Survey of Grauer's Gorillas (Gorilla gorilla graueri) and Eastern Chimpanzees (Pan troglodytes schweinfurthi) in the Kahuzi-Biega National Park Lowland Sector and Adjacent Forest in Eastern Democratic Republic of Congo

We describe the distribution and estimate densities of Grauer's gorillas (Gorilla gorilla graueri) and eastern chimpanzees (Pan troglodytes schweinfurthi) in a 12,770-km ² area of lowland forest between the Lowa, Luka, Lugulu, and Oku rivers in eastern Democratic Republic of Congo, the site of the largest continuous population of Grauer's gorillas. The survey included a total of 480 km of transects completed within seven sampling zones in the Kahuzi-Biega National Park lowland sector and adjacent Kasese region and approximately 1100 km of footpath and forest reconnaissance. We estimate total populations of 7670 (4180–10,830) weaned gorillas within the Kahuzi-Biega lowland sector and 3350 (1420–5950) individuals in the Kasese survey areas. Within the same area, we estimate a population of 2600 (1620–4500) chimpanzees. Ape nest site densities are significantly higher within the Kahuzi-Biega lowland sector than in the more remote Kasese survey area in spite of a significantly higher encounter rate of human sign within the lowland sector of the park. Comparison of our data with information obtained by Emlen and Schaller during the first rangewide survey of Grauer's gorillas in 1959 suggests that gorilla populations have remained stable in protected areas but declined in adjacent forest. These findings underscore the key role played by national parks in protecting biological resources in spite of the recent political and economic turmoil in the region. We also show that forest reconnaissance is a reliable and cost-effective method to assess gorilla densities in remote forested areas.     

Population structure and group composition of western lowland gorillas in North‐Western Republic of Congos

Population studies are an essential part of conservation actions. Under exceptional observation conditions we studied a western lowland gorilla population visiting the Maya salt‐clearing (north of the Parc national d'Odzala, P.N.O., Congo) over an 8 month period; 36 groups and 18 solitary individuals (a total of 420 individuals) have been identified visiting the clearing, which suggests a high gorilla density in the region. Ninety‐six percent of the gorillas entered the clearing in groups. One‐male groups had a mean size of 11.2. Ninety percent of solitary individuals were silver‐back males. Compared with other populations of both lowland gorillas and mountain gorillas, the Maya population had the highest immature rate and the highest number of infants per female. Ecological correlates that could explain the attractiveness of the Maya clearing are discussed. The present status and the renewal rate of the Maya population indicate the need for further studies and confirm the importance of developing eco‐tourism in this region as part of the sustainable park management activities developed by the ECOFAC programme (European Union). The results also provide arguments to support the proposal for extending the P.N.O. to include this region, which is rich in salt‐clearings and attracts many other key‐species of mammal such as forest elephants. Am. J. Primatol. 48:1–14, 1999. © 1999 Wiley‐Liss, Inc.     

An empirical evaluation of the African elephant as a focal species for connectivity planning in East Africa

Aim Large, charismatic and wide‐ranging animals are often employed as focal species for prioritizing landscape linkages in threatened ecosystems (i.e. ‘connectivity conservation’), but there have been few efforts to assess empirically whether focal species co‐occur with other species of conservation interest within potential linkages. We evaluated whether the African elephant (Loxodonta africana), a world‐recognized flagship species, would serve as an appropriate focal species for other large mammals in a potential linkage between two major protected area complexes. Location A 15,400 km² area between the Ruaha and Selous ecosystems in central Tanzania, East Africa. Methods We used walking transects to assess habitat, human activity and co‐occurrence of elephants and 48 other large mammal species (> 1 kg) at 63 sites using animal sign and direct sightings. We repeated a subset of transects to estimate species detectability using occupancy modelling. We used logistic regression and AIC model selection to characterize patterns of elephant occurrence and assessed correlation of elephant presence with richness of large mammals and subgroups. We considered other possible focal species, compared habitat‐based linear regression models of large mammal richness and used circuit theory to examine potential connectivity spatially. Results Elephants were detected in many locations across the potential linkage. Elephant presence was highly positively correlated with the richness of large mammals, as well as ungulates, carnivores, large carnivores and species > 45 kg in body mass (‘megafauna’). Outside of protected areas, both mammal richness and elephant presence were negatively correlated with human population density and distance from water. Only one other potential focal species was more strongly correlated with species richness than elephants, but detectability was highest for elephants. Main conclusions Although African elephants have dispersal abilities that exceed most other terrestrial mammals, conserving elephant movement corridors may effectively preserve habitat and potential landscape linkages for other large mammal species among Tanzanian reserves.     

