Changes of community composition at multiple trophic levels due to hunting in Nigerian tropical forests

Hunting in tropical forests decimates large mammals, and this may have direct and indirect effects on other trophic levels and lead to trophic cascades. We compared replicated sites of hunted and protected forests in southeastern Nigeria, with respect to community composition of primates, other mammals, birds, plant seedlings, and mature trees. We make predictions regarding the community composition at the different trophic levels. In forests where large primates are rare, we hypothesize that their ecological role will not be fully compensated for by small frugivores. We apply multivariate methods to assess changes in community composition of mammals, birds, and seedlings, controlling for any differences between sites in the other groups, including mature trees. Medium and large (4–180 kg) primates were much rarer in hunted sites, while porcupine and rock hyrax increased in abundance with hunting. In contrast, the community composition of birds was similar in both types of forests. Seedling communities were significantly related to the community composition of mammals, and thus strongly affected by hunting. In protected forests primate dispersed plant seedling species dominated, whereas in hunted forests the seedling community was shifted towards one dominated by abiotically dispersed species. This was probably both a consequence of reduced seed dispersal by primates, and increased seed predation by rodents and hyrax. Hence we found no evidence for buffering effects on tree regeneration through functional compensation by non‐hunted animals (such as birds). Our results highlight how seedling communities are changed by the complex plant–animal intera ctions, triggered by the loss of seed dispersers. The results predict a rarity of primate‐dispersed trees in future tropical forest canopies; a forest less diverse in timber and non‐timber resources.     

Mapping Bushmeat Hunting Pressure in Central Africa

Hunting and trade of wild animals for their meat (bushmeat), especially mammals, is commonplace in tropical forests worldwide. In West and Central Africa, bushmeat extraction has increased substantially during recent decades. Currently, such levels of hunting pose a major threat to native wildlife. In this paper, we compiled published data on hunting offtake of mammals, from a number of studies conducted between 1990 and 2007 in Cameroon, Central African Republic, Democratic Republic of Congo, Equatorial Guinea, Gabon, and Republic of Congo. From these data sources, we estimated annual extraction rates of all hunted species and analyzed the relationship between environmental and anthropogenic variables surrounding each hunting rate and levels of bushmeat extraction. We defined hunting pressure as a function of bushmeat offtake and number of hunted species and confirm that hunting pressure is significantly correlated with road density, distance to protected areas and population density. These correlations are then used to map hunting pressure across the Congo Basin. We show that predicted risk areas show a patchy distribution throughout the study region and that many protected areas are located in high‐risk areas. We suggest that such a map can be used to identify areas of greatest impact of hunting to guide large‐scale conservation planning initiatives for central Africa.     

A bioeconomic analysis of bushmeat hunting

Unsustainable bushmeat hunting is a major threat to mammal species, particularly in West/Central Africa. We developed a multispecies dynamic simulation model of hunter behaviour, parameterized using data from the Ashanti region, Ghana. The model distinguishes between two hunting techniques, snaring and gun hunting. We analyse the impact of key economic parameters on off-takes. Economic incentives determine the effort devoted to hunting, the choice of hunting technique, and the species that are consumed domestically or traded in markets. These factors, together with the growth rates and catchabilities of hunted species, determine the ecological impact of hunting. The results suggest that increased bushmeat prices are likely to lead to a switch from snaring, which is cheaper but less efficient, to gun hunting, with a consequent impact on vulnerable species. Increases in agricultural prices have an ambiguous effect on hunter behaviour, depending on the balance between incentives to invest in agriculture and increased consumption as incomes improve. Penalties are more effective if they target bushmeat sales, rather than the act of hunting. This model represents a step forward because it explicitly considers bushmeat as a component of the household economy. This has important implications as regards the development of policies to conserve species hunted for bushmeat.     

