Density Estimation of the Endangered Udzungwa Red Colobus (Procolobus gordonorum) and Other Arboreal Primates in the Udzungwa Mountains Using Systematic Distance Sampling

Estimates of population density and abundance are essential for the assessment of nonhuman primate conservation status, especially in view of increasing threats. We undertook the most extensive systematic primate survey yet of the Udzungwa Mountains of Tanzania, an outstanding region for primate endemism and conservation in Africa. We used distance sampling to survey three arboreal monkey species, including the endangered and endemic Udzungwa red colobus (Procolobus gordonorum). Overall, we encountered 306 primate clusters over 287 km walked along 162 line transects. We found the lowest cluster densities for both red colobus and Angolan colobus (Colobus angolensis; 0.8 clusters/km²) in the least protected forest (Uzungwa Scarp Forest Reserve, US), while we found the highest densities (3.2 and 2.6 clusters/km² for red colobus; 3.2 and 2.7 clusters/km²for Angolan colobus) in two large and protected forests in the national park. Unexpectedly, Magombera, a small forest surrounded by plantations, had the highest densities of red colobus (5.0 clusters/km²), most likely a saturation effect due to the rapid shrinking of the forest. In contrast, Sykes’ monkey (Cercopithecus mitis monoides/moloneyi) had more similar densities across forests (3.1–6.6 clusters/km²), including US, suggesting greater resilience to disturbance in this species. For the endemic red colobus monkey, we estimated an abundance of 45–50,000 individuals across all forests, representing ca. 80% of the global population. Though this is a relatively high abundance, the increasing threats in some of the Udzungwa forests are of continued concern for the long-term survival of red colobus and other primates in the area.     

Long-term dynamics of wild primate populations across forests with contrasting protection in Tanzania

Anthropogenic activities driving tropical forests' loss imperil global biodiversity and provision of ecosystem services. In this context, systematic monitoring programs evaluating wildlife trends are essential. Non-human primates are relevant conservation targets since they represent vital components of tropical forests by serving as pollinators and seed dispersers. Here, we present primate group counts data collected over 19 years in a primate hotspot in Tanzania. We analyzed data with a hierarchical dynamic model accounting for imperfect detection that estimates local group abundance and temporal rates of change, to assess whether habitat protection explained trends of the arboreal and diurnal Peters' Angola colobus (Colobus angolensis palliatus), Udzungwa red colobus (Piliocolobus gordonorum), and Tanzania Sykes' monkey (Cercopithecus mitis subsp. moloneyi). We targeted populations occurring in two forest blocks with contrasting protection regimes, with one block impacted by targeted poaching of the two Colobus species. We found that these latter species were much less abundant in the more human impacted forest, underwent a rapid decline, and subsequently remained at low abundance and without signs of recovery once this forest was granted greater protection. Instead, Sykes' abundance did not differ between forests, and improved protection was associated with a slight increase in abundance. Age class composition for social groups of both Colobus species differed between forests, indicating altered births and survivorship rates in the impacted forest. Results suggest that targeted hunting can prevent recovery for several years and even after increased habitat protection. Our approach appears valuable to monitor population dynamics over the long term, highlighting species-specific variations in both vulnerability to anthropogenic disturbance and recovery patterns in primate populations.     

Demographic Variation Among Udzungwa Red Colobus in Relation to Gross Ecological and Sociological Parameters

We evaluated variation in group size and composition of Udzungwa red colobus (Procolobus gordonorum) in relation to gross-habitat and sociological parameters. The endangered species is endemic to the Udzungwa Mountains and nearby forests in the Kilombero Valley of south-central Tanzania. We counted 63 groups in 10 forests, ranging in altitude from 250 to 1,800 m. Group size ranged from 7 to 83 (x = 36.3) individuals and adult sex ratios (females/male) ranged from 1.5 to 7.3 (x = 3.5), excluding solitary individuals. Group size was influenced by several habitat parameters, including tree density, degree of deciduousness, and forest size. Groups were largest in large blocks of mature, moist, mixed evergreen and semideciduous forests, but group size is not correlated with altitude. Groups in a highly degraded forest appeared to have fission-fusion societies. The effect of habitat quality on age-sex composition of groups was most apparent in natality and less so in survivorship of adult females and juveniles. The number of adult males in groups accounted for 50% of the variance in group size and 34–39% of the variance in numbers of adult females in groups. Habitat quality affects natality more than demographic parameters do. Groups with a low proportion of adult females had greater juvenile survivorship, perhaps because of lower food competition between these two classes. We proffer hypotheses based on our study and previous intertaxonal comparisons to explain cause and effect relationships between habitat quality and demography. Finally, our study demonstrates the importance of large blocks of old-growth forest to the conservation of Udzungwa red colobus, and we make recommendations for conservation and research.     

