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Current project details and status

Current project: Lemur conservation

Start Date: 14 July 2009
Target Completion Date: early February 2010
Special Notes: This will be a summary page for the new Lemur article. See Project Lemur for project overview. It will be published at the same time as the Lemur article re-write.

Progress:

• Research and note taking –  Done
• Page organization –  Done
• Organization of research notes –  Done
• Wikifying all references for use in text  Done
• Body written –  Not done
• Lead written  Not done
• Spell-checking and copyediting  Not done
• Publication  Not done

Main article: Lemur

...

New research prior to writing:

• None

Note: detailed stats about Madagascar (poverty, roads, economics, politics, etc.) can be obtained at: http://www.estandardsforum.org/system/briefs/275/original/brief-Madagascar.pdf?1261005963 (from http://www.estandardsforum.org/)

• for pet trade news from Northern Ireland: http://news.bbc.co.uk/2/hi/uk_news/northern_ireland/8527153.stm

Overview

...

oldWiki: Most lemurs are listed as endangered or threatened species. Many species have gone extinct in the last centuries, mainly due to habitat destruction (deforestation) and hunting. Conservation of lemurs in Madagascar is a high priority, but the country's poor economic situation and the lemurs' limited range make it an uphill battle. In 2008, a total of 99 living lemur species were formally recognized, with more species likely to be discovered or differentiated in the future. ... One of the foremost lemur research facilities is the Duke Lemur Center.

-"Georaphically restricted genera are all monotypic; these include the still extant Allocebus and Lemur, as well as extinct taxa (Babakotia, Archaeoindris, and Hadropithecus). About half of the extant genera have representatives in every major eco-geographic zone of Madagascar, and the same was probably true of the extinct genera."

-"Madagascar is the only place where primates genera are the dominant group overall. On a global scale primates rank 5th behind rodents, bats, carnivores, and even - toed hoofed mammals."

-"How did Madagascar achieve this position? Madagascar was one of the last great habitable land masses settled by humans. According to the comprehensive review by Burney et al. (2004), multiple points of evidence date the earliest presence of humans at ca. 350 yr BC. A decline in megafauna at around 230-410 AD is followed by large increases in charcoal particles in sediments signalling increased human impact on the landscape. When Madagascar was discovered by Europeans in 1500, almost all of the Malagasy megafaua - pygmy hippos, elephant birds, giant tortoises, large lemurs - had already disappeared. This was the result of the synergistic combination of human impact, nonlinear natural responses and environmental change, population fragmentation, and local extirpation"

-"On a world wide scale, four of these lemurs (Greater bamboo lemur, White - collared lemur, Silky sifaka, Perrier’s sifaka) rank in the top 25 most endangered primates"

-world's top 25 most endangered primates for 2008-2010 include: Prolemur simus, Eulemur cinereiceps, Eulemur flavifrons, Lepilemur septentrionalis, and Propithecus candidus

-"Lemur conservation equals forest conservation. Unique as lemurs are biologically, diverse, fluffy and sympathetic, they may serve as flagship species for the whole island. With regionally occurring endemic species they may even serve as flagships for different regions within Madagascar... Protecting and conserving lemurs will help to protect a plethora of other species as collateral effect. Because lemur conservation is forest conservation, the protection of lemurs also helps to grant important services by forests, such as reduced erosion, clear and sustainable water proliferation – a better life for humans."

-"Direct archaeological evidence for human habitation is surprisingly late for Madagascar--scarecely more than a millennium ago. Indirect clues point to human visitation of some sort beginning almost twice that long ago. These include human-modified bones of hippo and elephant bird and stratigraphic evidence for 1) pollen of plants introduced by humans, 2) a suddent drastic increase in microscopic charcoal particles, and 3) local vegetation and soil disrubance." (Natural History, p. 49)

-"At present, it is clear that there was a period of one to several centuries when people and the now extinct subfossils were coexisting on the island of Madagascar. It remains difficult to link the colonizing people with the specific processes responsible for the extinctions. Hunting may have played a role, but there is little evidence in the archaeological record to document it. Habitat transformation by clearing, burning, or livestock may have been important, but it is wise to remember that fairly large areas of Madagascar were covered by intact forest well into the twentieth century. The archaeological evidence suggests that human activities differed from one region to another, and so we can imagine that the human role in environmental changes must also have differed in distant parts of the island. It may well be that the mix of human impacts and the timing of the extinctions were not the same across the entire island." See entire section for details. (Natural History, p. 122)

