Effects of selective logging on tree diversity,composition and plant functional type patterns in a Bornean rain forest

Abstract. The effects of selective logging on tree diversity, changes in tree species composition and plant functional types were studied with the use of seven permanent plots in virgin and in logged forest. All plots were located in a lowland dipterocarp rain forest in East Kalimantan on the island of Borneo. Just after logging and during the following 20 yr tree diversity measured as Fishers’α was not significantly affected in logged forest plots. Temporal shifts in tree species composition were analysed with Principal Component Analysis (PCA). Logged forest plots had much larger changes over time than virgin forest plots. In the smallest diameter class, some logged forest plots showed a distinct trajectory in PCA space compared to virgin forest plots, while in larger diameter classes movement of logged plots in PCA space was random. This suggests that there is no predetermined community to which logged forest plots tend to shift when recovering from logging. We found a significant negative correlation between diameter increments and the species‐specific wood densities of tree species. Species‐specific wood density and potential tree height were used to assign species to five PFTs. As expected, logging increased the fraction of softwood stems in small diameter classes. In the largest diameter classes (≥ 50 cm DBH) a strong decrease of softwood emergent stems was found in logged forest plots. After more than 20 yr no recruitment was found of softwood emergent stems in selectively logged forest.     

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.     

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.     

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.     

Impacts of Intensive Logging on the Trophic Organisation of Ant Communities in a Biodiversity Hotspot

Trophic organisation defines the flow of energy through ecosystems and is a key component of community structure. Widespread and intensifying anthropogenic disturbance threatens to disrupt trophic organisation by altering species composition and relative abundances and by driving shifts in the trophic ecology of species that persist in disturbed ecosystems. We examined how intensive disturbance caused by selective logging affects trophic organisation in the biodiversity hotspot of Sabah, Borneo. Using stable nitrogen isotopes, we quantified the positions in the food web of 159 leaf-litter ant species in unlogged and logged rainforest and tested four predictions: (i) there is a negative relationship between the trophic position of a species in unlogged forest and its change in abundance following logging, (ii) the trophic positions of species are altered by logging, (iii) disturbance alters the frequency distribution of trophic positions within the ant assemblage, and (iv) disturbance reduces food chain length. We found that ant abundance was 30% lower in logged forest than in unlogged forest but changes in abundance of individual species were not related to trophic position, providing no support for prediction (i). However, trophic positions of individual species were significantly higher in logged forest, supporting prediction (ii). Consequently, the frequency distribution of trophic positions differed significantly between unlogged and logged forest, supporting prediction (iii), and food chains were 0.2 trophic levels longer in logged forest, the opposite of prediction (iv). Our results demonstrate that disturbance can alter trophic organisation even without trophically-biased changes in community composition. Nonetheless, the absence of any reduction in food chain length in logged forest suggests that species-rich arthropod food webs do not experience trophic downgrading or a related collapse in trophic organisation despite the disturbance caused by logging. These food webs appear able to bend without breaking in the face of some forms of anthropogenic disturbance.     

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.     

Effects of Forest Management on Diversity and Abundance of Fruit-feeding Nymphalid Butterflies in South-eastern Côte d'Ivoire

The effects of forest management on vegetation structure and capture frequencies of nymphalid butterflies were studied in a logged rain forest in south-east Côte d'Ivoire. An experimental compartment, where liberation thinning was carried out 3 years before, and a 5-year-old mono-dominant tree plantation were compared to a regenerating control compartment; 3642 specimens of 97 species were trapped. Accumulated species richness and diversity indices were lower in the control compared to the liberation thinning compartment, but lowest in the plantation. However, the habitat preference for traps situated in the control compartment was negatively correlated with the size of the species geographical range. Four out of five species with a lower capture frequency in the liberation thinning compartment showed preferences for mature succession stages and were either Upper Guinean endemics or Guinea–Congolian–restricted species. The seven species with higher capture frequencies in the liberation thinning compartment were all geographically widespread. Five of them showed higher frequencies in younger succession stages. Eight species, three canopy specialists and two Guinea savannah species, significantly preferred the plantation, while 17 species avoided this management type. Liberation thinning seems to affect the more specialised species with smaller geographic ranges, thus risking loss of regional diversity.     

Changes in avian communities of Budongo Forest Reserve after 70 years of selective logging

Owiunji, I. 2000. Changes in avian communities of Budongo Forest Reserve after 70 years of selective logging. Ostrich 71 (1 & 2): 216–219.

Budongo Forest Reserve, located in the west of Uganda, has been selectively logged for timber for over 70 years, and has a well documented management history. Changes in the avian community, species diversity and relative densities were assessed in two unlogged and three logged and arboricidally treated compartments. Tree-species diversity was highest in the disturbed forest. Over 100 forest bird species were recorded (including a new record for East Africa, Puvel's Illadopsis Illadopsis puveli. Both point counts and mist-netting showed that bird species diversity was higher in the logged and treated than in the unlogged forest. Five species had significantly higher densities in logged forest, three species had significantly higher densities in unlogged forest and 14 species showed no significant change in densities. The response of Budongo Forest birds to disturbance was species-specific.     

