Coalitionary mate guarding by male chimpanzees at Ngogo, Kibale National Park, Uganda

Cooperative mate guarding by males is unusual in mammals and birds, largely because fertilizations are non-shareable. Chimpanzees live in fission-fusion communities that have cores of philopatric males who cooperate in inter-group aggression and in defending access to the females in their community. Male contest mating competition is restrained within communities, but single high-ranking males sometimes try to mate guard estrous females. Data from an unusually large chimpanzee commmunity at Ngogo, Kibale National Park, Uganda, that contains more males than any previously studied community show new variation in chimpanzee mate-guarding behavior. Contrary to expectation given the large number of males, mate guarding was as common as, or more common than, at other sites, and males other than the alpha male guarded more often. More strikingly, pairs or trios of top-ranking males sometimes engaged in cooperative aggression to prevent estrous females from mating with other males, but tolerated each other's mating activities. Both single males and coalitions mostly guarded periovulatory females. Mate-guarding coalitions were previously unknown in chimpanzees. Coalitions occurred in large mating parties, seemingly because these often contained too many males for single males to maintain exclusive access to estrous females. Coalition members gained higher shares of copulations than they could have expected from solo mate guarding, and suffered lower per capita costs of guarding (as inferred from aggression rates). Two males who most often participated in coalitions formed two-male coalitions at about the point where the number of males present made it unlikely that either could get 50% or more of total copulations on his own, and formed trios when this value dropped below 33%. Kin selection could be a factor in cooperation among male chimpanzees, but coalition members were not necessarily close relatives and the apparent structure of payoffs fit that of mutualism. Furthermore, reliance of male chimpanzees on support from allies to maintain high rank could have led to trading of mating exclusivity for support against mating competitors. Received: 28 May 1997 / Accepted after revision: 16 May 1998     

Interdemic variation in mixed-species association patterns: common diurnal primates of Kibale National Park, Uganda

We used interdemic variation in the tendency to form mixed-species groups to examine the costs and benefits of association among the primates of Kibale National Park, Uganda. A year-long survey of six sites revealed that the amount of time that the five common diurnal primates [red colobus (Procolobus tephrosceles), black-and-white colobus (Colobus guereza), redtail monkeys (Cercopithecus ascanius), blue monkeys (Cercopithecus mitis), and grey-cheeked mangabeys (Lophocebus albigena)] spent in mixed-species groups varied dramatically among sites. In many cases, the proportion of time that species associated was positively related to their densities. By using detailed behavioral observations of redtail monkeys and red colobus made over 4 years (2660 h) at four sites, we were able to reject the null hypothesis that associations occur by chance for only one of four sites. However, a correlative approach exploring the costs and benefits of association suggests that ecological variables do influence association patterns. We found that redtail monkeys and red colobus overlapped in diet (19.2% of their foraging effort) and traveled further when in mixed-species groups than when alone. Having demonstrated this, we examined the applicability of the ecological constraints model for predicting the proportion of the time spent in mixed-species groups based on food availability. For this analysis we concentrated on red colobus from the site with 35 months of observation and demonstrated that their tendency to be in mixed- species groups was related to food availability. We used two methods to examine if mixed-species associations function to decrease predation risk. First, chimpanzees are known to prey heavily on red colobus, but rarely kill other primates. The time red colobus spent in mixed-species groups was correlated to chimpanzee density, but it was not for the other monkey species, suggesting that mixed-species groups serve to decrease predation risk. Second, when red colobus groups contain more infants and are presumably at the greatest risk of predation, they form mixed-species groups most often. These results demonstrate that the costs and benefits of mixed-species associations vary dramatically over small spatial and temporal scales. If such variation is generally the case, then studies conducted at different locations or different times could easily highlight the importance of difference selective agents in favoring mixed-species associations. Received: 10 February 1999 / Received in revised form: 16 September 1999 / Accepted: 2 October 1999     

