The Potential Impact of Small-Scale Physical Disturbance on Seedlings in a Papuan Rainforest1

Physical disturbance, such as falling debris, is an important cause of mortality to rain forest seedlings. I used artificial seedlings to quantify physical disturbance in a rain forest at the Crater Mountain Biological Research Station in Papua New Guinea. Single (N= 418) and clustered artificial seedlings (N= 428) were placed on eight transects spanning varied topography to assess the area's disturbance regime, to test if ground slope affected disturbance, and to test if clustering affected the incidence of damage. Two transects showed a significant departure from random in the spatial distribution of damaged seedlings within the transect, suggesting that some areas of experience more disturbance than others. Artificial seedlings on level ground experienced damage less often than artificial seedlings on slopes; disturbance increased with ground slope. Artificial seedlings in dry washes experienced the greatest probability of damage. Artificial seedlings in clusters experienced significantly less damage than single ones, suggesting clustered seedlings might be less vulnerable to damage than lone seedlings. The disturbance rate at this site (34.7% damaged in one year) was markedly lower than that measured at the La Selva Biological Reserve in Costa Rica (82.4%).     

The Role of Arboreal Seed Dispersal Groups on the Seed Rain of a Lowland Tropical Forest1

Most tropical plants produce fleshy fruits that are dispersed primarily by vertebrate frugivores. Behavioral disparities among vertebrate seed dispersers could influence patterns of seed distribution and thus forest structure. This study investigated the relative importance of arboreal seed dispersers and seed predators on the initial stage of forest organization–seed deposition. We asked the following questions: (1) To what degree do arboreal seed dispersers influence the species richness and abundance of the seed rain? and (2) Based on the plant species and strata of the forest for which they provide dispersal services, do arboreal seed dispersers represent similar or distinct functional groups? To answer these questions, seed rain was sampled for 12 months in the Dja Reserve, Cameroon. Seed traps representing five percent of the crown area were erected below the canopies of 90 trees belonging to nine focal tree species: 3 dispersed by monkeys, 3 dispersed by large frugivorous birds, and 3 wind‐dispersed species. Seeds disseminated by arboreal seed dispersers accounted for ca 12 percent of the seeds and 68 percent of the seed species identified in seed traps. Monkeys dispersed more than twice the number of seed species than large frugivorous birds, but birds dispersed more individual seeds. We identified two distinct functional dispersal groups, one composed of large frugivorous birds and one composed of monkeys, drop dispersers, and seed predators. These groups dispersed plants found in different canopy strata and exhibited low overlap in the seed species they disseminated. We conclude it is unlikely that seed dispersal services provided by monkeys could be compensated for by frugivorous birds in the event of their extirpation from Afrotropical forests.     

Tropical Rain Forest Recovery from Cyclone Damage and Fire in Samoa1

In 1990 and 1991, Samoa was struck by two cyclones, Ofa and Val. In the Tafua Rain Forest Preserve on the island of Savai'i, one part of the forest also burned after the first cyclone. Here we report on patterns of regeneration and changes in tree species composition in the Tafua lowland rain forest after five years of recovery from cyclone and fire disturbance. In the unburned area, tree canopy cover increased from 27 percent after the last cyclone to 58 percent, and in the burned area from below 12 to 49 percent. Nine of the ten most common tree species decreased in relative abundance in the entire forest after the last cyclone. One fast growing pioneer species, Macaranga harveyana now makes up 42 percent of the total number of trees (>5 cm DBH) in the unburned area and 86 percent in the burned area. Large interspecific differences occur in size distribution and there are at least four distinguishable regeneration patterns, which may be related to shade tolerance. Mean number of species per plot was generally higher in the unburned area than in the burned area, while the Shannon evenness index was higher in the unburned than in the burned area only for trees above 1 cm DBH. Species with fruits known to be fed upon by birds and/or bats generally made up a larger proportion of all trees in the burned than in the unburned area. In contrast to other studies of post‐cyclone regeneration, in which recovery is often rapid due to resprouting of trees, recovery in the Tafua forest was a slow process with regeneration more dependent on vertebrate seed dispersal than on resprouting.     