Pneumatic osteolysis in a cetacean (Orcaella brevirostris)

Pneumatic osteolysis (excessive pneumatisation of cranial bones) previously reported for certain terrestrial mammals (rhinoceroses, elephants, gorilla) is here reported in the Irrawaddy dolphin (Orcaella brevirostris). The osteological evidence provided by all mammal forms studied indicates unequivocally that, in the complex process of bone pneumatization (whether normal or excessive in range), the initiative lies with the peculiarly specialized mucosa of the pneumatic spaces concerned.     

Diversity of medium and large mammals in the Loka Abaya National Park,southern Ethiopia

We evaluated the richness, diversity, and composition of the medium and large mammal community in the Loka Abaya National Park (LANP), southern Ethiopia, and how these parameters differ among four habitat types: wooded grassland, riverine forest, hilly scrubland and wetland, and between seasons. We recorded a total of 2,573 individual animals of 28 medium and large mammal species in the park. This included three globally threatened species: the endangered African wild dog (Lycaon pictus), the vulnerable Leopard (Panthera pardus), and Hippopotamus (Hippopothamus amphibius). Season had little effect on species richness, diversity, and composition both across and within habitat types. However, species richness across seasons was significantly different among the four habitat types, in the declining order of the following: wooded grassland > riverine forest > hilly scrubland > wetland. The strongest similarity in species composition, both across and within seasons, was found between wooded grassland and riverine forest. In terms of relative abundance, mammal assemblage of the wooded grassland and wetland habitats had more evenly distributed number of species with different relative abundance categories. Overall, Anubis Baboon (Papio anubis), Grivet Monkey (Chlorocebus aethiops), and Greater Kudu (Tragelephus strepsiceros) were the three most abundant species across habitat types. In conclusion, findings of our study reveal that LANP plays an important role in Ethiopia's mammal conservation. Our findings will serve as baseline information for managers of the park to make effective conservation decisions and as a baseline for researchers wishing to conduct related ecological studies.     

Variation in behavioral traits of two frugivorous mammals may lead to differential responses to human disturbance

Human activities can lead to a shift in wildlife species’ spatial distribution. Understanding the specific effects of human activities on ranging behavior can improve conservation management of wildlife populations in human‐dominated landscapes. This study evaluated the effects of forest use by humans on the spatial distribution of mammal species with different behavioral adaptations, using sympatric western lowland gorilla and central chimpanzee as focal species. We collected data on great ape nest locations, ecological and physical variables (habitat distribution, permanent rivers, and topographic data), and anthropogenic variables (distance to trails, villages, and a permanent research site). Here, we show that anthropogenic variables are important predictors of the distribution of wild animals. In the resource model, the distribution of gorilla nests was predicted by nesting habitat distribution, while chimpanzee nests were predicted first by elevation followed by nesting habitat distribution. In the anthropogenic model, the major predictors of gorilla nesting changed to human features, while the major predictors of chimpanzee nesting remained elevation and the availability of their preferred nesting habitats. Animal behavioral traits (body size, terrestrial/arboreal, level of specialization/generalization, and competitive inferiority/superiority) may influence the response of mammals to human activities. Our results suggest that chimpanzees may survive in human‐encroached areas whenever the availability of their nesting habitat and preferred fruits can support their population, while a certain level of human activities may threaten gorillas. Consequently, the survival of gorillas in human‐dominated landscapes is more at risk than that of chimpanzees. Replicating our research in other sites should permit a systematic evaluation of the influence of human activity on the distribution of mammal populations. As wild animals are increasingly exposed to human disturbance, understanding the resulting consequences of shifting species distributions due to human disturbance on animal population abundance and their long‐term survival will be of growing conservation importance.     

