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.     

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.     

Feeding ecology and dietary differences between blue monkey (Cercopithecus mitis stuhlmanni Matschie) groups in logged and unlogged forest, Budongo Forest Reserve, Uganda

Several studies have investigated the effects of logging on primate population density, and attempted to relate density differences to changes in vegetation composition. As population density in some forest primates may be considerably influenced by dietary quality, it is important to understand how the effects of vegetation changes commonly associated with logging influence primate feeding ecology. Results are presented from a study carried out in the Budongo Forest Reserve in Uganda to investigate differences in blue monkey (Cercopithecus mitis stuhlmanni) feeding ecology in forest blocks with different logging history. Dietary composition differed between logged and unlogged forest, with a higher proportion of unripe fruit consumed in logged forest. By contrast, the proportion of young leaves, invertebrates and seeds in the diet was higher in unlogged forest. Variations in the intake of fruit was also higher in unlogged forest, but feeding patterns on other dietary items were more selective. These differences in diet reflect differences in availability of different categories of food items, as determined by tree species composition, abundance and plant phenological patterns. The observed differences between feeding ecology in logged and unlogged forest are discussed in relation to primate feeding ecology and habitat composition in Budongo.     

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.     

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.     

Density‐dependent habitat selection predicts fitness and abundance in a small lizard

Density‐dependent habitat selection has been used to predict and explain patterns of abundance of species between habitats. Thermal quality, a density‐independent component of habitat suitability, is often the most important factor for habitat selection in ectotherms which comprise the vast majority of animal species. Ectotherms may reach high densities such that individual fitness is reduced in a habitat due to increased competition for finite resources. Therefore, density and thermal quality may present conflicting information about which habitat will provide the highest fitness reward and ectotherm habitat selection may be density‐independent. Using ornate tree lizards Urosaurus ornatus at 10 sites each straddling two adjacent habitats (wash and upland), we tested the hypothesis that habitat selection is density‐dependent even when thermal quality differs between habitats. We first tested that fitness proxies decline with density in each habitat, indicating density‐dependent effects on habitat suitability. We also confirmed that the two habitats vary in suitability (quantified by food abundance and thermal quality). Next, we tested the predictions that habitat selection depends on density with isodar analyses and that fitness proxies are equal in the two habitats within a site. We found that monthly survival rates decreased with density, and that the wash habitat had more prey and higher thermal quality than the upland habitat. Lizards preferred the habitat with more food and higher thermal quality, lizard densities in the two habitats were positively correlated, and fitness proxies of lizards did not differ between habitats. These patterns are consistent with density‐dependent habitat selection, despite differences in thermal quality between habitats. We expect that density‐dependent habitat selection is widespread in terrestrial ectotherms when densities are high and temperatures are close to their optimal performance range. In areas where thermal quality is low, however, we expect that depletable resources, such as food, become less limiting because assimilating resources is more difficult.     

Population census of the chimpanzees in the Kalinzu Forest,Uganda: Comparison between methods with nest counts

The population of chimpanzees in the Kalinzu Forest, Uganda was censused with nest counts using the line-transect method. Four methods were examined to estimate density. The estimated densities, ranging from 2.0 to 4.7 chimpanzees per km², all indicated a high population density, in comparison with other chimpanzee habitats. Moreover, the density in the logged area of the Kalinzu Forest was higher than that in the unlogged area. Several factors are thought to contribute to the high density in the Kalinzu Forest. Notable are the mosaic forest structure dominated by mature forest with patches of logged areas, selective logging of non-food tree species of chimpanzees, and low hunting pressure. These results suggest that selective logging may be a practical means of primate conservation in places where timbers are exploited.     

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.     

Inter-annual dynamics and persistence of small mammal communities in a selectively logged tropical forest in Borneo

Understanding temporal change and long-term persistence of species and communities is vital if we are to accurately assess the relative values of human-modified habitats for biodiversity. Despite a large literature and emerging consensus demonstrating a high conservation value of selectively logged tropical rainforests, few studies have taken a long-term perspective. We resampled small mammals (≤?1 kg) in a heavily logged landscape in Sabah, Borneo between 2011 and 2016 to investigate temporal patterns of species-level changes in population density. We found that small mammal population density in heavily logged forest was highly variable among years, consistent with patterns previously observed in unlogged forest, and uncovered evidence suggesting that one species is potentially declining towards local extinction. Across nine species, population densities varied almost sevenfold during our 6-year study period, highlighting the extremely dynamic nature of small mammal communities in this ecosystem. Strictly terrestrial murid species tended to exhibit strong temporal dynamics, whereas semi-arboreal foraging species such as treeshrews had more stable dynamics. We found no relationships between population density and fruit/seed mass, and therefore no evidence that our patterns represent responses to inter-annual mast fruiting of the dominant canopy dipterocarp trees. This may be due to the removal of most of the canopy during logging, and hence the dipterocarp seed resource, although it possibly also reflects spatiotemporal limitations of our data. Our results underline the importance of understanding long-term variability in animal communities before developing conservation and management recommendations for human-altered ecosystems.     

