Effects of rain forest logging on species richness and assemblage composition of small mammals in Southeast Asia

Aim The effects of logging and habitat degradation on the richness and abundance of small mammals in Asian rain forests are largely unknown. This work compares the species richness, dominance and evenness of small non‐volant mammals between logged and unlogged forests, and assesses whether assemblage variability (β‐diversity) is similar between forest types. Location Southeast Asia, northern Borneo (Sabah, Malaysia), Sunda‐shelf. Methods We surveyed species‐rich assemblages of small non‐volant mammals in three unlogged and three logged forests for 2 years. At each forest site, we sampled a permanently marked transect and two additional sites in three trapping sessions. All analyses were performed at both levels to include the effects of local abundances and point estimates, separately from the relative abundances of species on a more regional scale. Results We trapped a total of 1218 individuals of 28 species. Eleven common species accounted for 95% of all captures. Species richness and diversity were significantly higher in unlogged forest (27 species) than in logged forest (17 species). This was mainly attributable to the smaller number of rarely recorded species in logged forest (five compared with 16 in unlogged forest, with a total of fewer than 10 captures). However, all common species were present in both logged and unlogged forests, and our analyses revealed similar patterns of dominance, evenness and fluctuations in abundance. Hence overall assemblage composition in multivariate space did not differ greatly between forest types. Assemblages of Muridae and Tupaiidae showed similar population fluctuations in space and time, indicating that the ecology of these taxa may be partially driven by the same environmental factors. Main conclusions Although species were distributed patchily within sites, analyses at local and regional scales revealed similar patterns in diversity and assemblage variability, suggesting that effects of forest modification did not differ extensively locally and regionally, but had a profound effect on rare species. Our results emphasize the importance and conservation value of logged forest stands that are able to hold a large proportion of the small mammals also found in unlogged forests. Rare and more specialized species are more vulnerable to forest degradation than commonly caught species, resulting in the complete loss, or a decrease in numbers, of certain groups, such as arboreal small mammals and Viverridae.     

Unfavorable successional pathways and the conservation value of logged tropical forest

Conservation agencies are increasingly advocating that the survival of many tropical forest species may depend on disturbed forests and are directing scarce conservation resources to managing logged forests. This requires critical evaluation. In this 10-year study, we quantify tree community dynamics in three selectively logged areas harvested at different intensities and compare their recovery to two unlogged areas in Kibale National Park, Uganda. Over the 10 years, 527 of the original 4840 tagged trees (10cm DBH) died; while the mortality rate was highest in the heavily logged area, it was only significantly greater than one of the unlogged sites. The density of new trees varied significantly among areas being highest in the lightly logged area, but the density of new recruits in the heavily logged area did not differ from the unlogged areas. Overall, the more heavily logged areas had higher growth rates than the unlogged or lightly logged areas. However, there was no difference among areas in the magnitude of the change in basal area; and in both 1990 (20+ years post-harvest) and 2000 the basal area in the heavily logged area was significantly less than the unlogged areas. In general, findings from the heavily logged area suggest that the expectation that in logged areas a number of processes of forest regeneration will be accelerated relative to unlogged forest was not met. Thus, 30+ years after logging evidence suggests that forest recovery will be slower than expected. We also present evidence to suggest that the climate in the region is changing, which may account for changes in the population dynamics of certain species in the unlogged forests.     

Effects of Selection Logging on Birds in Northern Belize1

We compared bird diversity and frequency in selection logged and unlogged forest to determine the effects of recent selection logging on avian biodiversity in a subtropical, moist evergreen forest. We used a combination of mist netting and fixed-radius point counts to assess bird communities in February and March 1993 in northwestern Belize. Vegetation structure and composition was similar in logged and unlogged forest. The 66 most common species occurred with statistically similar frequency in logged and unlogged forest although 13 species were two times more frequent in intact forest. Numbers of total bird species were similar between logging gaps and the logged forest matrix, and between the logged forest matrix and unlogged forests. A comparison of numbers of species in 26 guilds based on migration strategy, diet, foraging substrate, and height strata also showed them to be similar regardless of logging history. Our results differed from previous studies that reported lower bird species richness and abundance of individual species in logged tropical forests than in unlogged forest. The differences might be explained by the lower logging intensity and/or greater levels of natural disturbance in our study area compared to previous studies.     