Predatory behavior of crowned hawk-eagles (Stephanoaetus coronatus) in Kibale National Park, Uganda

Evaluating the nature and significance of predation on populations of wild primates has been difficult given a paucity of data regarding the phenomenon. Here we addressed this problem in a 37-month study of the predatory behavior of crowned hawk-eagles living at the Ngogo study site in Kibale National Park, Uganda. We collected prey remains underneath the nests of two pairs of eagles and census data on potential prey species to investigate prey selection and the ecological impact of predation on the Ngogo primate population. Results indicate that primates form the vast majority of all prey items. Eagles prey selectively on monkeys according to sex and species. Male primates were taken more often than females, while two species, redtail monkeys and mangabeys, were captured significantly more and less, respectively, than chance expectation. In addition, there was no bias in the age of prey: adult and non-adults were killed in numbers roughly equal to their proportional representations in the forest. Further analyses indicate that a non-trivial fraction of the entire primate population at Ngogo succumbs to crowned hawk-eagle predation each year. These results reveal both parallels and contrasts with those reported previously. Some of the parallels are due to similarities in prey availability, while contrasts are likely related to methodological differences between studies, inter- individual variations in predator hunting styles, and differences in prey abundance, demography, and behavior. Received: 29 March 2000 / Revised: 6 June 2000 / Accepted: 15 October 2000     

Male chimpanzees compromise the foraging success of their mates in Kibale National Park,Uganda

Sexual conflict develops when the optimal reproductive strategy for one sex inflicts fitness costs upon the other sex. Among species with intense within-group feeding competition and high costs of reproduction, females are expected to experience reduced foraging efficiency by associating with males, and this may compromise their reproductive ability. Here, we test this hypothesis in chimpanzees, a species with flexible grouping patterns in which female avoidance of large subgroups has been attributed to their relatively high costs of grouping. In an >11-year study of the Kanyawara community of East African chimpanzees (Pan troglodytes schweinfurthii) in the Kibale National Park, Uganda, the availability of sexually receptive females was a key determinant of the number of males in parties. In turn, females experienced significantly lower C-peptide of insulin levels, indicative of reduced energy balance, during periods when they associated with more males. Female associates did not produce the same negative effect. C-peptide levels positively and significantly predicted female ovarian steroid production, indicating that the costs of associating with males can lead to downstream reproductive costs. Therefore, we conclude that Kanyawara chimpanzees exhibit sexual conflict over subgroup formation, with the large groupings that allow males to compete for mating opportunities inflicting energetic and reproductive costs on females. Because association with males is central to female chimpanzees’ anti-infanticide strategy, and males may confer other benefits, we propose that reproductive success in female chimpanzees hinges on a delicate balance between the costs and benefits of associating with male conspecifics.     

Effects of Selective Logging on Bat Communities in the Southeastern Amazon

Abstract:  Although extensive areas of tropical forest are selectively logged each year, the responses of bat communities to this form of disturbance have rarely been examined. Our objectives were to (1) compare bat abundance, species composition, and feeding guild structure between unlogged and low-intensity selectively logged (1–4 logged stems/ha) sampling grids in the southeastern Amazon and (2) examine correlations between logging-induced changes in bat communities and forest structure. We captured bats in understory and canopy mist nets set in five 1-ha study grids in both logged and unlogged forest. We captured 996 individuals, representing 5 families, 32 genera, and 49 species. Abundances of nectarivorous and frugivorous taxa (Glossophaginae, Lonchophyllinae, Stenodermatinae, and Carolliinae) were higher at logged sites, where canopy openness and understory foliage density were greatest. In contrast, insectivorous and omnivorous species (Emballonuridae, Mormoopidae, Phyllostominae, and Vespertilionidae) were more abundant in unlogged sites, where canopy foliage density and variability in the understory stratum were greatest. Multivariate analyses indicated that understory bat species composition differed strongly between logged and unlogged sites but provided little evidence of logging effects for the canopy fauna. Different responses among feeding guilds and taxonomic groups appeared to be related to foraging and echolocation strategies and to changes in canopy cover and understory foliage densities. Our results suggest that even low-intensity logging modifies habitat structure, leading to changes in bat species composition.     

