of tree size and local crowding. The effect of local crowding was assumed to be one-sided through light competition, where the basal area for all trees larger than a target tree inside the circle of 10 m radius around the target was employed as the index of crowding. Estimated parameters were similar between the two plots. Tree mortality was expressed by descending function of tree size with asymptotic mortality for large trees corresponding to the gap formation rate. One parameter specifying the survival of trees at gap formation, which was required for the landscape-level simulation of a shifting-gap mosaic, was left undetermined from plot census data. Through simulation, this parameter was estimated so as to best fit the observed among-patch variation in terms of local basal area. The overall time course of simulation and tree size structure were not sensitive to this parameter, suggesting that one-sided competition along the vertical forest profile is a stronger determinant of average forest structure than among-patch horizontal heterogeneity in this forest. Simulated dynamic steady state successfully reproduced the observed forest architecture in the gap-dynamic landscape. It took about 400 years for a vacant landscape to be replaced by a steady-state architecture of forest. Sensitivity analysis suggests that steady-state basal area and biomass are most sensitive to changing gap formation rate and intrinsic size growth rate.

Mortality and growth of trees in peat-swamp and heath forests in Central Kalimantan after severe drought

Lowland forests in Central Kalimantan, Indonesian Borneo, are endangered by land conversion and the increasing frequency of severe drought. Knowledge of the tolerance of tropical trees to drought is urgent for the management of these lowland habitats. The short-term effects of drought on tree demography (mortality and growth) were investigated in an ever-wet riparian peat-swamp forest and a heath forest on coarse sandy soil after the 1997 El Nino Southern Oscillation (ENSO) event. This drought was unusually severe because little rain fell during the following rainy season. However, forest-wide mortality following the drought (1997–1999) was not critically high in the peat-swamp (6.13% yr−1) or heath (4.26% yr−1) forest. In both forests, standing trees frequently died during the dry season following the drought. The riparian peat-swamp forest was not flooded until 1998, after the prolonged drought in 1997. The hummock–hollow microtopography resulted in differential mortality of peat-swamp trees. On tall hummocks, standing death increased two-fold (4.99% yr−1) during the dry season, whereas uprooting decreased by one-third (0.85% yr−1) during the following rainy season. In contrast, tree growth was not affected by hummock height. Common canopy species were concentrated on tall hummocks and died standing more often than did understory species found in hollows, indicating species-specific mortality after the drought. The large stand basal area relative to the forest-wide growth rate in diameter suggested less resilience to drought by peat-swamp (45.6 m2 ha−1 and 0.0186 ln[cm] yr−1) than heath (27.9 m2 ha−1 and 0.0232 ln[cm] yr−1) forest. A single severe drought did not cause dramatic changes in the peat-swamp and heath forests; however, an increasing frequency of droughts similar in severity to that of the 1997 ENSO event may have the potential to alter the community structure and dynamics, leading to a consistent decline in Bornean lowland forests.