Effects of selective logging on the arboreal ants of a Bornean rainforest

We examined the effect of selective logging and corresponding forest canopy loss on arboreal ant diversity in a tropical rainforest. Arboreal ants were collected from an unlogged forest plot and from forest plots selectively logged 14 years and 24 years earlier in Danum Valley, Sabah, Malaysia, using a canopy fogging method. Selective logging was associated with a significant decrease in canopy cover and an increase in understory vegetation density relative to unlogged forest. Our study showed that selective logging in primary forest might not dramatically decrease total species number and overall abundance of arboreal ants; however, it may influence the species composition and dominance structure of the ant community, accompanied by an increase in abundance of shrub‐layer species and trophobiotic species. In view of the results of this study, management techniques that minimize logging impact on understory vegetation structure are likely to help maintain the conservation value of logged forests for arboreal ants. Our results also suggest that accurate assessment of the impacts on biodiversity should not be based only on measurement of species number and overall abundance, but also on analysis of species composition and community structure.     

Forest Regeneration in Logged and Unlogged Forests of Kibale National Park, Uganda

Processes of forest regeneration in two unlogged areas and in three areas that were logged nearly 25 years ago were quantified in Kibale National Park, Uganda. For forests to recover from logging, one would predict recruitment and growth processes to be accelerated in logged areas relative to unlogged areas, facilitating increased recruitment of trees into the adult size classes. We examined this prediction first by determining the growth of 4733 trees over a 51 to 56 month period and found that growth rates in the most heavily logged area were consistently slower than in the two unlogged areas. In contrast, the lightly logged forest had similar growth rates to unlogged areas in the small size classes, but trees in the 30 to 50 cm DBH size cohort exhibited elevated growth rates relative to the unlogged areas. Mortality was highest in the heavily logged areas, with many deaths occurring when healthy trees were knocked over by neighboring treefalls. We found no difference in the density or species richness of seedlings in the logged and unlogged forests. The number of seedlings that emerged from the disturbed soil (seed bank+seed rain) and initially seed-free soil (seed rain) was greater in the logged forest than in the unlogged forest. However, sapling density was lower in the heavily logged areas, suggesting that there is a high level of seedling mortality in logged areas. We suggest that the level of canopy opening created during logging, the lack of aggressive colonizing tree species, elephant activity that is concentrated in logged areas, and an aggressive herb community, all combine to delay vegetation recovery in Kibale Forest.     

Effects of Historical Selective Logging on Anuran Communities in a Wet Evergreen Forest,South India

Vast areas of tropical evergreen forests have been selectively logged in the past, and many areas continue to be logged. The impacts of such logging on amphibians are poorly understood. I examined the response of anuran communities to historical selective logging in a wet evergreen forest in south India. Anuran assemblages in unlogged forest were compared with assemblages in selectively logged forest. Forty 10 m × 10 m quadrats in forest, riparian zones, and streams of unlogged and selectively logged forests were searched at night for anurans. Species richness did not appear to be affected by logging. However, anuran density varied significantly and was 42 percent lower in selectively logged forests compared to unlogged forests. Anuran densities also varied significantly across microhabitats, with highest densities in streams of both selectively logged and unlogged forests. Patterns of niche overlap varied with selective logging as niche breadth either expanded, contracted, or remained neutral for different species. Ordination analysis explained 95 percent of the variation in species assemblage across selectively logged and unlogged forests. The assemblage in selectively logged forest was nested within unlogged forest. Among the habitat characteristics, litter thickness and water depth had the highest influence on the assemblage. This was followed by litter/water temperature, air temperature, and lastly relative humidity. It appears that species richness and composition of anurans in selectively logged forests is converging with unlogged forests, but the effects of historical logging seem to persist on anuran densities and their niche characteristics even ca 40 yr since logging ceased.     