The Bushmeat Harvest Alters Seedling Banks by Favoring Lianas, Large Seeds, and Seeds Dispersed by Bats, Birds, and Wind

We evaluated predictions that hunters favor lianas, large seeds, and seeds dispersed by bats, small birds, and mechanical means for seedling banks in central Panama. We censused 3201 trees in 20 1-ha plots and 38,250 seedlings in the central 64 m² of each plot. We found significant differences in the species composition of the seedling bank between nine protected sites in the Barro Colorado Nature Monument and 11 hunted sites in the contiguous Parque Nacional Soberanía. Lianas, species with large seeds, and species with seeds dispersed by bats, small birds, and mechanical means were all overrepresented at hunted sites. The latter two findings could also be evaluated relative to the species composition of reproductively mature adults for canopy trees. The tree species present in the seedling bank had significantly heavier seeds than the tree species present as adults at hunted sites but not at protected sites. The representation of seed dispersal modes among the species present in the seedling bank did not reflect pre-existing differences in the local species composition of adults. We hypothesize that hunting large seed predators favors large seeds by reducing predation and increasing survival. We also hypothesize that the harvest of large birds and mammals that disperse many seeds favors other species whose seeds are dispersed by bats, small birds, and mechanical means. This process also favors lianas because the seeds of disproportionate numbers of liana species are dispersed by wind.     

African rainforests: past,present and future

The rainforests are the great green heart of Africa, and present a unique combination of ecological, climatic and human interactions. In this synthesis paper, we review the past and present state processes of change in African rainforests, and explore the challenges and opportunities for maintaining a viable future for these biomes. We draw in particular on the insights and new analyses emerging from the Theme Issue on ‘African rainforests: past, present and future’ of Philosophical Transactions of the Royal Society B. A combination of features characterize the African rainforest biome, including a history of climate variation; forest expansion and retreat; a long history of human interaction with the biome; a relatively low plant species diversity but large tree biomass; a historically exceptionally high animal biomass that is now being severely hunted down; the dominance of selective logging; small-scale farming and bushmeat hunting as the major forms of direct human pressure; and, in Central Africa, the particular context of mineral- and oil-driven economies that have resulted in unusually low rates of deforestation and agricultural activity. We conclude by discussing how this combination of factors influences the prospects for African forests in the twenty-first century.     

Hunting of Mammals Reduces Seed Removal and Dispersal of the Afrotropical Tree Antrocaryon klaineanum (Anacardiaceae)

Throughout the tropics, mammalian seed dispersers are being driven to local extinction by intense hunting pressure, generating concern not only about the loss of these species, but also about the consequences for the plants they disperse. We compared two rain forest sites in Cameroon—one with heavy hunting pressure and one protected from hunting—to appraise the loss of mammalian seed dispersers and to assess the impact of this loss on seed removal and seed dispersal of Antrocaryon klaineanum (Anacardiaceae), a mammal-dispersed tree. Surveys of arboreal frugivores indicate that three of the five monkey species, as well as chimpanzee and gorilla, have been extirpated from the hunted forest. Diaspore counts underneath A. klaineanum adults (six trees per site) indicate that seed removal is severely reduced in the hunted forest. Finally, genetic maternity exclusion analysis (using 3–7 nuclear microsatellite loci) of maternally inherited endocarp tissue from diaspores collected under the canopies of 12 fruiting "mother" trees (six trees per site) revealed that seed dispersal in the hunted forest is also greatly reduced. In the hunted forest with reduced mammal dispersal agents, only 1 of the 53 assayed endocarps (2%) did not match the mother and was determined to be from a dispersed diaspore. By contrast, in the protected forest, 20 of the 48 assayed endocarps (42%) were from dispersed diaspores. This study provides strong evidence that loss of dispersal agents can lead to reduced seed removal and loss of seed dispersal, disrupting the seed dispersal cycle.     

Rural Bushmeat Consumption Within Multiple-use Protected Areas: Qualitative Evidence from Southwest Madagascar

Ensuring the sustainability of bushmeat consumption is critical for both biodiversity conservation and poverty alleviation in tropical developing countries, yet we know little about the role of hunting and bushmeat consumption in the daily lives of rural communities. We provide the first detailed, qualitative examination of bushmeat hunting activities conducted by a rural community within one of Madagascar’s new, multiple-use protected areas, in order to inform appropriate management strategies. Results suggest that most species are eaten, but that few are favoured above domestic meat. Hunting is generally a secondary pursuit, carried out opportunistically during the course of other activities, although its importance does increase in times of food stress. Management focused on increasing domestic meat availability and directing hunting effort away from sensitive species may improve the sustainability of hunting, but development interventions to reduce forest dependence may be required to promote conservation and poverty alleviation simultaneously.     