-"Given this remarkable rate of new mammal species being described from the island on one hand and the continued destruction of the remaining habitat of these animals on the other, it is possible that mammal species may become extinct before they are known to science." (Natural History, p. 1186)

-"Madagascar is one of the world's highest-priority areas for biodiversity conservation. It ranks among the top eight megadiversity countries and is one of the five highest-priority biodiversity hot spots. When one takes into account its msall land area (587,045 km2) realtieve to that of many of the other high-priority countries such as Brazil, Indonesia, Colombia and Australia, its conceration of life forms becomes even more striking. ... The majority of the country's natural vegetation has already been lost, with estimates of what remains usually on the order of 9-15%." see test for comparison of taxa count, country size, and levels of endemism (Natural History, p. 1538)

-"Madagascar is so important that the World Conservation Union's Species Survival Commission (IUCN/SSC) Primate Specialist Group has recoginzed the island nation as its own major priamte region for more than 20 years." (Natural History, p. 1538)

-"Many of the lemur speices that have been recently described but whose conservation status has not yet been fully assessed appear to have restricted distributions. Some of them could very well be threatened, but for the moment they remain in the Data Deficient category." (Natural History, p. 1541)

-"The Ring-tailed Lemur is the flagship species, an icon synonymous with its island home. Paradoxically it is far from being typical; it is the most terrestrial of Madagascar's primates, lives in the largest social groups and tolerates a variety of extreme habitats that no other lemurs can." (Garbutt, p. 146)

-Madagascar is "one of the world's most endagered "hotspots" and places it among the top 5-6 of the world's 18 "megadiversity" countries. What's more, the most recent information on endemic families and genera indictates that it is far and away the world leader in these categories as well, with far more unique evolutionary lineages than any other hotspots." (Mittermeier, p. 53)

-mention umbrella species

Threats in the wild

...

-more details on specific threats to lemurs (Mittermeier, p. 56-71) --use below as needed...

-"Today, the major threats to lemurs include deforestation due to slash-and-burn agriculture (known as tavy in Madagascar), logging fuelwood collection, charcoal production, the seasonal burning of dry forests to create cattle pasture, and live capture of lemurs as pets. Sadly, the hunting of lemurs as a source of food, a threat that we used to consider less severe than many others, is now emerging as a major problem in many areas as new data become available, and needs special attention." (Mittermeier, p. 15)

-Habitat destruction is the greatest concern, but hunting also occurs (see text). Both factors probably drove the largest species extinct.(IUCN, p. 11)

-"All endemic vertebrates on Madagascar over approximately 10 kg are extinct, as well as many smaller ones. ... There is no evidence in the prehistoric record suggesting that the "extinction window" has closed or that the sad changes wrought in vegetation, soils, and local climate have reached their end point." (Natural History, p. 50)

-"Like most lemurs, Eulemur species are primarily threatened by a reduction in forest area and increasing fragmentation of suitable habitats. This situation is exacerbated in some species and regions by hunting, the pet trade, and extermination as agricultural pests." (Natural History, p. 1324)

-"the entire remaining habitat available to lemurs is at best no more than 10% of the country's land area--60,000 km2... Furthermore, at this time, only 3% of Madagascar's land area is protected, an area of about 17,000 km2... and not all of that is effectively managed for the long term..." (Mittermeier, p. 17)

Effects of human overpopulation and poverty: -"As in Africa, poverty is widespread in Madagascar and the rate of population increase here is even higher than in Africa, at 3.2% per year." (Sussman, p. 267)

Habitat destruction, degradation, and fragmentation

-"Masoala National Park faces far more managerial challenges given all of its waterways and the difficulty of accessing the eastern boundary. However, land cleared for marijuana plots has recently been observed within the remote northern sector a few kilometers from the park boundary"

-"...indicate that forest degradation and fragmentation have marked effects on the level of parasitism of Madagascar’s lemurs."