Assessing the use of butterflies as indicators of logging in Borneo at three taxonomic levels

Logging is an issue of major conservation concern. Less than 5% of tropical forests are currently protected, and many of these are in so-called "paper parks." Many species may therefore depend on exploited forests, and management decisions concerning these forests will be a major determinant of their survival. An important aspect of forest management will entail the use of reliable, practical, and inexpensive indicator taxa to monitor exploitation. Here, butterflies are proposed as such indicators. Species, generic, and subfamily richness was significantly higher in logged than unlogged forest and community composition differed significantly at all three taxonomic levels (species, genus, and subfamily). Richness estimators were, furthermore, highly correlated among all three taxonomic levels. Significant individual indicator taxa were found at all three taxonomic levels, but the best overall taxa (highest indicator values) were found at the generic level and included the butterfly genera Ragadia and Paralaxita as indicators of unlogged forest and the genera Ypthima, Allotinus, and Athyma as indicators of logged forest. The use of genera instead of species presents a number of practical advantages. Identification is faster, easier, and more reliable. Genera can, furthermore, usually be identified "on the wing," thereby preventing accidental mortality due to capture.     

Effects of isolation,logging and dispersal on woody-species richness of islands

Isolation effects on species richness of woody plants were investigated in a system of islands that were created by the filling of the Clarks Hill Reservoir, Georgia. This reservoir was built between 1946–1954. Some islands were logged and cleared of woody plants prior to the filling of the reservoir and others were not logged. The presence of logged versus unlogged islands in the same system allowed us to test whether and how geographical isolation interacts with island history and species-specific dispersal properties in determining patterns of among-island variation in species number. Thirty-six years after the islands were created, logged islands had significantly fewer species of woody plants than unlogged ones. On logged islands, total number of woody species was negatively correlated with distance to the closest mainland (r=–0.95). On unlogged islands, variation in species number was very low (CV=4.9%) and was not correlated with distance to the mainland. These results indicate that the studied system as a whole has not yet reached equilibrium. However, the mean number of species on unlogged islands was very close to the intercept of the regression obtained for logged islands, suggesting that islands close to the mainland have already reached their equilibrium species richness. This conclusion is consistent with predictions of island biogeography theory. When species representing different dispersal properties were analyzed separately, statistically significant distance effects were obtained for bird-dispersed species (r=0.88) and for species with no adaptations to bird or wind dispersal (r=0.81). Wind-dispersed species did not show a decrease in species number with increasing isolation, but their relative frequency was positively and significantly correlated with distance to the mainland (r=0.94). Historical factors, as well as differences among species in dispersal properties, are important in explaining patterns of among-island variation in species number.     

A comparison of streams in logged and unlogged areas of Great Smoky Mountains National Park

Second order streams draining areas of virgin forest in Great Smoky Mountains National Park, Tennessee and North Carolina, U.S.A., are compared to those which drain forests logged before the establishment of the park in the 1930's. Water quality of two main study streams (one unlogged and one formerly logged) was compared and the unlogged stream had generally higher levels of dissolved solids and lower levels of suspended particulates than the logged stream. Stream channel characteristics were compared on four logged and four unlogged streams. The unlogged streams had over four times more (by volume) of woody debris and 10 times more material in debris dams than the logged streams. Only minor differences in substrate composition were observed. Macroinvertebrate samples from the four logged and four unlogged streams showed that the logged streams contained greater numbers of organisms and more taxa. More detailed sampling on the two main study streams showed similar patterns of more individuals and more taxa in the logged stream, as well as differences in the composition of five functional groups. These differences in invertebrate fauna may be due to differences in quantity and quality of leaf litter inputs, although other explanations are also possible.     