The Sustainability of Subsistence Hunting by Matsigenka Native Communities in Manu National Park, Peru

Abstract:  The presence of indigenous people in tropical parks has fueled a debate over whether people in parks are conservation allies or direct threats to biodiversity. A well-known example is the Matsigenka (or Machiguenga) population residing in Manu National Park in Peruvian Amazonia. Because the exploitation of wild meat (or bushmeat), especially large vertebrates, represents the most significant internal threat to biodiversity in Manu, we analyzed 1 year of participatory monitoring of game offtake in two Matsigenka native communities within Manu Park (102,397 consumer days and 2,089 prey items). We used the Robinson and Redford (1991) index to identify five prey species hunted at or above maximum sustainable yield within the ∼150-km² core hunting zones of the two communities: woolly monkey (Lagothrix lagotricha), spider monkey (Ateles chamek), white-lipped peccary (Tayassu pecari), Razor-billed Currasow (Mitu tuberosa), and Spix's Guan (Penelope jacquacu). There was little or no evidence that any of these five species has become depleted, other than locally, despite a near doubling of the human population since 1988. Hunter–prey profiles have not changed since 1988, and there has been little change in per capita consumption rates or mean prey weights. The current offtake by the Matsigenka appears to be sustainable, apparently due to source–sink dynamics. Source–sink dynamics imply that even with continued human population growth within a settlement, offtake for each hunted species will eventually reach an asymptote. Thus, stabilizing the Matsigenka population around existing settlements should be a primary policy goal for Manu Park.     

Response of Savanna Fire Regimes to Changing Fire-Management Policies in a Large African National Park

Abstract:  Approaches to fire management in the savanna ecosystems of the 2-million ha Kruger National Park, South Africa, have changed several times over the past six decades. These approaches have included regular and flexible prescribed burning on fixed areas and a policy that sought to establish a lightning-dominated fire regime. We sought to establish whether changes in management induced the desired variability in fire regimes over a large area. We used a spatial database of information on all fires in the park between 1957 and 2002 to determine elements of the fire regime associated with each management policy. The area that burned in any given year was independent of the management approach and was strongly related to rainfall (and therefore grass fuels) in the preceding 2 years. On the other hand, management did affect the spatial heterogeneity of fires and their seasonal distribution. Heterogeneity was higher at all scales during the era of prescribed burning, compared with the lightning-fire interval. The lightning-fire interval also resulted in a greater proportion (72% vs. 38%) of the area burning in the dry season. Mean fire-return intervals varied between 5.6 and 7.3 years, and variability in fire-return intervals was strongly influenced by the sequencing of annual rainfall rather than by management. The attempt at creating a lightning-dominated fire regime failed because most fires were ignited by humans, and the policy has been replaced by a more pragmatic approach that combines flexible prescribed burning with lightning-ignited fires.     

Edge-Related Variation in Medicinal and Other "Useful" Wild Plants of Bwindi Impenetrable National Park, Uganda