Grasshopper assemblage response to a restored national park (Mountain Zebra National Park, South Africa)

Twelve grassland sites were sampled inside and outside the Mountain Zebra National Park (MZNP), South Africa to assess changes in grasshopper assemblages to grazing by indigenous mammals inside the park compared to grazing by domestic cattle outside. The MZNP has been restored from cattle-grazed farmland to indigenous mammal parkland for 62 years. The number of grasshopper species and families inside the park was not significantly different from outside the park, but the number of individuals inside the park was significantly higher. Multivariate statistics did not reveal any strong site groupings based on simple inside/outside comparisons, but there were clear groupings of sites based on vegetation characteristics and other environmental variables. The park boundary, therefore, does not significantly determine grasshopper assemblages, although intensity of grazing does. The indigenous mammals inside the park had the same effect on grasshoppers as the domestic cattle outside, and it was the level of defoliation and trampling that was important rather than the type of mammal. Intensive livestock grazing and trampling leads to bush encroachment and reduction in grass cover and/or disappearance of several grass species. In response to this pressure, grasshopper populations dropped, with localized extirpation of some species. Vegetation composition and structure, particularly grass height and percentage cover, had a significant effect on grasshopper assemblages. The MZNP is thus an area of localized, elevated grasshopper abundance in comparison with the surrounding farms, and presumably represents a situation prior to the current, intensive farming activities. Such elevated grasshopper abundances are important for maintaining soil quality and hence ecological integrity of this landscape which is poor in organics and nitrogen. The MZNP could be viewed as a centre where species with high mobility may seek refuge from anthropogenic pressures. The MZNP also serves as a reference illustrating the differences between restored-through-natural-succession and anthropogenically disturbed habitats, and compares desirable with undesirable ecosystem changes for herbivorous invertebrates such as grasshoppers.     

Community‐based wildlife management area supports similar mammal species richness and densities compared to a national park

Community‐based conservation models have been widely implemented across Africa to improve wildlife conservation and livelihoods of rural communities. In Tanzania, communities can set aside land and formally register it as Wildlife Management Area (WMA), which allows them to generate revenue via consumptive or nonconsumptive utilization of wildlife. The key, yet often untested, assumption of this model is that economic benefits accrued from wildlife motivate sustainable management of wildlife. To test the ecological effectiveness (here defined as persistence of wildlife populations) of Burunge Wildlife Management Area (BWMA), we employed a participatory monitoring approach involving WMA personnel. At intermittent intervals between 2011 and 2018, we estimated mammal species richness and population densities of ten mammal species (African elephant, giraffe, buffalo, zebra, wildebeest, waterbuck, warthog, impala, Kirk's dik‐dik, and vervet monkey) along line transects. We compared mammal species accumulation curves and density estimates with those of time‐matched road transect surveys conducted in adjacent Tarangire National Park (TNP). Mammal species richness estimates were similar in both areas, yet observed species richness per transect was greater in TNP compared to BWMA. Species‐specific density estimates of time‐matched surveys were mostly not significantly different between BWMA and TNP, but elephants occasionally reached greater densities in TNP compared to BWMA. In BWMA, elephant, wildebeest, and impala populations showed significant increases from 2011 to 2018. These results suggest that community‐based conservation models can support mammal communities and densities that are similar to national park baselines. In light of the ecological success of this case study, we emphasize the need for continued efforts to ensure that the BWMA is effective. This will require adaptive management to counteract potential negative repercussions of wildlife populations on peoples' livelihoods. This study can be used as a model to evaluate the effectiveness of wildlife management areas across Tanzania.     

Performance of individual species as indicators for large mammal species richness in Northern Tanzania

In order to prioritize areas for biodiversity conservation, conservation practitioners frequently employ a single species whose distribution is statistically related to overall species richness. However, the performance of single mammal species in terms of (1) their strength, (2) spatial and (3) temporal variability for predicting large mammal species richness has rarely been assessed. Drawing upon data from multiple vehicle-based surveys in four study sites with varying conservation management approaches in the Tarangire–Manyara ecosystem, we assessed the performance of thirteen candidate indicator species. Overall, we found that the association strength between the distribution of single large mammal species and overall large mammal species richness varied (1) considerably across four management units within the same ecosystem, (2) between seasons and (3) years. In contrast to a study carried out in central Tanzania, elephants performed poorly as an indicator of large mammal species richness. Applying our findings to conservation planning, we suggest that information on zebra and wildebeest distribution should be used for delineating corridors for large mammals between protected areas in this ecosystem. The distribution of these two species had a high correlation with overall large mammal species richness, and these correlations were relatively constant throughout time and space. More generally, our study suggests that the performance of indicator species (1) should be assessed across multiple seasons because snapshot surveys may provide biased estimates of indicator performance, (2), cannot necessarily be extrapolated to other ecosystems and (3) should be supplemented by ecological or functional considerations.     