Effects of hunting on guianan forest game birds

To assess the effect of on-going and of previous or near by hunting pressure on game birds in a neotropical rain forest (French Guiana), I compared species abundances between six hunted and disturbed areas, seven non-hunted and pristine areas and eight intermediate areas, undisturbed and not currently hunted, but formerly hunted or close to current hunting areas. I recorded all birds detected within 100 m-wide strip transects, walked at random through every forest type all day long. The frequency of records per 10 h and flock sizes on each transect were averaged over all surveys in each study site (N = 3025 h). Censused groups included terrestrial (tinamous, woodquails, curassow and trumpeter) and arboreal species (guans, macaws, parrots, toucans), but not waterbirds. The abundance of all game birds was higher in non-hunted than in hunted areas, though not always significantly for secretive understorey species (tinamous, guans, wood-quail) or canopy frugivores (parrots, macaws, toucans). The Black Curassow and Gray-winged Trumpeter exhibited the highest and most consistent increases (7–10-fold) and more so between intermediate and non-hunted sites than between hunted and intermediate sites. The mean flock size also increased along the hunting gradient, especially in flocking species (macaws, parrots, trumpeter). Low reproductive rates and/or seasonal movements may explain that hunting pressure could still be felt after hunting has ceased or when it persists only away from a given area. Such a persistent effect would affect some populations in small protected areas.     

Does illegal hunting affect density and behaviour of African grassland birds? A case study on ostrich (Struthio camelus)

Widespread bushmeat hunting represents one of the major threats to many mammals and birds in Africa. We studied the influence of illegal bushmeat hunting on large grassland birds in the Serengeti National Park (SNP) and adjoining protected areas, by using the ostrich (Struthio camelus) as a case study. First, we documented illegal hunting of both small and large birds by using a questionnaire in the villages on the western and eastern side of the SNP. Second, we studied the effect of illegal hunting on density by driving 4,659 km of transects inside SNP and on the adjacent protected areas, where the data were analysed by DISTANCE sampling. Last, we used flight initiation distance (FID, i.e. the distance between an approaching predator (human) and prey when flight is started), to assess possible impacts on behaviour from illegal hunting. We found that people from the western side of the SNP admitted to hunting both small and large grassland birds, and collect ostrich feathers and eggs. Although the Maasai also hunted small birds, only ostrich feathers and eggs of the large grassland birds were used. Surprisingly, we found no significant differences in densities between the SNP and adjoining partially protected areas, but ostriches had longer FID to an approaching human outside the SNP. Currently illegal hunting does not appear to affect the ostrich population, but given the extensive use of birds for consumption more awareness educational programs accompanied by provision of agricultural incentives within the protected areas are needed.     

Hunted Woolly Monkeys (Lagothrix poeppigii) Show Threat-Sensitive Responses to Human Presence

Responding only to individuals of a predator species which display threatening behaviour allows prey species to minimise energy expenditure and other costs of predator avoidance, such as disruption of feeding. The threat sensitivity hypothesis predicts such behaviour in prey species. If hunted animals are unable to distinguish dangerous humans from non-dangerous humans, human hunting is likely to have a greater effect on prey populations as all human encounters should lead to predator avoidance, increasing stress and creating opportunity costs for exploited populations. We test the threat sensitivity hypothesis in wild Poeppigi''s woolly monkeys (Lagothrix poeppigii) in Yasuní National Park, Ecuador, by presenting human models engaging in one of three behaviours “hunting”, “gathering” or “researching”. These experiments were conducted at two sites with differing hunting pressures. Visibility, movement and vocalisations were recorded and results from two sites showed that groups changed their behaviours after being exposed to humans, and did so in different ways depending on the behaviour of the human model. Results at the site with higher hunting pressure were consistent with predictions based on the threat sensitivity hypothesis. Although results at the site with lower hunting pressure were not consistent with the results at the site with higher hunting pressure, groups at this site also showed differential responses to different human behaviours. These results provide evidence of threat-sensitive predator avoidance in hunted primates, which may allow them to conserve both time and energy when encountering humans which pose no threat.     