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.     

An examination of scale of assessment,logging and ENSO-induced fires on butterfly diversity in Borneo

The impact of disturbance on species diversity may be related to the spatial scales over which it occurs. Here I assess the impact of logging and ENSO (El Niño Southern Oscillation) -induced burning and forest isolation on the species richness (477 species out of more than 28,000 individuals) and community composition of butterflies and butterfly guilds using small (0.9 ha) plots nested within large (450 ha) landscapes. The landscapes were located in three habitat classes: (1) continuous, unburned forest; (2) unburned isolates surrounded by burned forest; and (3) burned forest. Plots with different logging histories were sampled within the two unburned habitat classes, allowing for independent assessment of the two disturbance factors (logging and burning). Disturbance within habitat classes (logging) had a very different impact on butterfly diversity than disturbance among habitat classes (due to ENSO-induced burning and isolation). Logging increased species richness, increased evenness, and lowered dominance. Among guilds based on larval food plants, the species richness of tree and herb specialists was higher in logged areas but their abundance was lower. Both generalist species richness and abundance was higher in logged areas. Among habitat classes, species richness was lower in burned forest and isolates than continuous forest but there was no overall difference in evenness or dominance. Among guilds, generalist species richness was significantly lower in burned forest and isolates than continuous forest. Generalist abundance was also very low in the isolates. There was no difference among disturbance classes in herb specialist species richness but abundance was significantly higher in the isolates and burned forest than in continuous forest. Tree specialist species richness was lower in burned forest than continuous forest but did not differ between continuous forest and isolates.The scale of assessment proved important in estimating the impact of disturbance on species richness. Within disturbance classes, the difference in species richness between primary and logged forest was more pronounced at the smaller spatial scale. Among disturbance classes, the difference in species richness between continuous forest and isolates or burned forest was more pronounced at the larger spatial scale. The lower levels of species richness in ENSO-affected areas and at the larger (landscape) spatial scale indicate that future severe ENSO events may prove one of the most serious threats to extant biodiversity.     

Habitat of the white-footed dunnart Sminthopsis leucopus (Gray) (Dasyuridae: Marsupialia) in a logged,burnt forest near Bega,New South Wales

In the winter of 1981, 9 months after an intense fire in November 1980 in the recently logged Mumbulla State Forest, the numbers of a small population of Sminthopsis leucopus had slightly increased, while the common species of small mammals had declined sharply from the previous winter. The study reported in this paper aimed to identify the habitat of S. leucopus. Ninety study plots were divided equally among three classes of forest (unlogged forest, and forest logged in 1979 and 1980), between two aspects (north- and south-facing slopes), and among three topographic positions (ridge, midslope and gully). Plots were assessed on the basis of both floristic data and the structural components of ground, shrub and tree cover. The primary finding was that the habitat of S. leucopus was the treeless ridges and mid-slopes with less than 51% ground cover in the logged areas of burnt forest. The floristic analysis showed that the vegetation subcommunities restricted to gullies, or carrying a ferny ground layer of vegetation, were not selected by S. leucopus. An analysis of movements of male S. leucopus identified two classes. ‘Resident’ males occurred on logged ridges and midslopes characterized by a floristic subcommunity of open forest with a grassy understorey.‘Explorer’ males moved through a variety of ridge habitats, including unlogged forest, and were spread more widely through the floristic communities. The pattern of habitat selection identified in this study shows S. leucopus to be ecologically distinct from the other species of small mammals in the forest and thus in need of special consideration in management programmes if it is to be conserved. Under the current regime, the uniform treatment of the forest leads to widespread stands of dense regrowth to the exclusion of the species and is inimical to its survival.     

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.     

Effects of disturbance caused by selective timber extraction on fish communities in Sabah, Malaysia

To examine the effects of selective timber extraction on fish communities in Sabah, Malaysia, quantitative samples of fishes were taken from thirteen streams running through undisturbed rainforest or through forest that had been selectively logged 3–18 years previously. Multivariate analysis (canonical discriminant analysis and cluster analysis) indicated that mesohabitats within streams (riffles and pools) and differences in stream size were more important in determining community structure than logging history. Riffles in streams running through logged or undisturbed forest were indistinguishable using relative biomass or abundance data, as were pools from small streams (approximate order 2). Fish communities from pools in larger streams showed some separation in multivariate space corresponding to a complex set of relative changes in abundance and/or biomass between species. However it was difficult to unambiguously assign such changes to logging regime alone. There appeared to be some differences in fish communities between streams in recently-logged (3–7 years) and old-logged (17–18 years) areas related to abundance or biomass of three cyprinids (Garra borneensis, Lobocheilos bo and Osteochilus chini). Only one species, Pangio mariarum, was not found in streams in logged forest, but it was only found at one location in undisturbed forest. A number of other species showed significant differences in abundance or biomass between sites but most of these were only present at some sites and in low abundance. Principal components analysis of habitat data showed that riffle sites were homogeneous whatever their logging history as were pools in unlogged large streams. Pools in logged large streams were significantly more heterogeneous but in a random rather than systematic manner. It is concluded that the type of selective logging practices used locally have low impact on fish communities through mechanisms of persistence and/or rapid recolonisation.     