Genetic identification of five strongyle nematode parasites in wild african elephants (Loxodonta africana)

African savannah elephants (Loxodonta africana) are an ecologically and economically important species in many African habitats. However, despite the importance of elephants, research on their parasites is limited, especially in wild populations. Currently, we lack genetic tools to identify elephant parasites. We present genetic markers from ribosomal DNA (rDNA) and mitochondrial DNA (mtDNA) to identify five elephant-specific nematode parasites in the family Strongylidae: Murshidia linstowi, Murshidia longicaudata, Murshidia africana, Quilonia africana, and Khalilia sameera. We collected adult nematodes from feces deposited by wild elephants living in Amboseli National Park, Kenya. Using both morphologic and genetic techniques, we found that the internal transcribed spacer (ITS) region in rDNA provides a reliable marker to distinguish these species of strongyles. We found no evidence for cryptic genetic species within these morphologic species according to the cox-1 region of mtDNA. Levels of genetic diversity in strongyles from elephants were consistent with the genetic diversity seen within other strongyle species. We anticipate that these results will be a useful tool for identifying gastrointestinal nematode parasites in elephants.     

The conservation significance of the proposed Mbaéré-Bodingué national park,Central African Republic,with special emphasis on its primate community

The proposed Mbaéré-Bodingué National Park (872km²) is located in the southwestern part of the Central African Republic and consists of terra firma rain forests and seasonally flooded forests. We assessed the conservation significance of this site using data on wildlife and habitat conditions collected during a comprehensive primate survey. Scores were attributed to a system of variables and sub-variables to assess the conservation value of the proposed park at the national level. We identified a total of 10 diurnal primate species (eight monkeys, two apes) and flooded forests were considered as a key habitat for primates as they hold a greater number of species than terra firma forests. We recorded the presence of 12 mammal species listed under the 2000 IUCN Red List of Threatened Species, including four classified as Endangered (two Primates, one Proboscidae and one Insectivora). We found that the conservation value of the proposed park ranked second out of the 15 existing protected areas of the country. Primate abundance was found to be generally higher than in the nearby Dzanga-Ndoki National Park (the sole other park located in the same vegetation unit), in particular for the species occurring in flooded forests. This highlights the fact that the proposed Mbaéré-Bodingué National Park is an important area for the conservation of species associated with flooded forests, a habitat which is currently under-represented in the national protected area system. Poaching pressure was identified as the main current threat to the integrity of the proposed park.     

The influence of tree canopies and elephants on sub-canopy vegetation in a savannah

The apparent influence of elephants on the structure of savannahs in Africa may be enhanced by management activities, fire and other herbivores. We separated the effect elephants have on grasses, woody seedlings (<0.5 m) and saplings (0.5–2 m) from the effect of tree canopies (canopy effect), and herbivory (park effect). We defined the canopy effect as the differences between plant abundances and diversity indices under tree canopies and 20 m away from these. Our testing of the park effect relied on the differences in the sub-canopy plant indices inside and outside a protected area that supported a range of herbivores. We based our assessment of the elephant effect on sub-canopy vegetation indices associated with elephant induced reductions in tree canopies. The park and canopy effects were more pronounced than the elephant effect. The park effect suppressed the development of woody seedlings into saplings. Conditions associated with tree canopies benefited woody plants, but not the grasses, as their indices were lower under trees. Elephants reducing canopies facilitated grass species tolerant of direct solar radiation. We concluded that management should consider other agents operating in the system when deciding on reducing the impact that elephants may have on vegetation.     

Tijuca National Park: two pioneering restorationist initiatives in Atlantic forest in southeastern Brazil.