Abundance of Diurnal Primates in Mwanihana Forest, Udzungwa Mountains, Tanzania

Many individual researchers have used line transect counts to estimate forest primate abundance. They have devoted less attention to the interpretation of line transect data obtained by several observers, as is often the case in long-term monitoring programs. We present primate relative abundance data that 5 observers collected over 6 yr (not continuous) along 4 different transects each 4 km long in the Mwanihana Forest, Udzungwa Mountains National Park, Tanzania. Total distance walked during transect repetitions is ca. 700 km. The species we saw most frequently was the endemic Udzungwa red colobus Procolobus gordonorum (mean 0.59 groups/km walked), followed by the Angolan black-and-white colobus Colobus angolensis (0.43 groups/km) and Sykes’s monkey Cercopithecus mitis (0.35 groups/km). We sighted the endemic Sanje mangabey Cercocebus galeritus sanjei and the yellow baboon Papio cynocephalus infrequently, the latter being confined to the deciduous forest parts of the transects. We analyzed sighting frequency by gross habitat type, transect, season, and observer. Interobserver differences in the relative abundance of each species were moderate and the few cases of significant variations were due to discordance of only 1 observer from the others. Estimated distances of primate group sightings differ significantly among observers, thus preventing us from deriving estimates of absolute density. Frequency distributions of distance-class intervals are not significantly different among observers, which may indicate gross interobserver consistency in the width of the area sampled. We conclude that unless consistency in data collection is checked, as we did for 2 observers who collected data simultaneously, potential interobserver differences remain an underlying source of variance in the results that cannot be separated from other sources of variance.     

Primates Decline Rapidly in Unprotected Forests: Evidence from a Monitoring Program with Data Constraints

Growing threats to primates in tropical forests make robust and long-term population abundance assessments increasingly important for conservation. Concomitantly, monitoring becomes particularly relevant in countries with primate habitat. Yet monitoring schemes in these countries often suffer from logistic constraints and/or poor rigor in data collection, and a lack of consideration of sources of bias in analysis. To address the need for feasible monitoring schemes and flexible analytical tools for robust trend estimates, we analyzed data collected by local technicians on abundance of three species of arboreal monkey in the Udzungwa Mountains of Tanzania (two Colobus species and one Cercopithecus), an area of international importance for primate endemism and conservation. We counted primate social groups along eight line transects in two forest blocks in the area, one protected and one unprotected, over a span of 11 years. We applied a recently proposed open metapopulation model to estimate abundance trends while controlling for confounding effects of observer, site, and season. Primate populations were stable in the protected forest, while the colobines, including the endemic Udzungwa red colobus, declined severely in the unprotected forest. Targeted hunting pressure at this second site is the most plausible explanation for the trend observed. The unexplained variability in detection probability among transects was greater than the variability due to observers, indicating consistency in data collection among observers. There were no significant differences in both primate abundance and detectability between wet and dry seasons, supporting the choice of sampling during the dry season only based on minimizing practical constraints. Results show that simple monitoring routines implemented by trained local technicians can effectively detect changes in primate populations in tropical countries. The hierarchical Bayesian model formulation adopted provides a flexible tool to determine temporal trends with full account for any imbalance in the data set and for imperfect detection.     

Tana river red colobus and crested mangabey: Results of recent censuses

Censuses of the Tana River red colobus (Colobus badius rufomitratus) and crested mangabey (Cercocebus galeritus galeritus), primate species endemic to riverine forests of the lower Tana River in eastern Kenya, showed significant population declines for both species between the mid-1970s and 1980s. Red colobus declined in terms of group size and number of groups per forest; the mangabey population declined only in numbers of groups per forest. There was no significant change in mean group age/sex composition between the two time periods for either species. Differences in the feeding and ranging ecology of the two species may explain why the mangabey population suffered a less severe decline than the red colobus. Similarities in census results over 3 years in the late 1980s suggest that the primate populations are no longer decreasing.     