Abstract:  Harvesting of wild plants for nontimber uses is widespread in the tropics, but its impact is usually quantified only for one or a few species at a time. Thus, forest managers are never clear about how well their efforts are protecting such plants. We quantified abundance and edge-related variation in 91 species of useful wild plants commonly harvested by communities around Bwindi Impenetrable National Park (BINP), Uganda, to evaluate the effect of their harvest. Forty percent of these species were harvested exclusively for medicines, 22% for weaving, and 24% for other uses. Fourteen percent were harvested for combinations of uses. Plants were surveyed around the entire periphery of the park transects that extended out 1 km into the forest interior from the edge. Analyses of edge and interior distribution were controlled for effects of topography. Individually, nine (10%) species were very rare, occurring in <0.5% of the plots searched. Of the remaining 82 species, most (50%) decreased significantly away from the park boundary, whereas 4.9% increased and 45.1% showed no pronounced edge-related distributions. Rarer species were no more likely to be less abundant near the edge than commoner species. These results suggest that most plants used for nontimber purposes in BINP are not currently being harvested unsustainably. In this respect many of the species of useful wild plants we examined resembled animals commonly hunted in tropical forests for bushmeat because they increased in abundance in disturbed habitat. Conservation action should initially aim to understand what influences distributions of very rare species. Edge-based assessments of distributions may be valuable for revealing harvest impact on species of useful wild plants commonly harvested by people living around forest islands in the tropics.     

Ranging behavior of the mountain gorillas (Gorilla beringei beringei) in Bwindi Impenetrable National Park, Uganda: a test of the ecological constraints model

The ecological constraints model predicts that daily travel distance and home range size of social animals will increase as group size increases in order to meet the dietary needs of additional group members. This theory has been supported more predominantly by studies of frugivorous primate species than by studies of folivorous species. We examined the ranging patterns of mountain gorillas in Bwindi Impenetrable National Park, Uganda, who include both herbaceous vegetation and fruit in their diet, to determine how ecological, behavioral, and social parameters influence movement patterns. Data were collected from three groups of gorillas with overlapping home ranges at a low-altitude location (1,450–1,800 m) and one group at a high-altitude location (2,100–2,500 m) in Bwindi from September 2001 to August 2002. We analyzed daily travel distance and home range size in relation to group size, while also considering patterns of frugivory, rainfall, and location (proxy for food availability) within the park. Both daily travel distance and home range size were positively related to group size. In addition, the degree of frugivory positively influenced daily travel distance and home range size, while rainfall negatively influenced daily travel distance only. Finally, groups at the low-altitude location, with higher fruit availability, traveled less than the group at the high-altitude location. These results demonstrate that mountain gorillas in Bwindi provide support for the ecological constraints model, but further studies are needed to determine how fine-scale spatial and temporal availability of food resources influence movement patterns. Ranging patterns of Bwindi gorillas are compared to those observed in other gorilla populations in the context of the ecological constraints model.Communicated by D. Watts     

Long-Term Effects of Forestry on Birds of the Eastern Canadian Boreal Forests: a Comparison with Fennoscandia

Abstract: Logging is considered the most important threat to species in boreal forests. In contrast to eastern Canada, where most boreal forests remain largely untouched, in Fennoscandia it is possible to assess the cumulative, long-term effects of intensive forestry on wildlife. But harvesting of stands is rapidly changing Canadian boreal forests, which represent an important proportion of the world's boreal forests. We show that Fennoscandia and eastern Canada present striking similarities in terms of forest-age structure, natural-disturbance regime, and structure of bird assemblages, and we provide an assessment of the long-term effects of forestry on eastern Canadian birds of the boreal forest. We used life-history traits from habitat, nesting site, and geographical range to calculate an index of sensitivity to changes induced by modern forestry for boreal species of each region. Tropical migrants commonly found in eastern Canadian boreal forests have life-history traits that are not threat factors in relation to changes caused by modern forestry. Therefore, the general belief that tropical migrants in North America are more sensitive to landscape changes than those in Europe may not hold for species found in the boreal coniferous forests of eastern Canada. Nine Fennoscandian species present high levels of sensitivity, and at least eight eastern Canadian species are of similar concern. In both regions, most of the sensitive species are resident cavity nesters. Given the important similarities between the two regions, the northern expansion of commercial forestry in eastern Canada is likely to result in the significant decline of several resident species, as has occurred in Fennoscandia.     