海南霸王岭热带山地雨林森林循环与树种多样性动态

通过对海南岛霸王岭热带山地雨林的调查 ,研究了热带山地雨林树种多样性特征随森林循环的动态变化规律。结果表明 :( 1 )热带山地雨林森林循环不同阶段斑块在森林景观中所占的面积比例分别是 :林隙阶段 ( G)占 38.5 0 % ,建立阶段 ( B)占 2 8.5 0 % ,成熟阶段 ( M)占 2 7.0 0 % ,衰退阶段 ( D)占 6 .0 0 %。 ( 2 )热带山地雨林中乔木树种的密度随森林循环的变化趋势是由 G→B→M呈现出逐渐增加的趋势 ,以成熟阶段达到最大 ,而到衰退阶段又趋于下降。灌木树种则表现出 G阶段斑块的密度最大 ,B阶段的最小 ,从 B到 M有所增加 ,到 D又稍有下降。 ( 3)热带山地雨林中不同高度级和不同径级的树木的密度在森林循环的不同阶段表现出不同的增减趋势 ,其随森林循环过程呈现出的动态变化可能与不同阶段斑块内的空间、环境及物种生物学特性有关。 ( 4 )热带山地雨林中树木的平均胸径、平均高、平均胸高断面积、平均单株材积随森林循环过程呈现出不断增加的趋势 ,其中平均胸径和平均高随森林循环的变化较为平缓 ,而平均胸高断面积和平均单株材积之变化较为陡急。 ( 5 )热带山地雨林森林循环不同阶段的物种多样性指数不同 ,其中 G和 B阶段的物种丰富度和多样性指数值较接近 ,M阶段的物种丰富度达到最大 ,D阶段则最小。     

North—South Patterns within Arboreal Ant Assemblages from Rain Forests in Eastern Australia1

This paper describes the ant assemblages sampled from rain forest canopies ranging from southern Victoria through to Cape York Peninsula, Australia, and also in Brunei. Specifically, it examines the influence of decreasing latitude and variations in elevation on the character, richness, and abundance of the arboreal rain forest ant fauna, and also the relative contribution of ants to the total arthropod community. The sites that were examined included: cool temperate Nothofagus cunninghamii forest from a range of locations in Victoria; cool temperate N. moorei forest at both Werrikimbe and Styx River, New South Wales; notophyll vine forest in Lamington National Park, southeast Queensland; high elevation notophyll vine forest in Eungella National Park, central Queensland; complex notophyll vine forest at Robson Creek, Atherton Tablelands, north Queensland; complex mesophyll vine forest at Cape Tribulation, north Queensland; and mixed dipterocarp forest in Brunei. Although these sites represent a gradient increasingly tropical in character, botanically speaking, Eungella is less tropical than Lamington because of its high elevation. All samples were obtained by fogging the canopy with a rapid‐knockdown pyrethrin pesticide. In all cases, circular funnels were suspended beneath the foliage of individual trees or small plots of mixed canopy. Arthropods were collected four hours after fogging. Following ordinal sorting, ants were identified and counted to morphospecies level. The resulting catch were then standardized across sites as numbers caught per 0.5 m² sampling funnel. Generic and species richness were higher at the lowland tropical Cape Tribulation sites than at the sites to the south and was comparable with values in the Brunei site. Species richness was negatively correlated with latitude and elevation. Within the Australian rain forest, the lowland/highland break appears to be the strongest predictor of ant relative abundance, with a weaker latitudinal relationship superimposed.     

Influence of Edge Exposure on Tree Seedling Species Recruitment in Tropical Rain Forest Fragments1

Edge creation has a pronounced influence on the understory vegetation, but the effects of edges on seedling species recruitment are still poorly understood. In Central Amazonia, 9–19 years after fragmentation, we recorded species richness and net seedling recruitment rate in 1 ha blocks exposed to none, one, or multiple edges within forest fragments. One‐hectare blocks were located in the center (no edge), the edge (one edge), the corners (two edges) of 10 and 100 ha fragments, and in a 1 ha fragment (four edges). In 1991, we counted all tree seedlings 5–100 cm tall found within permanent 1 m² plots located within the 1 ha blocks. In May 1993, we manually removed all seedlings that were smaller than 1 m tall from the permanent plots. Six years and five months later (October 1999), all new seedlings recruited into the plots were counted and classified into distinct morphospecies. Species richness of recruited seedlings, scaled by total seedling density, declined from the center to the edge, the corner blocks, and then to the 1 ha fragment. Overall, the four‐edged, 1 ha fragment had the poorest species richness and the non‐edged 100 ha central block the highest. The total number of recruited individuals was 40 percent less than that previously present, with the 100 ha corner having the lowest recruitment. Pairwise comparisons showed that species similarity was related to edge number for the 100 and 1 ha fragments. Species rank/abundance curves showed that a subset of species was common in all blocks within the fragments, and that the 100 ha center held more rare species than any other 1 ha block. This study demonstrated that, in a given fragment patch, the number of tree seedling species recruited varied inversely with the number of edges.     