-"Countries of the Eurpoean Union are the most important market of tropical timber from Africa, accounting for almost 90% of the market." (Sussman, p. 266)

-"...the main threat to all species is habitat desruction. Forest is cut mainly for local subsistence purposes, though rently commerical mining has become a problem in some regions." (Sussman, p. 267)

-"In the eastern rain forests of Madagascar, conservation efforts have been directed toward natural reserves and already protected areas, but most of these reserves were established in remote, isolated, and steep areas so they have not impinged upon the current need for land.... In other regions of Madagascar, the need for firewood or charchoal for fuel, or to create grassland for cattle leads to forest cutting. In fact, the dry forest of the west and south are probably in more danger than are the eastern rain forests." (Sussman, p. 268)

-"Among lemurs there appear to be speices that are very sensitive to forest disturbance, such as Varecia variegata, whereas others have higher popultion densities in slightly disturbed habitats, such as Microcebus and Cheirogaleus." (Natural History, p. 1184)

-"Habitat destruction ... are the primary threats to Daubentonia populations." (Natural History, p. 1351)

-"the majority of the country's protected areas are still inadequately defended against encroaching small-scale agriculture (Tavy), clearing and buring for pastureland, and hunting, the three principal threats to wildlife and habitat in this country." (Natural History, p. 1542)

-"While loss of key fruit trees may not drive lemurs to extinction immediately, it may adversely affect reproducte success years after logging... This kind of "energetic debt" can affect reproduction decades after the logging has ceased." (Lemurs, p. 392-393)

Habitat fragmentation:

-"It is important to know which species can persist for how long in fragments of different sizes. Many mammals, and lemurs in particular, have relatively long generation times and poor dispersal abilities (when compared with birds) and can potentialally persist in small forest fragments for considerable time spans." ... "It could be estimates that lemur populations of about 40 adult individuals are unable to survive for more than 20 to 40 years in isolated forest fragments. Depending on their body mass, the lartgest lemur speices would require forest of around 1000 ha to maintain viable populations for more than 20 to 40 years." (Natural History, p. 1228 & 1234)

-"Like most lemurs, Eulemur species are primarily threatened by a reduction in forest area and increasing fragmentation of suitable habitats..." (Natural History, p. 1324)

-"However, when populations are extremely small or fragmented, stochastic events such as disease outbreaks and epizootics may have catastrophic effects." (Lemurs, p. 423)

Hunting and taboo

...

Poaching

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  Three dead bamboo lemurs (Hapalemur griseus), killed for bushmeat in northeast Madagascar
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-Aye-aye as a pest and taboo (see text) (Madagascar Natural History, p. 188)

-"Although hunting and eating lemurs is taboo for many Malagasy, these animals are hunted in many regions." (Sussman, p. 210)

-"Even though hunting of certain lemurs is traditionally taboo in many regions of Madagascar, and currently is against the law, this still remains a problem. The distribution of the larger species away from villages, even where suitable habitat exists, is testimonial to this fact. However, the small nocturnal forms are not usually hunted..." (Sussman, p. 267)

-Habitat destruction is the greatest concern, but hunting also occurs (see text). ...In several regions, it is taboo to kill lemurs, but people without those traditions are becoming more mobile. Black-and-white Ruffed Lemurs are hunted for meat. Aye-ayes are killed on sight (but not hunted) because local people think they bring bad luck. Smaller nocturnal species are not actively hunted, but can be caught in snares.(IUCN, p. 11)

-"Within both nonprotected areas and relatively remote poritions of reserves it is not uncommon to find lemur traps. Further, organized hunting parities utilizing firearms, slings, and blowguns are known from many areas of the island. ... organized hunting paries most of the year... E. fulvus are brought back to the villages for local consumption. The cost of a lemur is based on the animal's size... Further, a group of experienced hunters generally return with 8 to 20 lemurs. Such heavy hunting pressure is having a substantial impact on primate populations, an effect local hunters have also noted." (Natural History, p. 1184)

-"Local cultural taboos have both positive and negative effects on certain species. For several tribes the consumption of lemur flesh is forbidden based on local customs, and these animals are therefore not hunted. There are many examples of this across the island. However, with the immigration into a given area of another cultural group that does not follow the same tradition, or with change in local traditional practices associated with ecological crises, such as drought and associated famine, these customs are notably relaxed. ... One example of a taboo that has negative effects on mammal speices concerns the aye-aye. Among certain groups of Sakalava living in the northwest, the aye-aye is considered an extremely dangerous animal because it is reputed to attack, kill, and consume chickens living in the village. Further, it is thought to enter houses during the night through thatched roofs and murder the sleeping human occupants with its elongated finger that cuts the aortic vein in its victims. It is for this reason that Sakalava kill aye-aye when they have a chance." (Natural History, p. 1185)