Patterns of Richness and Abundance in a Tropical African Leaf‐litter Herpetofauna1

I compared species richness and habitat correlates of leaf‐litter herpetofaunal abundance in undisturbed and selectively logged forests, and an abandoned pine plantation in Kibale National Park, Uganda. I sampled 50 randomly located 25 m² litter plots in each area during the wet and dry seasons in 1997. Ten anuran, five lizard, and three snake species were captured in plots over the study. Assemblage composition was most similar at logged and unlogged sites. The logged forest herpetofauna had higher species richness and abundance than the unlogged forest, but diversity was greater in the unlogged forest due to greater evenness. In contrast, the pine plantation site had the highest richness, abundance, and evenness of the three study sites, but species composition was distinct from the other areas. Herpetofaunal densities were significantly lower in all three areas during the dry season than in the wet season. During the dry season, soil moisture, litter mass, topography, shrub cover, and number of fallen logs were significant positive predictors of herpetofaunal presence in litter plots, but only soil moisture was significant in the wet season. The interaction of moisture and topography appears to be important in determining seasonal patterns of litter herpetofaunal distribution. Comparison of litter herpetofaunal studies across the tropics have shown that mid‐elevation faunas generally support fewer species than lowland faunas. Compared with other tropical mid‐elevation litter faunas, Kibale supports an intermediate number of species, but at lower densities than observed at any other mid‐elevation site reported in the literature.     

Degraded lands worth protecting: the biological importance of Southeast Asia's repeatedly logged forests

Southeast Asia is a hotspot of imperilled biodiversity, owing to extensive logging and forest conversion to oil palm agriculture. The degraded forests that remain after multiple rounds of intensive logging are often assumed to be of little conservation value; consequently, there has been no concerted effort to prevent them from being converted to oil palm. However, no study has quantified the biodiversity of repeatedly logged forests. We compare the species richness and composition of birds and dung beetles within unlogged (primary), once-logged and twice-logged forests in Sabah, Borneo. Logging had little effect on the overall richness of birds. Dung beetle richness declined following once-logging but did not decline further after twice-logging. The species composition of bird and dung beetle communities was altered, particularly after the second logging rotation, but globally imperilled bird species (IUCN Red List) did not decline further after twice-logging. Remarkably, over 75 per cent of bird and dung beetle species found in unlogged forest persisted within twice-logged forest. Although twice-logged forests have less biological value than primary and once-logged forests, they clearly provide important habitat for numerous bird and dung beetle species. Preventing these degraded forests from being converted to oil palm should be a priority of policy-makers and conservationists.     

Natural regeneration of selected timber species in the Republic of Congo

Natural regeneration of timber species is critical to the sustainable management of tropical forests. To understand what determines regeneration success of timber species in the Congo Basin, we evaluated whether seedling recruitment rates differed between forest logged 30 years previously and unlogged forest and determined the environmental factors that influence seedling density, growth and survival. We monitored the fate of 2186 seedlings of seven timber species within 462, 25‐m² plots located along 21 transects. We characterized seedling plots by light availability, soil nutrient availability and pH, and abundance of mammalian herbivores and then used linear and generalized linear mixed models to evaluate the variables that influenced seedling density, growth and survival. Light availability and canopy openness were 18% and 81% higher in logged than unlogged forest, and concentration of soil nutrients varied between sites. Seedling density was 32% higher in unlogged than logged forest. Taking all species together, seedling survival was positively correlated with calcium and negatively with magnesium and available phosphorus. Rates of seedling growth increased with available light. Taken separately, seedlings of the selected timber species responded differently to abiotic and biotic factors, demonstrating species‐specific regeneration requirements.     