Vegetative predictors of primate abundance: utility and limitations of a fine-scale analysis

Determining ecological predictors of primate abundance is important for both theoretical and applied conservation management. For forest primates, research has focused on comparisons of primate abundance and vegetation in different forest blocks or forest compartments with different management histories. However, great variation in primate abundance often occurs within single forests, especially in mountainous areas or in areas with habitat mosaics due to past disturbance. Here we assess, for the first time, the usefulness and limitations of small-scale, within-transect analysis of vegetative parameters as predictors of primate abundance in a very heterogeneous forest habitat in the Udzungwa Mountains of Tanzania. Relative abundance of four species of diurnal primates was recorded over a period of 2.5 years by walking three census transects 48 times each. Tree size, density, species composition, and food plants were measured along the same census lines. The fine-scale relationship between primate abundance and vegetative variables was analyzed through generalized linear modeling applied to 58 segments of these three census lines. Each segment was 200 m in length. For all four primate species, we found significant associations between their abundance and selected vegetative variables. The abundance of the endemic and endangered Udzungwa red colobus Procolobus gordonorum was positively related to mean basal area of large trees (diameter at breast height greater than 20 cm) and to the species richness of their food plants. Considering the very great variation in primate abundance that was recorded among segments of the census lines, our approach proved useful in predicting the relationship between primate abundance and small-scale habitat differences. The main limitation of this study, however, was the relatively low-predictive power of the models for some species, especially the Angolan colobus Colobus angolensis. We discuss the potential reasons for this problem and suggest possible improvements for future studies.     

Primates in Human-Modified and Fragmented Landscapes: The Conservation Relevance of Modelling Habitat and Disturbance Factors in Density Estimation

Accurate density estimations of threatened animal populations is essential for management and conservation. This is particularly critical for species living in patchy and altered landscapes, as is the case for most tropical forest primates. In this study, we used a hierarchical modelling approach that incorporates the effect of environmental covariates on both the detection (i.e. observation) and the state (i.e. abundance) processes of distance sampling. We applied this method to already published data on three arboreal primates of the Udzungwa Mountains of Tanzania, including the endangered and endemic Udzungwa red colobus (Procolobus gordonorum). The area is a primate hotspot at continental level. Compared to previous, ‘canonical’ density estimates, we found that the inclusion of covariates in the modelling makes the inference process more informative, as it takes in full account the contrasting habitat and protection levels among forest blocks. The correction of density estimates for imperfect detection was especially critical where animal detectability was low. Relative to our approach, density was underestimated by the canonical distance sampling, particularly in the less protected forest. Group size had an effect on detectability, determining how the observation process varies depending on the socio-ecology of the target species. Lastly, as the inference on density is spatially-explicit to the scale of the covariates used in the modelling, we could confirm that primate densities are highest in low-to-mid elevations, where human disturbance tend to be greater, indicating a considerable resilience by target monkeys in disturbed habitats. However, the marked trend of lower densities in unprotected forests urgently calls for effective forest protection.     

A novel landscape genetic approach demonstrates the effects of human disturbance on the Udzungwa red colobus monkey (Procolobus gordonorum)

A comprehensive understanding of how human disturbance affects tropical forest ecosystems is critical for the mitigation of future losses in global biodiversity. Although many genetic studies of tropical forest fragmentation have been conducted to provide insight into this issue, relatively few have incorporated landscape data to explicitly test the effects of human disturbance on genetic differentiation among populations. In this study, we use a newly developed landscape genetic approach that relies on a genetic algorithm to simultaneously optimize resistance surfaces to investigate the effects of human disturbance in the Udzungwa Mountains of Tanzania, which is an important part of a universally recognized biodiversity hotspot. Our study species is the endangered Udzungwa red colobus monkey (Procolobus gordonorum), which is endemic to the Udzungwa Mountains and a known indicator species that thrives in large and well-protected blocks of old growth forest. Population genetic analyses identified significant population structure among Udzungwa red colobus inhabiting different forest blocks, and Bayesian cluster analyses identified hierarchical structure. Our new method for creating composite landscape resistance models found that the combination of fire density on the landscape and distance to the nearest village best explains the genetic structure observed. These results demonstrate the effects that human activities are having in an area of high global conservation priority and suggest that this ecosystem is in a precarious state. Our study also illustrates the ability of our novel landscape genetic method to detect the impacts of relatively recent landscape features on a long-lived species.     