-"Like most lemurs, Eulemur species are primarily threatened by ... This situation is exacerbated in some species and regions by hunting, ..., and extermination as agricultral pests." (Natural History, p. 1324)

-"Habitat destruction and killing by humans are the primary threats to Daubentonia populations. Animals are occasionally killed when stealing crops from gardens, but the greatest threat appears to be local taboos, or fady." (Natural History, p. 1351)

-"Although it is taboo for the human tribes who live in the same area as the (Verreaux's) sifakas to kill the animals, they are hunted by others." (Anatomy, p. 75)

-"Villagers hold aye-ayes in dread and will move away from places where they have been seen. In many areas of Madagascar, meeting an aye-aye means death or bad luck in the family." (Anatomy, p. 80)

-"All lemurs are threatened by habitat loss and hunting; the latter threat is particularly acute in some areas and has largely been overlookeed in the past. Larger species like sifakas are obvious targets, but so too are many of the smaller species. In many areas lemurs are hunted in remote localities to provide bushmeat, both locally and for more distant larger villages and towns." (Garbutt, p. 85-86)

-"Perhaps the most ominous of these is the fact that the majority of rural Malagasy people do not know what endangered means, do no not know that hunting lemurs is prohibited by law, and do not know that lemurs occur only in Madagascar. It is easy for a poacher to walk from the edge to the center of most of Madagascar's parks and reserves in part or all of a single day. These parks and reserves do not have adequate protection. Blowguns and slingshots are abundant and are used. The hunting of lemurs for food is practiced almost everywhere... the judicial and law enforcement agencies are currently not effective in controlling this exploitation. ... Although in many areas there are fady that protect some lemurs, they are suually very specific for only certain lemurs, not others. These fady are breaking down as human population movements within Madagascar increase and people move in from other regions where there are no fady. For example, the Sakalava tribesmen around Daraina in the northeast say that they do not hunt the ankomba malandy (P. tattersalli), but there has been a recent gold rush in the small area where this species occurs, and it has brought in many people who do not observe this taboo." (Natural Change, p. 154)

-"No one knows how many lemurs have been killed, but species such as the golden crowned sifaka—considered endangered by the International Union for Conservation of Nature—are being targeted." --see article

-"If there are no specific restrictions, the Natives eat all kinds of lemurs. Among the Vazimba and Mikea, the meat of lemurs is very common food. They are are caught in snares and traps, or killed with sticks and stones. It is even more common to shoot them with blow-pipes and bamboo-arrows."

Human overpopulation

...

-"the desire to have many descendants is almost universal in rural Madagascar... rural Malagasy people’s wish to have numerous descendants is intimately linked to their understanding of what, at a very fundamental level, represents a successful and meaningful life" -- conservation literature focuses too strongly on fady when it discusses Malagasy culture.

Pet trade

-as pets, H. griseus live an average of 2 months (Pictorial, p. 46)

-"Trade is not considered a threat, while Madagascar and Comoros have strict regulations controlling their export.(IUCN, p. 11)

-"Like most lemurs, Eulemur species are primarily threatened by ... This situation is exacerbated in some species and regions by ...the pet trade..." (Natural History, p. 1324)

-"Crowned lemurs are, unfortunately, widely used as pets by the Malagasy people." (Anatomy, p. 69)

Introduced pests and parasites

-"In large stable populations, disease is a normal part of population dynamics. However, when populations are extremely small or fragmented, stochastic events such as disease outbreaks and epizootics may have catastrophic effects. In some cases, veterinary intervention during a disease outbreak is essential to prevent extinction of the population." (Lemurs, p. 423)

-geographic isolation, disease transmission, immunologically naive populations, etc. (Lemurs, p. 424)

-introduced diseases (Lemurs, p. 425)

-"A review of the current literature provides scant information on the diseases of wild lemurs, No major epizootics have been reported." (Lemurs, p. 427)

-toxoplasmosis, herpes simplex, etc. (Lemurs, p. 433-434)

-"In most cases introduced ticks are those of cattle or carnivores (dog) and therefore rarely come into contact with native or introduced small mammals... No exchange of lice between endemic and and introduced small mammals has been observed to date." (Natural History, p. 1193)

Climate change

-"Although we know that there has been a dramatic dessiccation of western and highland regions in the past thousand years, there are indications that deforestation and fragmentation of forests are presently continuing to produce a drier climate in Madagascar. Although drought is known to be a natual phenomenon, especially in the south of Madagascar, droughts may be becoming more frequent." (Lemurs, p. 393)