Effects of selective logging (50 years ago) on habitat use and ranging behaviour of a forest understorey bird (red-tailed bristlebill, Bleda syndactyla) in Uganda

To understand the effects of selective logging on animals we compared habitat use and ranging behaviour of a common understorey passerine bird, the red‐tailed bristlebill (Bleda syndactyla), in logged and unlogged forest in the Budongo Forest Reserve, Uganda. The secondary forest had been selectively logged about 50 years ago, and differed in vegetation structure from the unlogged, primary forest in particular by having a denser understorey. Home range size of radio‐tagged bristlebills was 10–20% larger (depending on data sample used) in unlogged forest compared with logged forest, but the difference was not significant. Movement rates during 1‐h observation periods were highest in unlogged forest. The bristlebill has been characterized as a bird of dense understorey vegetation, and data from unlogged forest in the present study suggested that areas with dense understorey were used more often than expected. In logged forest, no habitat preferences were found, probably because the forest had a dense understorey throughout. Assuming that smaller home ranges and lower movement rates indicate better habitat, there was no evidence that bristlebills were negatively affected by logging. The preference for dense understorey in unlogged forest suggests that the bristlebill may benefit from selective logging because this leads to an increase in dense understorey.     

The effects of selective logging on the distribution of moths in a Bornean rainforest

The effects of selective logging on the diversity and species composition of moths were investigated by sampling from multiple sites in primary forest, both understorey and canopy, and logged forest at Danum Valley, Sabah, Malaysia. The diversity of individual sites was similar, although rarefied species richness of logged forest was 17% lower than for primary forest (understorey and canopy combined). There was significant heterogeneity in faunal composition and measures of similarity (NESS index) among primary forest understorey sites which may be as great as those between primary understorey and logged forest. The lowest similarity values were between primary forest understorey and canopy, indicating a distinct canopy fauna. A number of species encountered in the logged forest were confined to, or more abundant in, the canopy of primary forest. Approximately 10% of species were confined to primary forest across a range of species' abundances, suggesting this is a minimum estimate for the number of species lost following logging. The importance of accounting for heterogeneity within primary forest and sampling in the canopy when measuring the effects of disturbance on tropical forest communities are emphasized.     

Rehabilitation of Logged Rain Forests: Avifaunal Composition,Habitat Structure,and Implications for Biodiversity‐Friendly REDD+

The inclusion of carbon stock enhancements under the Reducing Emissions from Deforestation and Forest Degradation (REDD+) framework will likely drive a rapid increase in biosequestration projects that remove carbon from the atmosphere through rehabilitation of degraded primary rain forests. Such projects could also present an important opportunity to reverse losses of biodiversity from degraded rain forests, but concern has recently been expressed that management interventions to increase carbon stocks may conflict with biodiversity conservation. Focusing on a large‐scale rain forest rehabilitation project in northern Borneo, we examine: (i) how intensive rehabilitation of selectively logged forests affected patterns of bird community composition and (ii) whether changes in vegetation structure explain observed shifts in avian guild structure and species composition. Bird composition differed between unlogged, naturally regenerating logged, and rehabilitated logged habitats, with the avifauna of rehabilitated forest more similar to that of naturally regenerating forest. Crucially, rehabilitation did not adversely affect either those species that declined after logging or those species that are IUCN Red Listed. Rehabilitation reduced the prevalence of vines and shrubs within regenerating forest, and across all habitats, the abundance and species richness of all birds and of obligate frugivores were positively related to vine prevalence. In contrast, the abundance and richness of frugivore–insectivore generalists and of salliers were negatively related to vines, suggesting that avifaunal responses to forest rehabilitation were attributable to liberation cutting of vines. Management intervention to increase carbon stocks had little adverse effect on avian biodiversity and we therefore argue that rain forest rehabilitation should play a strong role in future REDD+ agreements.     

Differential diameter-size effects of forest management on tree species richness and community structure: implications for conservation

In this paper we tested the hypothesis that logging effects in the adult tree community reverberate upon the regeneration contingent. We examined the differences on the tree community between forest reserves and 10 year-old logged areas in the Yucatan Peninsula. We used a paired design in three independent sites to estimate the effects of logging on tree species richness, diversity, composition and structure. Analyses were conducted differentiating individuals of four diameter-size classes: 1–5, 5–10, 10–25, and >25 cm DBH. We found out that there were differential effects by size. Species richness in the smaller and larger diameter-size classes was significantly lower in logged areas. Floristic composition was also different between logged and unlogged areas, with a trend towards more secondary forest associated species and less primary forest associated species in logged areas, and a higher density of species represented by a single individual in unlogged reserves. In terms of structure, trees DBH <10 cm, lianas, and re-sprouting stumps were more abundant in logged areas. Our findings suggest that 10 years after logging, harvested areas show alterations in structure, and potentially a reduction in species richness. We suggest that to make timber extraction and forest conservation compatible at this site, it is necessary to gain a better understanding of the ecology and regeneration requirements of the less abundant species, and to assess whether current logging practices might hinder their permanence in the study area.