Impact of Commercial Hunting on Monkey Populations in the Taï region,Côte d'Ivoire1

We studied the impact of hunting on monkey species in the Taï National Park and adjacent forests in Côte d'Ivoire. The average wild meat consumption per capita per year was assessed from market surveys and interviews. We determined that the amount of primate wild meat being extracted in the Taï National Park and surrounding forests was 249 t in 1999. Hunting pressure was the highest on the larger primate species such as red colobus, Procolobus badius, black and white colobus, Colobus polykomos, and the sooty mangabey, Cercocebus torquatus atys. Estimates of population densities were based on line transect surveys. The maximum annual production of each species was calculated using the Robinson and Redford model (1991) and assuming unhunted conditions. Comparing current harvest levels with the maximum sustainable yield suggests that harvest of red colobus monkeys (Procolobus badius) is sustainable, whereas current off‐take of the black and white colobus (Colobus polykomos), the sooty mangabey (Cercocebus atys), diana monkeys (Cercopithecus diana), and Campbell's monkeys (Cercopithecus campbelli) exceeds sustainability by up to three times.     

Group Size Dynamics over 15+ Years in an African Forest Primate Community

Group size affects many aspects of the ecology and social organization of animals. We investigated group size stability for five primate species in Kibale National Park, Uganda from 1996 to 2011 at three nested spatial scales. Survey data indicated that group sizes did not change for most species, with the exception of red colobus monkeys (Procolobus rufomitratus), in which group size increased at all spatial scales. Mangabey (Lophocebus albigena) group size increased in old‐growth forest, but the sample size and increase were small. To augment this survey data, we collected several years of demographic data on three habituated groups of redtail monkeys (Cercopithecus ascanius), eight groups of black‐and‐white colobus (Colobus guereza), and one red colobus group. The red colobus group increased from 59 to 104 individuals, while redtail monkey and black‐and‐white colobus group sizes were stable, mirroring our survey results. To understand mechanisms behind group size changes in red colobus versus stability in other primates, we monitored forest dynamics at two spatial scales between 1990 and 2013, considered changes in predator population, and explored evidence of disease dynamics. The cumulative size of all trees and red colobus food trees increased over 24 yr, suggesting that changing food availability was driving group size changes for red colobus, while predation and disease played lesser roles. Overall, our results and evidence of changing primate densities suggest that the Kibale primate community is in a non‐equilibrium state. We suggest future conservation and management efforts take this into consideration.     

Population Sizes and Distribution of Primates in the Lower Tana River Forests,Kenya

We studied the population size and distribution of diurnal primates in the lower Tana River forests, Kenya. They are the only remaining habitats for 2 threatened primates: the Tana River red colobus (Procolobus rufomitratus) and the Tana River crested mangabey (Cercocebus galeritus galeritus). We conducted censuses in 73 forest patches from January through March 2001. We estimate population size of the red colobus to be 788 individuals in 82 groups and that of the crested mangabeys to be 2,070 individuals in 59 groups. The data suggest that over a 7-year period (1994-2001), there was an 18% increase in the crested mangabey population and a 5% decline in red colobus numbers. Further, the red colobus range has expanded both north and south, whereas that of crested mangabeys has only expanded south. Fifty-six percent of crested mangabeys and 46% of red colobus groups were inside the Tana River Primate National Reserve (TRPNR). Other primates encountered included 170 groups of Sykes' monkeys (Cercopithecus mitis), 70 groups of yellow baboons (Papio cynocephalus) and 4 groups of grivets [Chlorocebus (Cercopithecus) aethiops]. Mean group densities of the 2 endangered primates and of baboons were higher inside than outside the TRPNR, reinforcing the importance of TRPNR for their conservation. An intervention program is required to stem further decline in the red colobus population and to protect small isolated groups in forest patches outside TRPNR.     