Effects of Controlled Fire and Livestock Grazing on Bird Communities in East African Savannas

Abstract: In East Africa fire and grazing by wild and domestic ungulates maintain savannas, and pastoralists historically set fires and herded livestock through the use of temporary corrals called bomas. In recent decades traditional pastoral practices have declined, and this may be affecting biodiversity. We investigated the effects of prescribed fires and bomas on savanna bird communities in East Africa during the first and second dry seasons of the year (respectively before and after the rains that mark the onset of breeding for most birds). We compared abundance, richness, and community composition on 9‐ha burned plots, recently abandoned bomas, and control plots in the undisturbed matrix habitat over a 3‐year period. Generally, recently burned areas and abandoned bomas attracted greater densities of birds and had different community assemblages than the surrounding matrix. The effects of disturbances were influenced by interactions between primary productivity, represented by the normalized difference vegetation index, and time. Bird densities were highest and a greater proportion of species was observed on burned plots in the months following the fires. Drought conditions equalized bird densities across treatments within 1 year, and individuals from a greater proportion of species were more commonly observed on abandoned bomas. Yearly fluctuations in abundance were less pronounced on bomas than on burns, which indicate that although fire may benefit birds in the short term, bomas may have a more‐lasting positive effect and provide resources during droughts. Several Palearctic migrants were attracted to burned plots regardless of rainfall, which indicates continued fire suppression may threaten their already‐declining populations. Most notably, the paucity of birds observed on the controls suggests that the current structure of the matrix developed as a result of fire suppression. Traditional pastoralism appears critical to the maintenance of avian diversity in these savannas.     

Effects of Historic Livestock Grazing on Vegetation at Chaco Culture National Historic Park, New Mexico

Abstract:   Livestock grazing is the most ubiquitous land use in western North America, yet it rarely has been studied in a controlled manner because of the lack of large areas free of grazing. We compared the ecological effects of three grazing treatments—long-term protection, short-term protection, and currently grazed—at Chaco Culture National Historic Park in northern New Mexico. Chaco has a long history of human habitation and is now one of the largest grazing exclosures in the American West. We studied the effects of livestock grazing on the cover of plants, soil crusts, and plant species richness at six sites with different potential natural vegetation. Species richness was higher under long-term protection than under current grazing at all six sites. Trends in shrub and grass response varied significantly across the six sites. Shrub cover increased with long-term protection at four upland sites, and grass cover increased with protection at four sites. The response of Chaco vegetation to release from grazing varied significantly according to each site's ecological potential, determined in part by edaphic and topographic characteristics. These nuances in vegetation response represent natural ecological variation and contrast with the notions of widespread shrub "invasion" often inferred in the past.     

Tourism policy,biodiversity conservation and management: a case of the Kruger National Park,South Africa

Management of Kruger National Park (KNP) has recently made considerable progress in developing new policies to address biodiversity conservation and management challenges. These include tourism, water provision and elephant management policies. This study examines the integration of these three policies with regard to biodiversity conservation and management. Findings indicate that in their current fragmented form, the three policies actually undermine opportunities for effective biodiversity conservation and management in the KNP. This can be reversed by adopting a new management approach based on an environmental matrix organizational framework. Such a framework provides a mechanism for integrated policymaking and implementation and for improving biodiversity conservation and management in the KNP. The study concludes that an integrated approach is necessary due to the interlinkages between policies that affect biodiversity conservation at the KNP.     