Analysis of patterns of bushmeat consumption reveals extensive exploitation of protected species in eastern Madagascar

Understanding the patterns of wild meat consumption from tropical forests is important for designing approaches to address this major threat to biodiversity and mitigate potential pathways for transmission of emerging diseases. Bushmeat consumption has been particularly poorly studied in Madagascar, one of the world's hottest biodiversity hotspots. Studying bushmeat consumption is challenging as many species are protected and researchers must consider the incentives faced by informants. Using interviews with 1154 households in 12 communes in eastern Madagascar, as well as local monitoring data, we investigated the importance of socio-economic variables, taste preference and traditional taboos on consumption of 50 wild and domestic species. The majority of meals contain no animal protein. However, respondents consume a wide range of wild species and 95% of respondents have eaten at least one protected species (and nearly 45% have eaten more than 10). The rural/urban divide and wealth are important predictors of bushmeat consumption, but the magnitude and direction of the effect varies between species. Bushmeat species are not preferred and are considered inferior to fish and domestic animals. Taboos have provided protection to some species, particularly the Endangered Indri, but we present evidence that this taboo is rapidly eroding. By considering a variety of potential influences on consumption in a single study we have improved understanding of who is eating bushmeat and why. Evidence that bushmeat species are not generally preferred meats suggest that projects which increase the availability of domestic meat and fish may have success at reducing demand. We also suggest that enforcement of existing wildlife and firearm laws should be a priority, particularly in areas undergoing rapid social change. The issue of hunting as an important threat to biodiversity in Madagascar is only now being fully recognised. Urgent action is required to ensure that heavily hunted species are adequately protected.     

Zoonotic Disease Risk and the Bushmeat Trade: Assessing Awareness Among Hunters and Traders in Sierra Leone

The bushmeat industry has been a topic of increasing importance among both conservationists and public health officials for its influence on zoonotic disease transmission and animal conservation. While the association between infectious diseases and the bushmeat trade is well established in the research community, risk perception among bushmeat hunters and traders has not been well characterized. I conducted surveys of 123 bushmeat hunters and traders in rural Sierra Leone to investigate hunting practices and awareness of zoonotic disease risk associated with the bushmeat trade. Twenty-four percent of bushmeat hunters and traders reported knowledge of disease transmission from animals to humans. Formal education did not significantly affect awareness of zoonotic disease transmission. Individuals who engaged exclusively in preparation and trading of bushmeat were more likely to accidentally cut themselves compared to those who primarily engaged in bushmeat hunting (P < 0.001). In addition, women involved in the bushmeat trade were at greater risk of exposing themselves to potential zoonotic pathogens through accidental self-cutting compared to men (P < 0.01). This study collected preliminary information on risk perception among bushmeat hunters that could guide the creation of a future public health-based education program to minimize zoonotic disease transmission risk among vulnerable communities.     

Patterns and determinants of monkey densities in Peru and Bolivia,with notes on distributions

A comparative study of species assemblages and population densities was conducted on Amazonian monkey communities in 16 areas, ranging from 3°S latitude in northern Peru to 18°S latitude in southern Bolivia. The habitats ranged from several types of tropical rain forest in the more northern latitudes to dry, deciduous forest in the southernmost study area. The monkey populations of three of the study areas have historically received light hunting pressure; the rest have been moderately to heavily hunted. A transect census technique was used to estimate the relative and absolute densities of all diurnal monkey species except Cebuella pygmaea. The number of coexisting monkey species ranged from 4–6 in the southern areas to 12–14 in the northern areas. The reduction in species richness in central and southern areas of Bolivia is probably attributable to several inimical habitat factors. Predation by humans was found to be the single most important factor affecting monkey densities. Monkey densities, and especially biomasses, were much lower in areas not protected from hunting than in protected areas. Hunting did not affect all species equally. Larger-sized species are hunted more and have severely reduced numbers in unprotected areas, whereas the densities of smaller species are not noticeably diminished in unprotected areas. Large, herbivorous monkey species contributed the major proportion of the total monkey biomass in protected areas. The strong influence of hunting has largely obscured the effects of other factors on population densities.     