-"Severe drought years have been shown to impact dry, western forests in Madagascar. But droughts also affect the rainforest by extending the dry season and reducing annual rainfall. Drought in rainforests has resulted in high canopy tree mortality, reproductive failure, fruit crop failure, and decrease in young leaf abundance." (Lemurs, p. 413)

-"No long-term studies have been conducted yet on the effects of natural disasters such as drought and cyclones on fauna and flora in the Manombo region, However, direct cyclone hits can result in complete defoliation, blowdowns of all canopy trees, landslides, and flooding. As a consequence, neither fruits nor leaves may be available for consumption until the following spring, leaving only crisis foods, such as epiphytes, for the lemurs to eat." (Lemurs, p. 413)

Effects of political instability

Main article: Illegal logging in Madagascar

... (material relocated to Sandbox9)

-"James MacKinnon, senior technical director at the Madagascar office of Conservation International, says the looting includes illegal mining and poaching of rare animals for the international pet trade. "There's a rush on high-value resources," he says. "They're taking advantage of the situation to make a quick buck."

-"At the same time that northern rainforests were being pillaged for their timber, a disturbing new trade emerged: commercial bushmeat hunting of lemurs. In August, Conservation International (CI) released photos showing piles of dead lemurs that had been confiscated from traders and restaurants in northern towns. ... And it is not for subsistence, but rather to serve what is certainly a 'luxury' market in restaurants of larger towns..."

-"Tourist arrivals dropped 50 to 60 percent for the year, badly affecting workers employed in the tourism industry. ... In Ranomafana, tourist arrivals are down by more than 40 percent. Before the coup, in 2008, 24,000 tourists visited the park, generating $1.72 million in revenue"

-"Conservation in Madagascar is highly dependent on income from tourism. Half of park entrance fees are returned to communities living in and around protected areas. Without this source of income, locals in some areas may be forced to turn to conservation areas for timber, fuelwood, and agricultural land."

-re-visit this source for more info under this section...

Conservation efforts

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-See conservation history (IUCN, p. 12)

-"(the rural, relatively isolated human populations), not just the central government, need to have input into programs that have a direct input into their daily lives. However, there are often conflicting goals and aspirations among the scientific community, international conservation organizations, global agencies, national and local governments, and the local people. These conflicts are often extremely complex, with each group having real and legitimate problems related to conservation. Because of this, conservation often becomes as much, or more, a human social problem as it is a problem of purely the biological and natural sciences." (Sussman, p. 266)

-"Considerable world attention has been drawn to the island's primates, and biologists and conservationists have used these animals as a "flagship" for the plight of Madagascar's unique and endangered biota." (Natural History p. 1184)

-"Varecia is also an indictaor, umbrella, and flagship species." (more in text) to frugivores susceptible to fragmentation (p. 1334) (Natural History, p. 1336)

-"The Malagasy government declared all lemurs "protected" in 1927." (Natural History, p. 1351)

-"At present, about 2.9% (16,855 km2) of the country's land surface is represented in protected areas -- these total 49 including 16 PN, 7 RNIs, 23 RSs, and 3 small Reserves Privees. With the exception of the priamte reserves, the netowrk is managed by the Association Nationale pour la Gestion des Aires Protegees (ANGAP), a quasi-governmental body created in 1989 and funded mainly by interational donor agencies. ... Lemurs are reasonably well covered (by the ANGAP network), with all but a few taxa currently recognized occurring in one or more parks and reserves." (Natural History, p. 1542)

-"Another key element in a long-term strategy to conserve lemurs and their habitats in Madagascar is the creation of conservation corridors to link parks and reserves in broader landscapes. However, the first piority must be to ensure the long-term viability of the parks and reserves themselves and to fast-track creation of new ones in the highest priority areas." (Natural History, p. 1542)

-conservation work by the MFG (detailed) (Natural History, p. 1543–1545)

-H. g. alaotrensis, 3500-5500 individuals left, limited to 220 km2, no national park system to protect -- "Durrell Wildlife Conservation Trust have demonstrated long-term commitment in the region by promoting awareness and maintaining a steady stream of research and monitoring programs..... a breeding program managed by Jersey Zoo is currently in place." (Lemurs, p. 378)