Ecology of the Zanzibar Red Colobus Monkey: Demographic Variability and Habitat Stability

We examined the Zanzibar red colobus' (Procolobus kirkii) social structure and population dynamics in relation to the density, diversity and dispersion of food resources in ground-water forest and agricultural land, which we characterized in terms of red colobus food species density, diversity, basal area and dispersion. We used transect sampling and group follows to describe population dynamics and social systems. Two agricultural areas, SJF Shamba and Pete Village, had higher densities and more uniformly dispersed red colobus food tree species than those of the ground-water forest. Red colobus at these two sites had greater population densities and natality, and smaller home ranges than red colobus in the ground-water forest. However, these findings apply to a very small area of agricultural land (approximately 18 ha) that is contiguous with an area of the forest reserve having a high density of red colobus. It is not representative of agricultural areas elsewhere on Zanzibar which support much lower densities or no red colobus. Although agricultural areas contiguous with the forest reserve had high densities of red colobus, they appear to be very unstable. Within the agricultural areas, we observed higher intergroup variation in group size and composition, study groups that decreased dramatically in size and disappeared from the study site, significantly lower levels of juvenile recruitment, and red colobus food trees that exhibited definite signs of overbrowsing. This apparent instability in the subpopulation of red colobus utilizing agricultural systems probably reflects the lower basal area of food trees and the greater fragmentation of suitable habitat and floristic dynamics due to human activities in these areas. A fusion-fission social system occurred only in the ground-water forest subpopulation, which we hypothesize to be due to highly clumped food resources.     

Patch depletion behavior differs between sympatric folivorous primates

Food competition in group-living animals is commonly accepted as a critical determinant of foraging strategies and social organization. Here we examine food patch depletion behavior in a leaf-eating (folivorous) primate, the guereza (Colobus guereza). Snaith and Chapman (2005) studied the sympatric folivorous red colobus (Procolobus rufomitratus), which shares many food resources with the guereza. They determined that red colobus deplete the patches (feeding trees) they use, while we found contrary evidence for guerezas using the same methods. We found that the time guerezas spent feeding in a patch was affected by neither tree size, an indicator of food abundance, nor the size of the feeding group, an indicator of feeding competition. For their principal food item (young leaves), intake rate remained constant and coincided with a decrease in the distance moved to find food within a patch, implying that guerezas do not deplete patches. This points to a fundamental difference in the use of food by guerezas and red colobus, which may be linked to the large difference in their group sizes and/or to a disparity in their digestive physiologies. However, further analyses revealed that the number of feeders within a patch did not affect patch depletion patterns in either species, leaving the potential for a physiological basis as the most plausible explanation. Our research highlights the need for a more critical examination of folivorous primate feeding ecology and social behavior, as all folivorous primates are typically lumped into a single category in socioecological models, which may account for conflicting evidence in the literature.     

Estimating the population size of the Sanje mangabey (Cercocebus sanjei) using acoustic distance sampling

The Sanje mangabey (Cercocebus sanjei) is endemic to the Udzungwa Mountains, Tanzania, and is classified as Endangered due to its putatively declining population size, habitat degradation and fragmentation. Previous population size estimates have ranged from 1,350 to 3,500 individuals, with the last direct survey being conducted 15 years before the present study. Previous estimates are now thought to have underestimated the population due to a limited knowledge of group and habitat size, nonsystematic approaches and the use of visual methods that are not suitable for surveying the Sanje mangabey with its semi-terrestrial and elusive behaviors. We used an acoustic survey method with observers recording the distinctive “whoop-gobble” vocalization produced by mangabeys and point transect distance sampling to model a detection function and estimate abundance. Twenty-eight surveys were conducted throughout the two forests where Sanje mangabeys are found: Mwanihana forest in the Udzungwa Mountains National Park (n = 13), and the Uzungwa Scarp Nature Reserve (n = 15). Group density was found to be significantly lower in the relatively unprotected Uzungwa Scarp forest (0.15 groups/km²; 95% CI: 0.08–0.27) compared to the well-protected Mwanihana forest (0.29 groups/km²; 95% CI: 0.19–0.43; p = .03). We estimate that there are 1,712 (95% CI: 1,141–2,567) individuals in Mwanihana and 1,455 (95% CI: 783–2,702) in the Uzungwa Scarp, resulting in a total population size of 3,167 (95% CI: 2,181–4,596) individuals. The difference in group density between sites is likely a result of the differing protection status and levels of enforcement between the forests, suggesting that protection of the Uzungwa Scarp should be increased to encourage recovery of the population, and reduce the threat of degradation and hunting. Our results contribute to the reassessment of the species' IUCN Red List status and informing management and conservation action planning.     

