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Hunting is one of the greatest threats to nonhuman primates worldwide. Despite Madagascar’s status as a primate mega-diversity country, a critical lack of information on the hunting of lemurs at the national scale persists. Here, we synthesize the current state of knowledge of the annual rates of household-level lemur hunting near ten protected areas, representing most ecoregions in Madagascar. We examine geographic and taxonomic variation in lemur hunting, including an analysis of hunting relative to species density, extinction risk, and intrinsic ecological characteristics of species. We found that lemurs are commonly hunted across Madagascar; the rural households in our study ate, on average, more than one lemur each year, or a median of 4.1% of the lemur species’ population size where densities are known. However, this pressure varied significantly across sites and species, reaching its highest levels in the northeastern rainforest region. While hunting levels are concerning for numerous threatened species, hunting pressure was driven primarily by species availability, and among ecological traits, small body size was related to increased hunting; however, conservation status showed no such relationship. This first national-level assessment of hunting, including one-third of Madagascar’s lemur species and more than a tenth of the world’s primates, identifies regional variation and lemur taxa at acute risk from hunting—important steps toward developing targeted strategies to conserve one of the world’s most threatened groups of vertebrates.

Introduced predators currently threaten endemic animals on Madagascar through predation, facilitation of human-led hunts, competition, and disease transmission, but the antiquity and past consequences of these introductions are poorly known. We use directly radiocarbon dated bones of introduced dogs (Canis familiaris) to test whether dogs could have aided human-led hunts of the island’s extinct megafauna. We compare carbon and nitrogen isotope data from the bone collagen of dogs and endemic “fosa” (Cryptoprocta spp.) in central and southwestern Madagascar to test for competition between introduced and endemic predators. The distinct isotopic niches of dogs and fosa suggest that any past antagonistic relationship between these predators did not follow from predation or competition for shared prey. Radiocarbon dates confirm that dogs have been present on Madagascar for over a millennium and suggest that they at least briefly co-occurred with the island’s extinct megafauna, which included giant lemurs, elephant birds, and pygmy hippopotamuses. Today, dogs share a mutualism with pastoralists who also occasionally hunt endemic vertebrates, and similar behavior is reflected in deposits at several Malagasy paleontological sites that contain dog and livestock bones along with butchered bones of extinct megafauna and extant lemurs. Dogs on Madagascar have had a wide range of diets during the past millennium, but relatively high stable carbon isotope values suggest few individuals relied primarily on forest bushmeat. Our newly generated data suggest that dogs were part of a suite of animal introductions beginning over a millennium ago that coincided with widespread landscape transformation and megafaunal extinction. Introduction Materials and Methods - Specimen and Site Selection - 14C and Stable Isotope Analyses - Data Analysis Results - Chronological Data - Stable Isotope Data Discussion

Bushmeat consumption in Madagascar is increasingly acknowledged as one of the major threats to its wild vertebrates. Nevertheless, few studies have examined the drivers of the consumption of protected versus legally huntable wild species, or examined its variance across Madagascar’s protected and unprotected areas. This research provides a novel study of the consumption of protected, unprotected, and fish/eel species between forest types (deciduous and rainforest), as well as across a gradient of protected habitat (National Park, Reserve, Unprotected). Members of 1750 households were interviewed across four regions, including two national parks, two reserves, and two unprotected forests. Household demographic, socioeconomic, cultural, and geographic variables were explored as possible predictors of bushmeat consumption. We found that poorer households reported consuming greater quantities of protected species whereas wealthier households reported consuming greater quantities of fish and eel. Households located inside Andasibe-Mantadia National Park, Madagascar’s most visited protected area, reported consuming the greatest quantities of protected species. Interviewees’ most favoured meat was from livestock, and fish. The consumption pattern of wild species reflected interviewees’ stated preference for species that are either unlisted (e.g. tilapia fish) under Malagasy species protection laws, classified as pest (e.g. bushpig) and/or game species (e.g. tenrec). Most protected species (such as lemurs and carnivorans) were interviewees’ least favoured wild meats. Given the lack of cultural affinity, and low preference for the consumption of most protected species, our results suggest that improving accessibility to domestic meat is a promising strategy for reducing the consumption of protected species.