As a contribution to the environmental history of the Tijuca National Park, we report on two pioneering restorationist initiatives and list its the mammal species now found in this urban park. The Tijuca National Park (TNP), a 3,200 ha urban park covered by secondary tropical forest, is located within Rio de Janeiro, in southeastern Brazil. The two restorationist initiatives were a pioneer tropical forest restoration project in the nineteenth century and a fauna management project in the 70' s. The mammal list presented here was based on specimens in the Museu Nacional do Rio de Janeiro and on publications. The mammal community of TNP is composed of 49 species, of which 11 are on regional red lists, and four are on the 2003 IUCN Red List of Threatened Species. Occurrence of these threatened species and the park history itself made the TNP a priority site for studying conservation, management, and monitoring. Besides maintaining fauna and flora (including threatened species) diversity, the park benefits the population of Rio de Janeiro by providing water, green areas, and recreational and touristic opportunities.     

Patterns of elephant impact on woody plants in the Hluhluwe-Imfolozi park, Kwazulu-Natal, South Africa

This study identifies patterns of elephant Loxodonta africana africana impacts upon tree species and woody plant communities in Hluhluwe-Imfolozi Park, a South African savannahs/woodlands area. Elephants were reintroduced there from 1981, following more than 80 years of absence. Data were collected in 2003 on elephant impact on woodland in the Park. Different vegetation types were susceptible to different types and levels of damage by elephants, suggesting that elephants will not homogenize the vegetation. Elephants targeted larger stems for all types of damage, with a strong preference for some of the less abundant species such as Albizia versicolor (breaking and toppling) and Cordia caffra and Schotia brachypetala (debarking). Elephant impacts tended to be distributed evenly across the park landscape, irrespective of stem density or proximity to permanent water. Overall, elephants have little impact on slowing or reversing the spread of undesirable woody species, but are having a marked impact on certain less common tree species and larger tree size-classes in the Hluhluwe-Imfolozi Park.     

Elephants of south‐east Angola in war and peace: their decline,re‐colonization and recent status

Angola’s intermittent 27‐year civil war displaced over four million people and decimated wildlife populations. During the 1980s, African elephants (Loxodonta africana Blumenbach) in Angola drew international alarm with reports of 100,000 elephants killed. Luiana Partial Reserve (PR), a conservation area in south‐east Angola, was the military operations centre for UNITA (National Union for the Total Independence of Angola), which used elephant ivory to pay for arms and meat. However, the full impact of the civil war on elephants is uncertain because there are no reliable estimates of Angolan elephant populations. Following the end of the civil war in 2002, our three aerial surveys of Luiana PR indicated that elephant numbers are increasing rapidly, from 366 in January 2004 to 1827 in November 2005, and expanding their range in the Reserve. Concurrently, elephants tagged with satellite collars in northern Botswana and the Caprivi Strip, Namibia, moved into Luiana PR. To facilitate re‐colonization and conservation of elephants and other wildlife in Luiana PR, we recommend: (i) realignment of the veterinary fence on the Botswana–Namibia border; (ii) development of effective land use management and anti‐poaching programmes; (iii) clearing of landmines; (iv) designation of the Reserve a national park; and (v) development of ecotourism and community conservation programmes.     

Newly discovered Bale monkey populations in forest fragments in southern Ethiopia: evidence of crop raiding, hybridization with grivets, and other conservation threats

Until recently, the Bale monkey (Chlorocebus djamdjamensis), an arboreal primate endemic to the southern Ethiopian highlands, remained virtually unstudied, and its distribution pattern inadequately documented. To broaden our knowledge of the species' distribution and abundance, we carried out interviews with local people and total count surveys for Bale monkeys across 67 fragmented forest sites in human-dominated landscapes in the Oromia and Southern Nations, Nationalities, and People's Regions, Ethiopia. From January 2010 to May 2011, we discovered 26 new Bale monkey populations inhabiting forest fragments at elevations ranging from 2,355 to 3,204 m asl. Across these populations, we recorded 37 groups ranging in size from 9 to 29 individuals (Mean = 19.5, SD = 4.5), for a total of 722 individuals. Black-and-white colobus monkeys (Colobus guereza) were sympatric with Bale monkeys at all sites, while grivet monkeys (Chlorocebus aethiops) were found only at sites where Bale monkeys did not occur. All of the newly discovered Bale monkey sites once contained bamboo forest, though at 35% of the sites bamboo forest had been eliminated during the past two decades. The persistence of Bale monkeys at fragmented sites lacking bamboo suggests greater habitat flexibility for the species than previously thought, though the long-term viability of populations both with and without bamboo remains uncertain. Human hunting in response to crop raiding, a behavior the monkeys engaged in at all sites, represents a major threat facing the newly discovered Bale monkey populations. Furthermore, despite their current lack of sympatry, apparently hybrid individuals between Bale monkeys and grivets were noted at three sites, posing yet another potential obstacle to Bale monkey conservation. Community conservation programs aimed at (1) protecting remaining habitat fragments, (2) planting bamboo and trees within and between fragments, and (3) reducing crop raiding represent the only hope for survival of the newly discovered Bale monkey populations.     