Group size in folivorous primates: ecological constraints and the possible influence of social factors

The ecological-constraints model assumes that food items occur in depletable patches and proposes that an increase in group size leads to increased day range due to more rapid patch depletion. Smaller groups become advantageous when an increase in travel costs is not repaid by an increase in energy gained or some other fitness advantage. On the other hand, we also know that group size can be influenced by social factors. Here we contrast the diet and group size of red colobus (Procolobus badius) and black-and-white colobus (Colobus guereza) in Kibale National Park, Uganda to consider how ecological and social factors are affecting their group sizes. Subsequently, we examine whether the insights gained from this detailed comparison can provide an understanding of why the social organization and group size of mantled howlers (Alouatta palliata) and black howlers (A. pigra) differ. Two groups of red colobus and two groups of black-and-white colobus were studied over 10 months. Red colobus groups were larger (48 and 24) than black-and-white colobus groups (9 and 6). The two groups of red colobus overlap home ranges with the two groups of black-and-white colobus; 75% and 95% of their home ranges were within red colobuss home range. There was a great deal of similarity in the plant parts eaten by the two species and both species fed primarily on young leaves (red colobus 70%, black-and-white colobus 76%). In terms of the actual species consumed, again there was a great deal of similarity between species. The average dietary overlap among months for the two neighboring groups of red colobus was 37.3%, while the dietary overlap between the red colobus and the black-and-white colobus group that had its home range almost entirely within the home range of the red colobus groups averaged 43.2% among months. If ecological conditions were responsible for the difference in group size between the two colobine species, one would expect the density of food trees to be lower in the home ranges of the black-and-white colobus monkeys, since they have the smaller group size. We found the opposite to be true. Both black-and-white colobus groups had more food trees and the cumulative size of those trees was greater than those in the red colobuss home ranges. We quantify how these differences parallel differences in mantled and black howlers. The average group size for mantled howlers was 12.9 individuals, and for black howlers it was 5.3 individuals. We explore possible social constraints, such as infanticide, that prevent black-and-white colobus and black howlers from living in large groups.This revised version was published online in April 2005 with corrections to the cover date of the issue.     

Coping with Forest Fragmentation: The Primates of Kibale National Park,Uganda

A goal of conservation biology is to determine which types of species are most susceptible to habitat disturbance and which types of disturbed habitats can support particular species. We studied 20 forest fragments outside of Kibale National Park, Uganda, to address this question. At each patch, we determined the presence of primate species, tree species composition, patch size, and distance to nearest patch. We collected demographic, behavioral, and dietary data for Abyssinian black-and-white colobus (Colobus guereza). Black-and-white colobus and red-tailed guenons (Cercopithecus ascanius) were in almost all fragments; Pennant's red colobus (Procolobus pennantii) and chimpanzees (Pan troglodytes) were in some fragments; and blue monkeys (Cercopithecus mitis) and gray-cheeked mangabeys (Lophocebus albigena) were absent from all fragments. No species characteristics—home range, body size, group size, or degree of frugivory—predicted the ability of species to live in patches. No characteristics of patches—area, distance to the nearest patch, distance to Kibale, or number of food trees present—predicted the presence of a particular species in a patch, but distance to Kibale may have influenced presence of red colobus. Black-and-white colobus group size was significantly smaller in the forest patches than in the continuous forest of Kibale. For a group of black-and-white colobus in one patch, food plant species and home range size were very different from those of a group within Kibale. However, their activity budget and plant parts eaten were quite similar to those of the Kibale group. The lack of strong predictive variables as well as differences between other studies of fragmentation and ours caution against making generalizations about primate responses to fragmentation.     