As with many other species in the primate order, ring-tailed lemurs (Lemur catta) are threatened with extinction. Our articles documented declines in wild ring-tailed lemur populations and noted that fewer than 2,500 wild ring-tailed lemurs are known to persist in 32 [Gould and Sauther: Primate Conservation 2016; 30: 89–101] and 34 [LaFleur et al.: Folia Primatologica 2017; 87: 320–330] sites. A criticism of our articles [Murphy et al.: International Journal of Primatology 2017; 38: 623–628] suggested that we have inadequately sampled ring-tailed lemur populations and habitats, and misused the literature. We disagree, and provide both a detailed rebuttal and responses to specific critique points herein. Moreover, we restate our case outlining a dramatic decline of ring-tailed lemurs resulting from anthropogenic pressures (deforestation, severe habitat fragmentation, extraction for the pet and bushmeat trades). We pose several thought-provoking questions as to when is the appropriate time for researchers to “sound the alarm” about a species’ decline, and remain committed to understanding the drivers of unsustainable exploitation of this emblematic lemur, and preventing their extinction in the wild.

The main threats to lemurs are habitat loss and hunting. Conservation policies often assume that people will decrease lemur hunting if they understand government prohibitions on hunting, are educated and/or involved in ecotourism, have access to affordable meat, and/or are healthy and financially secure. Yet these assumptions are often not well tested where conservation policies are implemented. We interviewed every member of a focal village in one of the most biodiverse places on earth, the Masoala peninsula of Madagascar. The factors that best predicted the decision to hunt lemurs were poverty, poor health, and child malnutrition. Knowledge of laws, level of education, involvement in ecotourism, traditional cultural values, taste preferences, opportunity, and human–wildlife conflict had no impact on lemur hunting. Our results suggest that the welfare of humans and lemurs are linked. The key to discouraging illegal hunting and improving the viability of Endangered lemur populations may be improving rural human health and welfare.

SUMMARYThe role of wild meat for subsistence or as a luxury good is debated. We investigated the role of wild meat in food security in Madagascar, where consumption is poorly understood in urban areas and at regional scales. Using semi-structured interviews (n = 1339 heads-of-households, 21 towns), we aimed to: (1) quantify the amount and purpose of, (2) understand the drivers of, and (3) examine changes in wild meat consumption. Few respondents preferred wild meat (8 ± 3%) but most had eaten it at least once in their lifetime (78 ± 7%). Consumption occurred across ethnic groups, in urban and rural settings. More food insecure areas reported higher rates of wild meat consumption in the 6–8 months prior to interviews. Consumption was best explained by individual preferences and taboos. Less than 1% of respondents had increased consumption during their lifetimes. Wild meat prices showed no change from 2005–2013. Most consumption involved wild pigs and smaller-sized animals, though they were consumed less in the years following the 2009 coup. These data illustrate the differences between urban and rural communities, the occasions in which wild meat is used a source of food security, and provide evidence that some taxa are not hunted sustainably in Madagascar.

Lemur catta is the most reported illegal captive lemur. We document 286 L. catta that were held in illegal captive conditions in Madagascar. Coastal tourist destinations are “hot spots” for sightings. Many of the L. catta reported were in businesses (49%) and were perceived to be held captive for the purpose of generating income (41%). Infant/juvenile L. catta were overwhelmingly observed annually in December (41%) and may suffer high mortality rates given that they are not weaned during this month of the year. Population growth modeling suggests that known capture rates may be sustainable in all but small populations of 500 individuals and when infants/juveniles are targeted. However, of the seven remaining populations of L. catta with more than 100 individuals, only one is known to contain more than 500 animals, and we present evidence here that infants/juveniles are targeted. Moreover L. catta face significant other threats including habitat loss, bushmeat hunting, and climate change. Several actions could reduce the illegal capture and ownership of L. catta in Madagascar such as tourist behavior change initiatives, enforcement of laws, and alternative livelihoods for local people. These interventions are urgently needed and could be adapted to protect other exploited wildlife in the future.