The high value of logged tropical forests: lessons from northern Borneo

The carbon storage and conservation value of old-growth tropical forests is clear, but the value of logged forest is less certain. Here we analyse >100,000 observations of individuals from 11 taxonomic groups and >2,500 species, covering up to 19?years of post-logging regeneration, and quantify the impacts of logging on carbon storage and biodiversity within lowland dipterocarp forests of Sabah, Borneo. We estimate that forests lost ca. 53% of above-ground biomass as a result of logging but despite this high level of degradation, logged forest retained considerable conservation value: floral species richness was higher in logged forest than in primary forest and whilst faunal species richness was typically lower in logged forest, in most cases the difference between habitats was no greater than ca. 10%. Moreover, in most studies >90% of species recorded in primary forest were also present in logged forest, including species of conservation concern. During recovery, logged forest accumulated carbon at five times the rate of natural forest (1.4 and 0.28?Mg?C?ha^?1?year^?1, respectively). We conclude that allowing the continued regeneration of extensive areas of Borneo??s forest that have already been logged, and are at risk of conversion to other land uses, would provide a significant carbon store that is likely to increase over time. Protecting intact forest is critical for biodiversity conservation and climate change mitigation, but the contribution of logged forest to these twin goals should not be overlooked.     

Multiple stages of tree seedling recruitment are altered in tropical forests degraded by selective logging

Tropical forest degradation is a global environmental issue. In degraded forests, seedling recruitment of canopy trees is vital for forest regeneration and recovery. We investigated how selective logging, a pervasive driver of tropical forest degradation, impacts canopy tree seedling recruitment, focusing on an endemic dipterocarp Dryobalanops lanceolata in Sabah, Borneo. During a mast‐fruiting event in intensively logged and nearby unlogged forest, we examined four stages of the seedling recruitment process: seed production, seed predation, and negative density‐dependent germination and seedling survival. Our results suggest that each stage of the seedling recruitment process is altered in logged forest. The seed crop of D. lanceolata trees in logged forest was one‐third smaller than that produced by trees in unlogged forest. The functional role of vertebrates in seed predation increased in logged forest while that of non‐vertebrates declined. Seeds in logged forest were less likely to germinate than those in unlogged forest. Germination increased with local‐scale conspecific seed density in unlogged forest, but seedling survival tended to decline. However, both germination and seedling survival increased with local‐scale conspecific seed density in logged forest. Notably, seed crop size, germination, and seedling survival tended to increase for larger trees in both unlogged and logged forests, suggesting that sustainable timber extraction and silvicultural practices designed to minimize damage to the residual stand are important to prevent seedling recruitment failure. Overall, these impacts sustained by several aspects of seedling recruitment in a mast‐fruiting year suggest that intensive selective logging may affect long‐term population dynamics of D. lanceolata. It is necessary to establish if other dipterocarp species, many of which are threatened by the timber trade, are similarly affected in tropical forests degraded by intensive selective logging.     

Spatial Ecology of the Endangered Milne-Edwards’ Sifaka (Propithecus edwardsi): Do Logging and Season Affect Home Range and Daily Ranging Patterns?

Primates often live in human-altered habitats; Malagasy lemurs are no exception. It is important to understand if habitat alteration affects primates’ space use patterns across multiple spatial and temporal scales, as this drives population density. We quantified the daily, seasonal, and annual space-use of seven groups of Milne-Edwards’ sifaka (Propithecus edwardsi) living in unlogged and logged rain forest in Ranomafana National Park, Madagascar between December 2002 and November 2003. Concurrent data showed that sifakas consumed higher quality foods in the unlogged than in logged forests; thus we explored how space use patterns were related to energy use strategies. Sifaka groups in the logged rain forest traveled 7–13% less per day than groups in the unlogged rain forest, despite their larger home ranges (median: 46.12 and 23.52 ha, in the logged and unlogged forests, respectively). Sifakas may thus use an energy-minimizing strategy at the scale of the individual day but an energy-maximizing strategy at the annual home range scale. Sifakas exhibited fidelity to the home range across seasons, but their core area of use shifted considerably with season. We found no difference in population density between sites. However, given the interannual variability in sifaka foods, a multiyear study is needed to assess if energy strategies observed in this study are consistent across longer time periods. Our findings suggest that lemurs may persist in logged habitats by altering spatial use patterns; future work should attempt to quantify the threshold level of forest regeneration from logging that will allow lemurs to persist at similar densities as in unlogged forest.     