Differences in abundance and species richness between shrews and rodents along an elevational gradient in the Udzungwa Mountains,Tanzania

Small mammals (shrews and rodents) were surveyed along an elevational transect in the Udzungwa Scarp Forest Reserve, in the Udzungwa Mountains, Tanzania. Trap lines and pitfall lines were installed at 600, 910, 1460, and 2000 m a.s.l. In a total of 10341 sample nights (7448 trap-nights and 2893 bucket-nights) 343 specimens (148 shrews, 205 rodents) were captured representing 9 shrew and 14 rodent species for a total of 23 species. While overall species diversity generally increased with elevation, this pattern was not constant for each group sampled. For rodents, both species richness and abundance were lowest at 600 m and greatest at 2000 m a.s.l., and were significantly correlated with elevation. While the highest species number and abundance for shrews was at 2000 m, there was no correlation of these two values with elevation. Rainfall appears to have affected the capture of shrews, but not rodents, and capture success of individual buckets and traps indicated a lack of capture independence. Eastern Arc endemics such asCrocidura desperata Hutterer, Jenkins and Verheyen, 1991 andMyosorex kihaulei Stanley and Hutterer, 2000 were more abundant at 2000 m a.s.l., than at lower elevations. Implications of results of this survey for analyses of future biotic surveys are discussed.     

Thirty Years of Research in Kibale National Park,Uganda, Reveals a Complex Picture for Conservation

Kibale National Park, Uganda, has a rich and abundant primate community and a complicated history of anthropogenic disturbance. Moreover, it has been the focus of over 30 yr of research and has received considerable attention from nongovernmental and governmental conservation organizations. As a result, Kibale serves as a valuable case study with which to evaluate the factors that regulate primate population density and the challenges of deriving generalizations for conservation. We review the impact of logging and forest fragmentation on primate population density and trace the efficacy of various conservation strategies. A 28-yr comparison of primate abundance in logged and unlogged forests and a 10-yr study of forest dynamics showed that primate recovery in logged areas is generally slow or not occurring at all for some species, which is likely driven by the fact that the forest is not recovering as expected. No primate species characteristic predicted their ability to live in forest fragments around Kibale. While a nutritional model was useful to predict the abundance of colobus in forest fragments outside of Kibale, a 5-yr study revealed that human land-use practices are more fundamentally shaping population dynamics. We evaluate data on primate abundance in light of Milton’s protein/fiber model to predict colobine biomass. We demonstrate that while the model can predict colobus biomass in pristine habitats, the 2 colobus species respond differently to disturbance. We offer suggestions for future conservation research and consider strategies to conserve forested national parks based on experiences gained over 30 yr.     

Primate population dynamics: variation in abundance over space and time

The rapid disappearance of tropical forests, the potential impacts of climate change, and the increasing threats of bushmeat hunting to wildlife, makes it imperative that we understand wildlife population dynamics. With long-lived animals this requires extensive, long-term data, but such data is often lacking. Here we present longitudinal data documenting changes in primate abundance over 45 years at eight sites in Kibale National Park, Uganda. Complex patterns of change in primate abundance were dependent on site, sampling year, and species, but all species, except blue monkeys, colonized regenerating forest, indicating that park-wide populations are increasing. At two paired sites, we found that while the primate populations in the regenerating forests had increased from nothing to a substantial size, there was little evidence of a decline in the source populations in old-growth forest, with the possible exception of mangabeys at one of the paired sites. Censuses conducted in logged forest since 1970 demonstrated that for all species, except black-and-white colobus, the encounter rate was higher in the old-growth and lightly-logged forest than in heavily-logged forest. Black-and-white colobus generally showed the opposite trend and were most common in the heavily-logged forest in all but the first year of monitoring after logging, when they were most common in the lightly-logged forest. Overall, except for blue monkey populations which are declining, primate populations in Kibale National Park are growing; in fact the endangered red colobus populations have an annual growth rate of 3%. These finding present a positive conservation message and indicate that the Uganda Wildlife Authority is being effective in managing its biodiversity; however, with constant poaching pressure and changes such as the exponential growth of elephant populations that could cause forest degradation, continued monitoring and modification of conservation plans are needed.