Densities of Bornean orang‐utans (Pongo pygmaeus morio) in heavily degraded forest and oil palm plantations in Sabah,Borneo

The conversion of forest to agriculture continues to contribute to the loss and fragmentation of remaining orang‐utan habitat. There are still few published estimates of orang‐utan densities in these heavily modified agricultural areas to inform range‐wide population assessments and conservation strategies. In addition, little is known about what landscape features promote orang‐utan habitat use. Using indirect nest count methods, we implemented surveys and estimated population densities of the Northeast Bornean orang‐utan (Pongo pygmaeus morio) across the continuous logged forest and forest remnants in a recently salvage‐logged area and oil palm plantations in Sabah, Malaysian Borneo. We then assessed the influence of landscape features and forest structural metrics obtained from LiDAR data on estimates of orang‐utan density. Recent salvage logging appeared to have a little short‐term effect on orang‐utan density (2.35 ind/km ²), which remained similar to recovering logged forest nearby (2.32 ind/km ²). Orang‐utans were also present in remnant forest patches in oil palm plantations, but at significantly lower numbers (0.82 ind/km ²) than nearby logged forest and salvage‐logged areas. Densities were strongly influenced by variation in canopy height but were not associated with other potential covariates. Our findings suggest that orang‐utans currently exist, at least in the short‐term, within human‐modified landscapes, providing that remnant forest patches remain. We urge greater recognition of the role that these degraded habitats can have in supporting orang‐utan populations, and that future range‐wide analyses and conservation strategies better incorporate data from human‐modified landscapes.     

Population densities of Sulawesi crested black macaques (Macaca nigra) on Bacan and Sulawesi,Indonesia: Effects of habitat disturbance and hunting

Population surveys of Sulawesi crested black macaques (Macaca nigra) were conducted on the Indonesian islands of Sulawesi and Bacan in 1992–1994 to assess the status of natural populations and determine habitat and anthropogenic factors affecting their population densities. We surveyed five sites for primates, including undisturbed and disturbed habitats. Data were collected on habitat structure and composition at two undisturbed and one disturbed forest site in which the primates were surveyed. The highest density of macaques was found in primary forest at Gunung Sibela Nature Reserve on Bacan (170.3 individuals/km²). Population density in logged forest on Bacan was high but significantly less than primary forest (133.4 individuals/km²). The high density of crested black macaques in primary forest on Bacan is best explained by the high carrying capacity found in primary forest. The lower food quantity and quality of food resources found in logged forest correlated with lower primate densities compared to primary forest. However, the large population of macaques in logged forest demonstrates the conservation value of such forest. Densities on Sulawesi at Tangkoko-Batuangas-DuaSudara Nature Reserve (TBDS) showed a continuing decline since earlier surveys. Primate densities were highest near the protected center of Tangkoko Reserve (66.7 individuals/km²). The peripheral areas of Batuangas and DuaSudara, even though adjacent and continuous, showed lower population densities of 46.4 and 23.5 individuals/km², respectively. The best explanation for the continued decline of Macaca nigra populations at TBDS is hunting. Unless conservation measures are implemented immediately, M. nigra on Sulawesi risks extinction in the near future. Am. J. Primatol. 44:89–106, 1998. © 1998 Wiley-Liss, Inc.     

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.     

Effects of reduced‐impact logging on physical habitat and fish assemblages in streams of Eastern Amazonia

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Assessing the function of geophagy in a Malagasy rain forest lemur

Geophagy has been observed in nearly every long‐term study of folivorous primates. Yet despite frequent observations of this behavior, conclusive explanations for soil consumption remain ambiguous. This study tests the most frequently proposed hypotheses for geophagy using data collected on the geophagic behavior of the Milne‐Edwards’ sifaka (Propithecus edwardsi) living in two forests with varying levels of disturbance within Ranomafana National Park, Madagascar. Field data were collected from December 2002 to November 2003, during which time soil samples were collected for mineral analyses from 102 sites selected for geophagy and 42 control sites at which no geophagy was recorded. Results indicate that control soils differ significantly between the two study sites and that this difference is primarily attributable to varying levels of habitat disturbance. The frequency of soil consumption also does not vary significantly by sex or between logged and unlogged habitats. Soil consumption significantly correlated with fruit/seed consumption overall, but to a lesser degree in logged compared with unlogged sites. Clay minerals found in soils likely prevent absorption of dietary toxins in the gut and/or may mediate gut pH. This provides strong evidence for the protection hypothesis for geophagy, which may be especially important in areas where logging, or other forms of habitat disturbance, has been experienced. Abstract in French is available with online material.     

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.     

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.     

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.     

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.     

Long-term consequences of changes in territory quality on feeding and reproductive strategies of vervet monkeys

1. Food availability and quality are important determinants of mammalian reproductive success, and long-term changes in food availability were assessed for their impact on diets and reproduction of three adjacent groups of vervet monkeys in Amboseli, Kenya in two periods spanning an interval of 9 years.
2. Diets were largely restricted to the products of two species of acacia trees ( Acacia xanthophloea and Acacia tortilis ), with food selection primarily determined by availability (tree density, size and seasonal production of foods).
3. Over this period the overall abundance of major foods, measured through absolute species density, declined while territory size increased.
4. Despite significant changes in food plant densities, diets remained relatively stable, suggesting a component of consistency in diet choice. Limited options or high costs for incorporation of novel foods are suggested as factors maintaining this stability, with deleterious consequences in the face of very long-term habitat changes.
5. This study suggests that the habitat deterioration, assessed by reduction in food densities, initiated local group extinction. An increased energy expenditure in foraging, high mortality and low reproductive rates ultimately led to a population crash under conditions of reduced food availability.