Effects of Selective Logging on the Butterflies of a Bornean Rainforest

Abstract: Selective logging has been the main cause of disturbance to tropical forests in Southeast Asia, so the extent to which biodiversity is maintained in selectively logged forest is of prime conservation importance. We compared the butterfly assemblages of Bornean primary rainforest to those of rainforest selectively logged 6 years previously. We sampled by means of replicated transects stratified into riverine and ridge forests and we included roads in the logged forest. There was a three-fold variation in species richness and abundance over the 8-month sampling period. More species and individuals were observed in the logged forest, although between-replicate variability was high. Rarefied species richness was positively correlated with canopy openness within the range of disturbance levels encountered at our forest sites. Within families, there was no significant difference in the number of species between primary and logged forest. There was a significant difference in the relative abundance of species, but this was due largely to the abundance of one or two species. Community ordination separated the sites along a gradient of disturbance and revealed strong differences between riverine and ridge-forest butterfly assemblages in primary forest that were obscured in logged forest. There was no evidence that logging has resulted in a change in the composition of the butterfly assemblages from species with a local distribution to more widespread species. We conclude that at a logged forest site in close proximity to primary forest, low intensities of logging do not necessarily reduce the species richness or abundance of butterflies, although assemblage composition is changed.     

Effects of Illegal Harvest of Eggs on the Population Decline of Leatherback Turtles in Las Baulas Marine National Park,Costa Rica

Abstract: Within 19 years the nesting population of leatherback turtles (Dermochelys coriacea) at Parque Nacional Marino Las Baulas declined from 1500 turtles nesting per year to about 100. We analyzed the effects of fishery bycatch and illegal harvesting (poaching) of eggs on this population. We modeled the population response to different levels of egg harvest (90, 75, 50, and 25%) and the effect of eradicating poaching at different times during the population decline. We compared effects of 90% poaching with those of 20% adult mortality because both of these processes were present in the population at Las Baulas. There was a stepwise decline in number of nesting turtles at all levels of egg harvest. Extirpation times for different levels of poaching ranged from 45 to 282 years. The nesting population declined more slowly and survived longer with 20% adult mortality (146 years) than it did with 90% poaching (45 years). Time that elapsed until poaching stopped determined the average population size at which the population stabilized, ranging from 90 to 420 nesting turtles. Our model predicted that saving clutches lost naturally would restore the population when adult mortality rates were low and would contribute more to population recovery when there were short remigration intervals between nesting seasons and a large proportion of natural loss of clutches. Because the model indicated that poaching was the most important cause of the leatherback decline at Las Baulas, protecting nests on the beach and protecting the beach from development are critical for survival of this population. Nevertheless, the model predicted that current high mortality rates of adults will prevent population recovery. Therefore, protection of the beach habitat and nests must be continued and fishery bycatch must be reduced to save this population.     

Variation in behavioral and hormonal responses of adult male gray-cheeked mangabeys (<Emphasis Type="Italic">Lophocebus albigena</Emphasis>) to crowned eagles (<Emphasis Type="Italic">Stephanoaetus coronatus</Emphasis>) in Kibale National Park,Uganda

Intensive study of arboreal forest-dwelling primates and their predators in Africa is increasingly revealing that crowned eagles (Stephanoaetus coronatus) are major predators of primates. Gray-cheeked mangabeys (Lophocebus albigena) are overrepresented in the diets of crowned eagles in Kibale National Park, Uganda, and adult male mangabeys are represented more than females. We focused on the behavior of adult male gray-cheeked mangabeys living in social groups in Kibale National Park (1) to clarify the interactions between mangabeys and eagles that might put adult males at greater risk and (2) to better understand individual variation in behavioral responses to predators. Adult male mangabeys in five groups responded to observer-confirmed presence of crowned eagles 88 times over a 13-month period. While all males gave alarm calls, only the highest-ranking male in each of four groups chased eagles. These males had elevated levels of fecal cortisol metabolites in the days immediately after they engaged in active defense, suggesting that they perceived such behavior as risky. In the one group where male ranks were unstable and there were no infants, no male was observed to chase eagles. We suggest that males pursue the dangerous tactic of chasing eagles only when they are likely to have offspring in the group. Males in larger groups also spent less time alarm calling to crowned eagles (from first to last call in a group), and our observations confirmed that the duration of their alarm calls was related to eagle presence. Thus, eagles spent less time around larger mangabey groups. Alarm calling by adult male mangabeys may signal to this ambush predator that it has been detected and should move on.