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.     

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.     

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.     

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.     

Can remnant frugivore species effectively disperse tree seeds in secondary tropical rain forests?

Seed dispersal by frugivores in tropical rain forests is important for maintaining viable tree populations. Over the years, vertebrate assemblages in tropical forests have been altered by anthropogenic disturbances, leading to concerns about the ability of remnant vertebrates to substitute for the lost or declining vertebrate populations. We compared vertebrate composition and frugivore visitation rates as an indirect measure of rate of seed dispersal in three tropical rain forests in Uganda, namely Mabira, Budongo and Kibale Forests. Mabira is highly disturbed, Kibale is little and Budongo is intermediate. The aim was to determine whether vertebrate assemblages in differentially disturbed forests had comparable abilities to disperse seeds and whether tree species were equally vulnerable to loss of seed dispersers. Assemblages of forest generalist species were similar in all forests, but specialists were less abundant in the heavily disturbed forest. Remnant frugivores in the heavily disturbed forest were mainly small-bodied species that spat seeds beneath fruiting trees compared to large-bodied species observed in the less disturbed forests that ingested and carried away the seeds. We postulate that the quantity of seeds dispersed in heavily disturbed forests is much reduced due to low visitation rates of frugivores and the absence of large frugivores that consume large quantities of fruit. The quality of seed dispersal is affected as well by the distance over which seeds are moved. Assessment of vulnerability of trees shows no evidence for disperser substitution for trees producing large fruits. Fruit trees with low nutritional contents and digestibility were least visited in frugivore-impoverished forests. The loss of large specialist frugivores is likely to affect recruitment of many trees, especially of species that cannot establish beneath adult conspecifics.     

Ecology of Pericopsis elata (Fabaceae), an Endangered Timber Species in Southeastern Cameroon

Pericopsis elata (Fabaceae) is a tall tree of high commercial value of the moist semi‐deciduous African forests. As a result of decades of logging it is now considered as threatened and listed on the IUCN Red List and CITES Appendix II, even though essential biological parameters controlling its population dynamics remain unknown. This study aims at improving the knowledge of the species' ecological parameters, and at assessing the impact of selective logging on its populations in an 118,052 ha forest in Cameroon. After inventorying the species in 1432 ha, mortality and growth were assessed over continuous 5‐ and 2‐yr periods in unlogged and logged areas, respectively. Phenology was monitored in the unlogged forest during 5 yr. The population structure shows high relative abundance of trees in medium size categories. Mean annual diameter increments in both environments did not differ significantly between unlogged and logged areas. P. elata is a deciduous species that flowers at the end of the main dry season. The minimum reproduction and effective flowering diameters were 32 and 37 cm, respectively. Fruit maturation took place during 7 mo. With a minimum logging diameter of 90 cm, the recovery rate computed over a 30‐yr period was > 100 percent. Selective logging harvested only 12.1 percent of the total number of seed trees and had little influence on the species' biological parameters. Securing sufficient regeneration as a post‐logging action is probably the most important consideration for achieving long‐term sustainability.     

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.     

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.     

Breeding birds in the tropical rain forests of Kibale National Park, Uganda

A combination of trapping and foot surveys was used to assess the breeding status of birds in unlogged and logged sites in the tropical rain forest of Kibale National Park, Uganda. Breeding of forest edge and gap species was greatly enhanced by logging, whereas crevice and hole nesting specialist breeders were adversely affected. Egg‐laying periods corresponded to higher rainfall regimes at the nearby field station. The relevance of these findings to bird conservation is discussed.     

Cordia millenii: on the risk of local extinction?

Selective logging of valuable tropical timber trees is a conservation concern because it threatens the long-term sustainability of forests. However, there is insufficient information regarding the postlogging recovery of harvested species. Here, I assessed the seed dispersal patterns, recruitment and abundance of Cordia millenii , a valuable timber tree in two Ugandan tropical rain forests that have been subjected to varying disturbance regimes. The aim was to determine the vulnerability of Cordia in these forests. The rate of seed dispersal was lower in the heavily disturbed Mabira Forest compared with the less disturbed Budongo Forest. Frugivores in Mabira were small-bodied individuals that spat seeds beneath fruiting trees, whereas 90% of the fruit in Budongo was consumed by large-bodied chimpanzees that disperse seeds over long distances. Juveniles of Cordia were not found in the closed forest, although they were found in forest gaps in Budongo but not Mabira. Mature tree density was higher in Budongo compared with Mabira. Lack of effective seed dispersal coupled with the inability of seedlings of Cordia to establish under closed canopy account for the arrested recruitment in Mabira. Enrichment planting in felling gaps is necessary to avoid local extinction of Cordia in forests without large vertebrates.     

The avifauna 23 years after logging in Kibale National park, Uganda

The bird fauna of logged and unlogged forest in Kibale National Park, western Uganda were studied and comparisons made. Species diversity and richness were higher in the logged areas. A single species, the Yellow-whiskered Greenbul Andropadus latirostris dominated in the logged compartment. However, the majority of individual birds found in the logged forest were generalist or forest-edge species. Over 84% of the forest interior specialist species that occurred in primary forest had recolonized or persisted in the logged forest; however, there were seven out of 48 understorey forest specialists that had not done so. Although there was moderate species overlap between the two habitats, the logged forest compartment had not fully recovered from logging after 23 years.     