Bushmeat and the Fate of Trees with Seeds Dispersed by Large Primates in a Lowland Rain Forest in Western Amazonia

In Neotropical forests, large fruit-eating primates play important ecological roles as dispersal agents of large seeds. Bushmeat hunting threatens to disrupt populations of primates and large-seeded trees. We test the hypothesis that otherwise intact Neotropical forests with depressed populations of large primates experience decline in recruitment of large-seeded trees. We quantify the proportion of small juveniles (> 0.5 m tall–1 cm diameter at breast height, DBH) of large primate-dispersed tree species found underneath heterospecifc trees that are also dispersed by large primates at two protected sites in Manu National Park and one hunted site outside Manu N.P. in southeastern Peru. The forests are comparable in edaphic and climatic qualities, successional stage, and adult tree species composition. We found that hunting locally exterminates populations of large primates, and reduced primates of intermediate body size (hereafter "medium primates") by 80 percent. Moreover, tree species richness was 55 percent lower and density of species dispersed by large and medium-bodied primates 60 percent lower in hunted than in protected sites. In addition, richness and density of abiotically dispersed species and plants dispersed by non-game animals are greater in hunted sites. Overhunting threatens to disrupt the ecological interactions between primates and the plants that rely on them for seed dispersal and recruitment. Sustainable wildlife management plans are urgently needed, because protected areas are at risk of becoming "island" parks if buffer zones become empty of animals and have impoverished flora.     

The importance of well protected forests for the conservation genetics of West African colobine monkeys

In tropical forests, anthropogenic activities are major drivers of the destruction and degradation of natural habitats, causing severe biodiversity loss. African colobine monkeys (Colobinae) are mainly folivore and strictly arboreal primates that require large forests to subsist, being among the most vulnerable of all nonhuman primates. The Western red colobus Piliocolobus badius and the King colobus Colobus polykomos inhabit highly fragmented West African forests, including the Cantanhez Forests National Park (CFNP) in Guinea-Bissau. Both species are also found in the largest and best-preserved West African forest—the Taï National Park (TNP) in Ivory Coast. Colobine monkeys are hunted for bushmeat in both protected areas, but these exhibit contrasting levels of forest fragmentation, thus offering an excellent opportunity to investigate the importance of well-preserved forests for the maintenance of evolutionary potential in these arboreal primates. We estimated genetic diversity, population structure, and demographic history by using microsatellite loci and mitochondrial DNA. We then compared the genetic patterns of the colobines from TNP with the ones previously obtained for CFNP and found contrasting genetic patterns. Contrary to the colobines from CFNP that showed very low genetic diversity and a strong population decline, the populations in TNP still maintain high levels of genetic diversity and we found no clear signal of population decrease in Western red colobus and a limited decrease in King colobus. These results suggest larger and historically more stable populations in TNP compared to CFNP. We cannot exclude the possibility that the demographic effects resulting from the recent increase of bushmeat hunting are not yet detectable in TNP using genetic data. Nevertheless, the fact that the TNP colobus populations are highly genetically diverse and maintain large effective population sizes suggests that well-preserved forests are crucial for the maintenance of populations, species, and probably for the evolutionary potential in colobines.     

Identifying Conservation Opportunities among Malinké Bushmeat Hunters of Guinea,West Africa

Bushmeat hunting plays an important role in many rural African households; however, hunting pressures are growing, threatening the survival of many bushmeat species. Wildlife resources are rapidly dwindling; yet effective conservation strategies have been difficult to develop and implement. Many dimensions of bushmeat resource use have not been sufficiently explored and are consequently ignored in conservation interventions. To improve understanding of hunting practice, we conducted semistructured interviews with 74 hunters in three Malinké villages in Guinea, West Africa regarding motivations to hunt and hunting processes. In addition, we investigated the local bushmeat trade in a nearby city. Using data from previous studies, and Robinson and Redford’s (1991) model, we find that Red-flanked duiker, bushbuck and buffalo are unsustainably harvested. Malinké hunters’ perspectives offer both opportunities and obstacles for conservation which will be valuable for the development of conservation strategies in this area.     