Studying seasonal hunting patterns can be critical for developing sound actions for conservation and public health. As availability of funds to implement conservation policy is limited, it is essential to focus efforts during the most critical times of year. During July 2011–June 2012 I recorded direct observations of hunting of forest mammals, and conducted daily 24-hour recall surveys (2 weeks per month over 11 months: August 2011–June 2012), and interviews of all households in a focal village on the Masoala Peninsula of Madagascar to investigate (1) what drives seasonal hunting patterns and (2) how seasonal variation in consumption of wildlife and domestic meat affects native species and people. There is marked seasonal variation in hunting of forest mammals and in the consumption of fish and domesticated livestock on the Masoala Peninsula. Hunters target bushpigs Potamochoerus larvatus and tenrec and lemur species during the austral winter (March–August), whereas more native and introduced carnivorans, fish and domesticated livestock are consumed during the austral summer (September–February). The results suggest that seasonal variation in hunting patterns is driven by the physical and behavioural characteristics of prey rather than seasonal scarcity of alternative meat. Seasonal hunting and meat consumption on the Masoala Peninsula may amplify the negative impact of hunting on native carnivorans and tenrecs (which are hunted when they are pregnant and lactating), and the positive impact of consumption of lemurs, bushpigs and tenrecs on human health. This study highlights an important aspect of hunting on the Masoala: the decision whether or not to hunt is made independently of decisions regarding when to hunt particular species.

Threats to primates result from the complex relationship between ecological processes and the direct and the indirect impacts of humans. Yet we know little about the proportional impacts of hunting and changes to habitat on individual primate species. This knowledge is critical to effective conservation. I used primate surveys, habitat analysis, interviews, and one year of direct observation of hunter behavior and catch to compare the relative impacts of altered habitat and snare trapping on two sympatric lemur species: the two largest-bodied and most endangered lemurs on the Masoala peninsula of Madagascar, Varecia rubra (the red ruffed lemur; Critically Endangered) and Eulemur albifrons (the white-fronted brown lemur; Endangered). Results indicate that alteration of habitat and hunting shape local faunal communities in species-specific ways. While alteration of habitat had a greater effect than snare trapping on the populations of V. rubra, snare trapping had a greater effect than habitat on the populations of E. albifrons. Therefore conservation action plans for V. rubra and E. albifrons may benefit from individual tailoring. These findings illustrate the need to consider the different manners in which habitat change and hunting affect sympatric primate species when designing conservation policy.

Environmental education is widely used to increase awareness of conservation issues. The theory is that increasing knowledge will improve attitudes towards the environment. Often, environmental education is aimed at children with the assumption that this can also impact adults through intergenerational transfer of knowledge and attitudes. However, there are few detailed studies evaluating the effectiveness of environmental education on changing knowledge and attitudes, and whether any changes do transfer between generations. We evaluate the effect of a school-based education programme run by Malagasy researchers aimed at promoting lemur conservation in Eastern Madagascar. We assess changes in the knowledge and attitudes of participating children and their parents (surveying 126 children and 88 parents across four matched villages, 1 year after two of the villages received environmental education). There was very low awareness of the law protecting lemurs. Attitudes towards lemurs varied between species; with the aye-aye (considered scary) and the eastern lesser bamboo lemur (considered a pest) being less preferred. Children in villages who received environmental education had higher knowledge about lemurs and more positive attitudes than children in the villages not exposed to the environmental education. Knowledge about lemurs among parents where children had received environmental education was also higher (although not attitudes). Environmental education programmes can have a lasting effect, certainly on knowledge, but engagement of the research and NGO community is needed to build the capacity of teachers in rural areas to enthuse their pupils about ecology and conservation issues.