Disturbance history mediates climate change effects on subtropical forest biomass and dynamics

The responses of forest communities to interacting anthropogenic disturbances like climate change and logging are poorly known. Subtropical forests have been heavily modified by humans and their response to climate change is poorly understood. We investigated the 9‐year change observed in a mixed conifer‐hardwood Atlantic forest mosaic that included both mature and selectively logged forest patches in subtropical South America. We used demographic monitoring data within 10 1 ha plots that were subjected to distinct management histories (plots logged until 1955, until 1987, and unlogged) to test the hypothesis that climate change affected forest structure and dynamics differentially depending on past disturbances. We determined the functional group of all species based on life‐history affinities as well as many functional traits like leaf size, specific leaf area, wood density, total height, stem slenderness, and seed size data for the 66 most abundant species. Analysis of climate data revealed that minimum temperatures and rainfall have been increasing in the last few decades of the 20th century. Floristic composition differed mainly with logging history categories, with only minor change over the nine annual census intervals. Aboveground biomass increased in all plots, but increases were higher in mature unlogged forests, which showed signs of forest growth associated with increased CO₂, temperature, and rainfall/treefall gap disturbance at the same time. Logged forests showed arrested succession as indicated by reduced abundances of Pioneers and biomass‐accumulators like Large Seeded Pioneers and Araucaria, as well as reduced functional diversity. Management actions aimed at creating regeneration opportunities for long‐lived pioneers are needed to restore community functional diversity, and ecosystem services such as increased aboveground biomass accumulation. We conclude that the effects of climate drivers on the dynamics of Brazilian mixed Atlantic forests vary with land‐use legacies, and can differ importantly from the ones prevalent in better known tropical forests.     

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.     

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.     

El Niño droughts and their effects on tree species composition and diversity in tropical rain forests

In this study I investigated the effects of the extreme, 1997/98 El Niño related drought on tree mortality and understorey light conditions of logged and unlogged tropical rain forest in the Indonesian province of East Kalimantan (Borneo). My objectives were to test (1) whether drought had a significant effect on tree mortality and understorey light conditions, (2) whether this effect was greater in logged than in undisturbed forest, (3) if the expected change in tree mortality and light conditions had an effect on Macaranga pioneer seedling and sapling densities, and (4) which (a)biotic factors influenced tree mortality during the drought. The 1997/1998 drought led to an additional tree mortality of 11.2, 18.1, and 22.7% in undisturbed, old logged and recently logged forest, respectively. Mortality was highest in logged forests, due to extremely high mortality of pioneer Macaranga trees (65.4%). Canopy openness was significantly higher during the drought than during the non-drought year (6.0, 8.6 and 10.4 vs 3.7, 3.8 and 3.7 in undisturbed, old logged and recently logged forest, respectively) and was positively correlated with the number of dead standing trees. The increase in light in the understorey was accompanied by a 30 to 300-fold increase in pioneer Macaranga seedling densities. Factors affecting tree mortality during drought were (1) tree species successional status, (2) tree size, and (3) tree location with respect to soil moisture. Tree density and basal area per surface unit had no influence on tree mortality during drought. The results of this study show that extreme droughts, such as those associated with El Niño events, can affect the tree species composition and diversity of tropical forests in two ways: (1) by disproportionate mortality of certain tree species groups and tree size classes, and (2) by changing the light environment in the forest understorey, thereby affecting the recruitment and growth conditions of small and immature trees.     

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 confounding effects of logging on tree seedling growth and herbivory in Central Amazon

Light availability is an important modulator of seedling growth and plant–herbivore dynamics. Logging increases light levels in forests, potentially altering herbivore–plant interactions that drive seedling establishment. We conducted a transplant experiment to evaluate how logging and herbivory affect seedling growth and survival in three shade‐tolerant tree species, at paired canopy gap and understory sites in logged forest and an adjacent unlogged area in central Amazonia (Brazil). Seedlings were either left exposed to naturally occurring insect herbivores or protected from insects by a fine netting structure. We measured the herbivore damage and growth rate of seedlings after 18 mo. In general, logged areas received more light than unlogged sites. Growth and herbivory rates were positively influenced by light, and herbivory was also influenced positively by logging. In gaps, increased growth mitigated foliar damage. Logging resulted in a loss of foliar tissue due to increased herbivory. Herbivory rates were higher in the understory of logged sites than in that of unlogged understory sites, but growth was similar in these areas. Thus, the understory of logged areas provided the least favorable sites for shade‐tolerant tree regeneration, due to higher herbivory rates. The effect of logging on biotic interactions can extend beyond the gaps it creates into untouched understory sites. To our knowledge, this is the first time such a pattern has been observed, highlighting the importance of evaluating the impact of logging on biotic interactions.     

Does selective logging change ground‐dwelling beetle assemblages in a subtropical broad‐leafed forest of China?

Selective logging with natural regeneration is advocated as a near‐to‐nature strategy and has been implemented in many forested systems during the last decades. However, the efficiency of such practices for the maintenance of forest species are poorly understood. We compared the species richness, abundance and composition of ground‐dwelling beetles between selectively logged and unlogged forests to evaluate the possible effects of selective logging in a subtropical broad‐leafed forest in southeastern China. Using pitfall traps, beetles were sampled in two naturally regenerating stands after clearcuts (ca. 50 years old, stem‐exclusion stage: selectively logged 20 years ago) and two mature stands (> 80 years old, understory re‐initiation stage: selectively logged 50 years ago) during 2009 and 2010. Overall, selective logging had no significant effects on total beetle richness and abundance, but saproxylic species group and some abundant forest species significantly decreased in abundance in selectively logged plots compared with unlogged plots in mature stands. Beetle assemblages showed significant differences between selectively logged and unlogged plots in mature stands. Some environmental characteristics associated with selective logging (e.g., logging strategy, stand age, and cover of shrub and moss layers) were the most important variables explaining beetle assemblage structure. Our results conclude that selective logging has no significant impacts on overall richness and abundance of ground‐dwelling beetles. However, the negative effects of selective logging on saproxylic species group and some unlogged forest specialists highlight the need for large intact forested areas for sustaining the existence of forest specialist beetles.