Effects of roads on small mammal diversity and abundance in the northern Serengeti,Tanzania

The high biodiversity of small mammal species in the Serengeti ecosystem provides this ecosystem with important conservation value. However, whether the extensive development of roads has negative impacts on the small mammal population has not been tested. Small mammal population diversity and abundance were examined in this study using live trapping at sites close to (experimental) and away (control) from the main gravel road during the short rainy seasons in November and December 2011 and 2012. A total of 138 individuals from three orders representing six families and fourteen species were collected over 4,860 trap nights. There were no significant differences in the species richness, diversity or abundance of small mammals between the control and experimental sites (p > 0.05), suggesting that the current gravel road does not have a significant impact on the small mammal population. These findings were ascribed to the availability of favourable habitats at both distances as a result of little road usage due to poor conditions. Should the road be improved, the control of anthropogenic activities in the area should be given high priority. Continuous monitoring of the small mammal populations in the area is recommended.     

Marsh rabbit mortalities tie pythons to the precipitous decline of mammals in the Everglades

To address the ongoing debate over the impact of invasive species on native terrestrial wildlife, we conducted a large-scale experiment to test the hypothesis that invasive Burmese pythons (Python molurus bivittatus) were a cause of the precipitous decline of mammals in Everglades National Park (ENP). Evidence linking pythons to mammal declines has been indirect and there are reasons to question whether pythons, or any predator, could have caused the precipitous declines seen across a range of mammalian functional groups. Experimentally manipulating marsh rabbits, we found that pythons accounted for 77% of rabbit mortalities within 11 months of their translocation to ENP and that python predation appeared to preclude the persistence of rabbit populations in ENP. On control sites, outside of the park, no rabbits were killed by pythons and 71% of attributable marsh rabbit mortalities were classified as mammal predations. Burmese pythons pose a serious threat to the faunal communities and ecological functioning of the Greater Everglades Ecosystem, which will probably spread as python populations expand their range.     

Effective sociodemographic population assessment of elusive species in ecology and conservation management

Wildlife managers are urgently searching for improved sociodemographic population assessment methods to evaluate the effectiveness of implemented conservation activities. These need to be inexpensive, appropriate for a wide spectrum of species and straightforward to apply by local staff members with minimal training. Furthermore, conservation management would benefit from single approaches which cover many aspects of population assessment beyond only density estimates, to include for instance social and demographic structure, movement patterns, or species interactions. Remote camera traps have traditionally been used to measure species richness. Currently, there is a rapid move toward using remote camera trapping in density estimation, community ecology, and conservation management. Here, we demonstrate such comprehensive population assessment by linking remote video trapping, spatially explicit capture–recapture (SECR) techniques, and other methods. We apply it to three species: chimpanzees Pan troglodytes troglodytes, gorillas Gorilla gorilla gorilla, and forest elephants Loxodonta cyclotis in Loango National Park, Gabon. All three species exhibited considerable heterogeneity in capture probability at the sex or group level and density was estimated at 1.72, 1.2, and 1.37 individuals per km² and male to female sex ratios were 1:2.1, 1:3.2, and 1:2 for chimpanzees, gorillas, and elephants, respectively. Association patterns revealed four, eight, and 18 independent social groups of chimpanzees, gorillas, and elephants, respectively: key information for both conservation management and studies on the species' ecology. Additionally, there was evidence of resident and nonresident elephants within the study area and intersexual variation in home range size among elephants but not chimpanzees. Our study highlights the potential of combining camera trapping and SECR methods in conducting detailed population assessments that go far beyond documenting species diversity patterns or estimating single species population size. Our study design is widely applicable to other species and spatial scales, and moderately trained staff members can collect and process the required data. Furthermore, assessments using the same method can be extended to include several other ecological, behavioral, and demographic aspects: fission and fusion dynamics and intergroup transfers, birth and mortality rates, species interactions, and ranging patterns.