Information on the extent of bushmeat hunting is needed to assess the likely impact on hunted species, to provide information on the opportunity cost to local people of conservation, and to judge the efficacy of interventions at reducing pressure. However, where hunting is illegal, or socially unacceptable, respondents may not answer honestly to direct questions about hunting or consumption of bushmeat. We adapted a specialized method for investigating sensitive behaviours (the randomized response technique, RRT) and questioned 1,851 people in Madagascar about their consumption of six species, using either RRT or direct questions. For most species at most sites RRT and direct questions returned similar estimates of the proportion of the population who had consumed bushmeat in the previous year. However, RRT resulted in significantly higher estimates of bushmeat consumption in communities surrounding a protected area, where conservation activities made such questions sensitive. RRT has been predominately used in Europe and the USA; we demonstrate that it can provide a valuable approach for studying rule-breaking among people with poor literacy in low income countries. Between 12 and 33% of people across our sites had eaten brown lemur (Eulemur spp.), and 12–29% had eaten sifaka (Propithecus spp.) in the previous year. These results add to the growing body of evidence that hunting of protected species in Madagascar is a serious problem requiring urgent action. Conservation interventions to tackle bushmeat hunting will make questions about hunting or consumption more sensitive, increasing the need for researchers to use appropriate approaches for asking sensitive questions.

Lemurs are the most threatened group of mammals on earth. Lemur catta (ring-tailed lemur) represents one of the most iconic lemur species and faces numerous anthropogenic threats in the wild. In this study, we present population estimates from 32 sites across the range of L. catta, collected from primary and secondary data sources, to assess the number of ring-tailed lemurs left in the wild. We estimate that there are approximately 2,220 individual L. catta remaining in the 32 sites considered. We note local extinctions of populations of L. catta in at least 12 of the 32 sites examined, and that significantly more extinctions occurred in areas without some form of protection. This decrease in extant populations could represent a decrease of more than 95% of all ring-tailed lemurs in Madagascar since the year 2000. While these results should be considered preliminary, we stress the rapid decline of the species and note that habitat loss, bushmeat hunting and the illegal pet trade are driving populations to local extinction. Based on the data presented here, urgent and immediate funding and conservation action are crucial to ensure the viability of the remaining wild populations of ring-tailed lemurs.

We assessed the consumption and hunting of wild animals by people in urban areas of western Madagascar using structured questionnaires in households and direct observations. Six wild mammal and five wild bird species were reported, or observed, to be sources of bushmeat although fish and domestic animals were the preferred and cheapest sources of animal protein. Bushmeat accounted for 10% of the meat consumed the day before our questionnaires were completed. Common tenrec Tenrec ecaudatus and bush pig Potamochoerus larvatus were the preferred wild meat and the former was also the most expensive type of meat. Taboos and strong dislikes limited the consumption of domestic pigs, bush pigs, goats, lemurs and fruit bats. Game species were hunted according to their availability, which coincided with the legal hunting season for fruit bats but only partly so for the other game species. Illegal hunting of Verreauxi’s sifaka Propithecus verreauxi is cause for concern and assessments of primate consumption may have been underestimated because of reluctance of interviewees to admit illegal activities.

In the 2003 Durban Vision the Malagasy government committed to tripling the amount of protected areas in Madagascar by 2009. This extensive expansion needs to involve an assessment of the potential impacts on the people who rely on forest resources for subsistence. Wildlife for human consumption (bushmeat) is one such resource that has received great attention on mainland Africa but has largely been ignored in Madagascar until recently. In terms of biomass, hunting in Madagascar appears to be on a lesser scale compared to areas of mainland Africa. However, because of the life-history characteristics associated with hunted primate and carnivore species in Madagascar even small-scale hunting is a major threat to long-term conservation. In this study I used semi-structured interviews to quantify annual rates of bushmeat harvest in 14 villages adjacent to the Makira Forest in north-eastern Madagascar. Interviews revealed that 23 mammal species were hunted for consumption, providing a new insight into the scale and frequency of bushmeat use. Harvest data and life-history information were sufficient to allow quantitative assessments of sustainability for four species of lemur (black and white ruffed lemur Varecia variegata, indri Indri indri, eastern bamboo lemur Hapalemur griseus and white-fronted brown lemur Eulemur albifrons) and a species of the carnivore family Eupleridae (fossa Cryptoprocta ferox). Model results suggest hunting of these species is probably unsustainable. This research presents clear evidence that hunting is a major conservation and livelihoods issue in Madagascar and needs to be considered in the planning stages of protected area development to address better the needs of local people.