The Effect of Logging on the Ground-Foraging Ant Community in Eastern Amazonia

In order to determine the suitability of ants as indicator organisms for habitat disruption in tropical forests, we studied the effects of both high and low impact logging on ant communities in northeastern Pará State, in the Brazilian Amazon. We collected ants from logged forests and unlogged forest sites with Winkler bags throughout the 1998 rainy season (January and April) and the following dry season (July and September). Both methods of timber harvesting showed impacts on ant community composition when compared with unlogged forest, although these impacts did not include modifications in total species richness or the relative contribution of each ant subfamily to the total number of species. Instead, logging induced alterations took place at the level of species and genera. A 2-fold reduction in the dominance of ants of the highly diverse genus Pheidole was associated with forest alterations in high-impact logging sites. Thus, logging in Amazonia can be seen to promote species shifts in ant communities, without, however, altering species richness. Ants of the genus Pheidole are potentially useful indicators for forest disturbances resulting from timber extraction.     

Impacts of logging on density-dependent predation of dipterocarp seeds in a South East Asian rainforest

Much of the forest remaining in South East Asia has been selectively logged. The processes promoting species coexistence may be the key to the recovery and maintenance of diversity in these forests. One such process is the Janzen-Connell mechanism, where specialized natural enemies such as seed predators maintain diversity by inhibiting regeneration near conspecifics. In Neotropical forests, anthropogenic disturbance can disrupt the Janzen-Connell mechanism, but similar data are unavailable for South East Asia. We investigated the effects of conspecific density (two spatial scales) and distance from fruiting trees on seed and seedling survival of the canopy tree Parashorea malaanonan in unlogged and logged forests in Sabah, Malaysia. The production of mature seeds was higher in unlogged forest, perhaps because high adult densities facilitate pollination or satiate pre-dispersal predators. In both forest types, post-dispersal survival was reduced by small-scale (1 m(2)) conspecific density, but not by proximity to the nearest fruiting tree. Large-scale conspecific density (seeds per fruiting tree) reduced predation, probably by satiating predators. Higher seed production in unlogged forest, in combination with slightly higher survival, meant that recruitment was almost entirely limited to unlogged forest. Thus, while logging might not affect the Janzen-Connell mechanism at this site, it may influence the recruitment of particular species.     

Leaf litter decomposition rates in degraded and fragmented tropical rain forests of Borneo

Previously extensive tracts of primary rain forest have been degraded by human activities, and we examined how the effects of forest disturbance arising from habitat fragmentation and commercial selective logging affected ecosystem functioning in these habitats by studying leaf litter decomposition rates in litter bags placed on the forest floor. The rain forests of Borneo are dominated by trees from the family Dipterocarpaceae, and we compared leaf litter decomposition rates of three dipterocarp species at eight forest fragment sites (area 3–3529 ha) that had different histories of disturbance pre‐fragmentation: four fragments had been selectively logged prior to fragmentation and four had been formed from previously undisturbed forest. We compared these logged and unlogged forest fragments with sites in continuous forest that had been selectively logged (two sites) and fully protected and undisturbed (two sites). After 120 d, undisturbed continuous forest sites had the fastest rates of decomposition (52% mass loss). Forest fragments formed from unlogged forest (32% mass loss) had faster decomposition rates than logged forest fragments (28% mass loss), but slower rates than continuous logged forest (39% mass loss). Leaves of a light‐demanding species (Parashorea malaanonan) decomposed faster than those of a shade‐tolerant species (Hopea nervosa), but decomposition of all three dipterocarp species that we studied responded similarly to logging and fragmentation effects. Reduced decomposition rates in logged and fragmented forest sites may affect nutrient cycling and thus have detrimental consequences for forest regeneration. Conservation management to improve forest quality should be a priority, particularly in logged forest fragments.