-"In light of this, it is particualrly encouraging that Madagascar's President, Marc Ravalomanana, has commited to tripling his country's protected area coverage over the next five years. This historic declaration, made at the World Parks Congress in Durban, South Africa, in September 2003..." (Mittermeier, p. 17) more on "The Durban Vision" (Mittermeier, p. 77-78)

-"The national netowrk of protected areas in Madagascar was initited in 1927 and now includes 45 terrestrial protected areas in 3 categories: national parks (18), special reserves (22), and strict or integral nature reserves (5).... Protected area management is overseen by the National Association for the Management of Protected Areas (ANGAP) but, in a number of cases, responsibility for managing individual parks and reserves is also shared by international non-governmental organizations such as Conservation International, the Wieldlife Conservation Society, and the World Wide Fund for Nature." (Mittermeier, p. 77)

-"Education and public awareness in Madagasar are priorities at all levels, from government decision-makers to city-dwellers to local people living in the most remote villages. Education efforts using lemurs and other speices, have been carried out sporadically in the past, employing materials such as t-shirts, posters, stickers, pins, brochures, and a variety of other products." (Mittermeier, p. 81)

-"With 70 percent of Madagascar’s population living below the poverty line, the country is one of the poorest in the world. Burdened with high levels of debt, Madagascar has limited domestic resources to address environmental degradation and preserve its unique and globally significant biodiversity. Debt-for-nature swaps, such as this one, are designed to free up resources in debtor countries for much needed conservation activities."

-P. simius: "Researchers in Madagascar have confirmed the existence of a population of greater bamboo lemurs more than 400 kilometers (240 miles) from the only other place where the Critically Endangered species is known to live, raising hopes for its survival."

-"Moreover, in June 2007, the World Heritage Committee has named a significant proportion of Madagascar’s eastern rainforests as one of three new UNESCO World Heritage List sites."

• Captive breeding and captive releases:

- see Global-Reintro-Perspectives.pdf for full update on Varecia reintroduction

-Project Betampona: release of captive Black-and-white Ruffed Lemurs, high mortality, especially from predation (but not wild population); free-ranging individuals (group 1) adapted better than cage-reared lemurs (group 2); better to restock young males since they disperse; habitat must be surveyed for adequate year-round resource availability. (Natural History, p. 1545–1551)

-Indriids: "None of these folivores survives well in captivity, although the DLC ... has been very successful with Propithecus." (Sussman, p. 200)

-"Indri has not been successfully managed in captivity." (Natural History, p. 1342)

-Conservation in captivity (Mittermeier, p. 78-80)

External links

• Madagascar Rainforest Massacre on YouTube. (Also available in French and Malagasy.)
• Field investigation into illegal logging in Madagascar – videos and other downloads from the Global Witness Media Library

References

• Ankel-Simons, Friderun (2007). Primate Anatomy (3rd ed.). Academic Press. ISBN 978-0-12-372576-9.
• Garbutt, Nick (2007). Mammals of Madagascar, A Complete Guide. A&C Black Publishers. ISBN 978-0-300-12550-4.
• Goodman, Steven M.; Benstead, Jonathan P., eds. (2003). The Natural History of Madagascar. University of Chicago Press. ISBN 0-226-30306-3.
• Goodman, Steven M.; Patterson, Bruce D., eds. (1997). Natural Change and Human Impact in Madagascar. Smithsonian Institution Press. ISBN 978-1560986829.
• Gould, Lisa; Sauther, Michelle L., eds. (2006). Lemurs: Ecology and Adaptation. Springer. ISBN 978-0387-34585-7.
• Harcourt, Caroline (1990). Thornback, Jane (ed.). Lemurs of Madagascar and the Comoros: The IUCN Red Data Book. World Conservation Union. ISBN 978-2880329570.
• Mittermeier, R. A. (2006). Lemurs of Madagascar. Illustrated by S.D. Nash (2nd ed.). Conservation International. ISBN 1-881173-88-7. {{cite book}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
• Nowak, R.M. (1999). Walker's Mammals of the World (6th ed.). Johns Hopkins University Press. ISBN 0801857899.
• Preston-Mafham, Ken (1991). Madagascar: A Natural History. Facts on File. ISBN 978-0816024032.
• Rowe, Noel (1996). The Pictorial Guide to the Living Primates. Pogonias Press. ISBN 978-0964882515.
• Sussman, Robert W. (2003). Primate Ecology and Social Structure. Pearson Custom Publishing. ISBN 978-0536743633.