Ants Induce Domatia in a Rain Forest Tree (Vochysia vismiaefolia)1

In Amazonian rain forest trees of Vochysia vismiaefolia (Vochysiaceae), ants were found to induce twig structures that resembled classical ant domatia. This phenomenon is novel for ant‐plants, which commonly develop domatia without the activity of ants. Eight species of ants were recorded inside the domatia of six individual trees, but domatia were most numerous and characteristic when induced and inhabited by an undescribed species of Pseudomyrmex on two trees. To investigate the mechanism of domatium growth, we drilled holes into young twigs: the expansion of the twig diameter surrounding the holes was significantly accelerated, comparable to domatia formation. Domatia induction is discussed as a putative step in the evolution of ant‐plants.     

Forest Succession in Tropical Hardwood Hammocks of the Florida Keys: Effects of Direct Mortality from Hurricane Andrew1

A tree species replacement sequence for dry broadleaved forests (tropical hardwood hammocks) in the upper Florida Keys was inferred from species abundances in stands abandoned from agriculture or other anthropogenic acitivities at different times in the past. Stands were sampled soon after Hurricane Andrew, with live and hurricane‐killed trees recorded separately; thus it was also possible to assess the immediate effect of Hurricane Andrew on stand successional status. We used weighted averaging regression to calculate successional age optima and tolerances for all species, based on the species composition of the pre‐hurricane stands. Then we used weighted averaging calibration to calculate and compare inferred successional ages for stands based on (1) the species composition of the pre‐hurricane stands and (2) the hurricane‐killed species assemblages. Species characteristic of the earliest stages of post‐agricultural stand development remains a significant component of the forest for many years, but are gradually replaced by taxa not present, even as seedlings, during the first few decades. This compositional sequence of a century or more is characterized by the replacement of deciduous by evergreen species, which is hypothesized to be driven by increasing moisture storage capacity in the young organic soils. Mortality from Hurricane Andrew was concentrated among early‐successional species, thus tending to amplify the long‐term trend in species composition.     

Post‐Establishment Seedling Success in the Brazilian Cerrado: A Comparison of Savanna and Forest Species1

Due to frequent fire, low nutrient availability, and prolonged drought, tropical savanna is a stressful environment for the survival and growth of woody plant seedlings. To understand why forest species do not succeed in this environment while savanna species are able to persist, the effects of fire and woody cover on seedlings of these two functional groups were investigated in the Brazilian Cerrado. Seedlings were established in experimental plots under three densities of woody cover, in sites protected from fire and sites to be subjected to fire. There was a clear difference in the ability of savanna and forest species to survive fire. None of the three forest species were able to survive fire during the first two years of life, whereas eight of the nine savanna species were able to resprout following fire. The small seed size of the ninth savanna species, Miconia albicans, predisposed its seedlings to be sensitive to fire, because there was a strong positive correlation between seed size and survivorship. Savanna species were less dependent on woody cover than were forest species, which exhibited higher growth and survival under tree canopies than in open grassland. The low rates of establishment and survival of forest trees in savanna, combined with high sensitivity to fire, appear sufficient to prevent the expansion of forest into savanna under current fire regimes in the Cerrado.     

Home Range Size of Sympatric Squirrel Species Inhabiting a Lowland Dipterocarp Forest in Malaysia1

Home range sizes and spatial overlap of four sympatric squirrel species were investigated in a lowland dipterocarp forest in Malaysia using a radio‐tracking method. The population density of Callosciurus caniceps was highest and C. notatus was next highest, while C. nigrovittatus and Lariscus insignis were scarce. C. caniceps was larger than C. nigrovittatus and C. notatus while L. insignis was extremely small. For females, home range size was smaller in L. insignis than Callosciurus spp., which may support the body weight hypothesis: larger species have larger home ranges. Among the three Callosciurus species, female C. caniceps had the smallest home range. These differences were accounted for by habitat characteristics rather than by density or body weight; C. caniceps was dominant in bushy areas and used crowded small trees while C. notatus and C. nigrovittatus used large trees in the forest. In this study, home range size did not change seasonally; this differs from studies in temperate regions, possibly because food availability is much less variable among seasons in tropical rain forest. Home range overlap among heterospecific individuals was common but different species seemed to partition space by using different vertical levels of the forest. Consequently, the home range size and spatial overlap of sympatric squirrel species may be affected by habitat diversity in tropical rain forest.     