Mapping suitable great ape habitat in and around the Lobéké National Park,South‐East Cameroon

As a result of extensive data collection efforts over the last 20–30 years, there is quite a good understanding of the large‐scale geographic distribution and range limits of African great apes. However, as human activities increasingly fragment great ape spatial distribution, a better understanding of what constitutes suitable great ape habitat is needed to inform conservation and resource extraction management. Chimpanzees (Pan troglodytes troglodytes) and gorillas (Gorilla gorilla gorilla) inhabit the Lobéké National Park and its surrounding forest management units (FMUs) in South‐East Cameroon. Both park and neighboring forestry concessions require reliable evidence on key factors driving great ape distribution for their management plans, yet this information is largely missing and incomplete. This study aimed at mapping great ape habitat suitability in the area and at identifying the most influential predictors among three predictor categories, including landscape predictors (dense forest, swampy forest, distance to water bodies, and topography), human disturbance predictors (hunting, deforestation, distance to roads, and population density), and bioclimatic predictor (annual precipitation). We found that about 63% of highly to moderately suitable chimpanzee habitat occurred within the Lobéké National Park, while only 8.4% of similar habitat conditions occurred within FMUs. For gorillas, highly and moderately suitable habitats occurred within the Lobéké National Park and its surrounding FMUs (82.6% and 65.5%, respectively). Key determinants of suitable chimpanzee habitat were hunting pressure and dense forest, with species occurrence probability optimal at relatively lower hunting rates and at relatively high‐dense forest areas. Key determinants of suitable gorilla habitat were hunting pressure, dense forests, swampy forests, and slope, with species occurrence probability optimal at relatively high‐dense and swampy forest areas and at areas with mild slopes. Our findings show differential response of the two ape species to forestry activities in the study area, thus aligning with previous studies.     

Possible Indirect Effects of Mammal Hunting on Dung Beetle Assemblages in Panama

Hunting often impacts rain forest mammal communities but little is known about its indirect effects on other taxa. We examined dung beetle assemblages using pitfall and flight-intercept traps at six rain forest sites in Panama that ranged in hunting intensity. Heavily hunted sites showed altered community composition, significantly fewer species (based on rarefaction), and lower abundances of dung beetles than did sites with little hunting. Our results suggest that intensive hunting affects nontarget forest taxa and is potentially altering ecosystem functioning.     

Assessing the impact of hunting pressure on population structure of Guinea baboons (Papio papio) in Guinea-Bissau

Guinea baboons are heavily hunted for bushmeat consumption in Guinea-Bissau. We investigated whether hunting-driven mortality has affected population structure in this generalist primate using two genetic markers. Sampling was conducted in protected areas separated by anthropogenic landscape features. We predicted significant genetic differentiation between samples and investigated whether genetic discontinuities in the data were concordant with the location of human infrastructures. Genetic diversity was not significantly reduced when compared with a neighbouring population in Senegal and we inferred historically female-biased dispersal and recent contact between localities. Evidence was found for a contact zone between genetically differentiated populations where gene-flow is unidirectional, admixed individuals are at a higher proportion and individuals differentiated for both genetic markers co-exist within the same social units. Genetic discontinuities were, however, unrelated to anthropogenic dispersal barriers and we could not explain the existence of a contact zone by geographic distance, habitat type or the effect of social structure. We propose that hunting practices have affected the population structure by increasing dispersal distances, facilitating contact between previously separated gene pools within social groups. We suggest that hunting-related density sinks found in areas where the quality of the habitat remains adequate could precipitate the immigration of genetically distinct individuals from distant populations. Alternatively, migrants found in protected areas might be avoiding hunters, in locations they may perceive as less disturbed. This study suggests that hunting practices must be considered when investigating genetic patterns in primates and underlines the utility of molecular approaches to detect population perturbations due to bushmeat hunting.