Mammalian Diet and Broad Hunting Strategy of the Dingo (Canis familiaris dingo) in the Wet Tropical Rain Forests of Northeastern Australia1

The diet of dingoes (Canis familiaris dingo) in the Australian Wet Tropics was examined by analyzing 383 dingo scats collected throughout the region for the presence of mammal prey remains. The scats yielded 29 native and 4 introduced mammal prey species from 14 families. The most important species in terms of percentage occurrence in the scats were Melomys cervinipes (22.2%), hoodon macrourus (17.0%), Perameles nasuta (12.5%), and Thylogale stigmatica (12.5%). The most important families were Muridae (37.1%), Peramelidae (29.5%), and Macropodidae (25.8%). Examination of small‐scale habitat preferences revealed species that preferentially use the forest edge ranked significantly higher in the diet than those that do not, and species that are terrestrial ranked higher in the diet than those that are arboreal. Relative abundance was also a significant factor in the ranked dietary occurrence of each species, with abundant species ranked significantly higher than those that are less abundant. These results suggest that dingoes in the Australian Wet Tropics are opportunistic predators of a wide variety of mammal species, with abundant terrestrial and forest edge‐dwelling taxa the most susceptible to predation.     

Edge Effects and Seedling Bank Depletion: The Role Played by the Early Successional Palm Attalea oleifera (Arecaceae) in the Atlantic Forest

In this study, we examined the impacts of Attalea oleifera on the structure of seedling bank and discuss potential mechanisms of palm influence. Seed rain, seedling bank, and palm leaf fall were assessed beneath the canopy and in the vicinity of 16 adult palms across the edges of a large fragment (3500 ha) of the Atlantic forest. Moreover, we examined A. oleifera impacts on seed germination and seedling mortality by experimentally submitting seeds and seedlings to prolonged palm-leaf covering. As expected, seedling bank beneath the adults exhibited reduced abundance and species richness at local and habitat scale. Small to large seeds (3.1–30 mm) were underrepresented in the seed rain below adults palms, while experimental leaf covering drastically reduced both seed germination and seedling survivorship. A. oleifera leaf fall occurred over the whole year (3.6±2.7 leaves/individual/yr), which resulted in deep leaf litter mounds (10.7±9.2 cm). Finally, adult palm density (21.6±11.9 individuals/ha) correlated negatively with seedling density across Attalea clusters. Our results suggest that A. oleifera exerts negative effects on the seedling bank by reducing seedling abundance and richness as a consequence of two complementary mechanisms: impoverished and size-biased seed rain plus reduced seed germination and increased seedling mortality due to prolonged covering by fallen leaves.     

Hyper‐abundance of Native Wild Pigs (Sus scrofa) in a Lowland Dipterocarp Rain Forest of Peninsular Malaysia1

This study reports extraordinarily high density estimates for the wild pig (Sus scrofa) from an aseasonal tropical forest site within the species'native range. At Pasoh Forest Reserve, a 2500 ha area of lowland dipterocarp rain forest in Peninsular Malaysia, line transects were used to estimate pig density from May to October in 1996 and 1998. In 1996, 44 sightings of S. scrofa consisting of 166 individuals were recorded along 81 km of transects. In 1998, 39 sightings documented 129 individuals along 79.9 km of transects. Estimated population density was 47.0 pigs/km² in 1996 and 27.0 pigs/km²‐ in 1998. Sus scrofa biomass in this forest was estimated at 1837 kg/km² and 1346 kg/ km² in 1996 and 1998, respectively. Differences between years were attributed to changes in the density of young pigs, coincident with a mast‐seeding year of dipterocarp trees in 1996. Pig densities at Pasoh Forest Reserve were much higher than at other forest locations within the species' native range in Europe and Asia. Because Pasoh Forest Reserve is a forest fragment, two factors likely account for the extremely high pig densities: (1) local extinction of natural predators (mainly tigers and leopards) and (2) an abundant year‐round food supply of African oil palm fruits from extensive plantations bordering the reserve.     

The Influence of Topography on Tree Growth,Mortality, and Recruitment in a Tropical Montane Forest1

To determine if there were consistent differences in growth, mortality, and recruitment on slopes and ridge crests in tropical montane forests, which could explain the (frequent but not universal) low stature of trees in the ridgetop forests, we analyzed data from long‐term plots in Jamaica (1990–1994; sixteen 200‐m² plots, six on ridge crests and five each on north and south slopes). Mortality was higher on north slopes, while growth and recruitment rates were not significantly different among positions. Soil pH and effects of recent disturbance by Hurricane Gilbert were positively correlated with growth and recruitment, while slope angle and disturbance effects were the best predictors of mortality. The patterns we found in Jamaica, that growth and recruitment were not higher on ridge crests than slopes, are different than those found by Herwitz and Young in Australia where growth and turnover were greater on a ridge crest. Therefore, it is not possible at present to make simple generalizations about dynamics of ridge crest versus slope forests in the montane tropics.