New Lemur Species

New Primates Described in 2005

The July issue of Primate Report (No. 71) and the November issue of the American Journal of Primatology (Vol. 67, No. 3) provided formal descriptions of three new lemurs – a giant dwarf lemur, a mouse lemur and a woolly lemur. Science published a description of the highland mangabey in its Volume 308 on 20 May 2005, and Sinha et al. (2005) described the Arunachal macaque in the fourth issue of Volume 26 of the International Journal of Primatology, published in August 2005.

Three New Species of Lemurs

The northern giant dwarf lemur
The northern giant dwarf lemur, Mirza zaza, was described in Kappeler et al. (2005) by Peter M. Kappeler and Christian Roos, both of the German Primate Center, Göttingen. It is smaller than the only other member of the genus currently recognized (Coquerel’s mouse lemur, M. coquereli) and occurs well to the north, in northwest Madagascar in Ambato and Pasandava (locations separated by several hundred kilometers) in the Province of Ansiranana. The holotype is an adult male collected on 25 September 1865 at Congoni, Ampasindava Peninsula, 13º40’S, 48º15’E (skin and skull in the Leiden Museum). The argument for the distinctiveness of M. zaza was based on comparisons of the morphology, genetics and behavior with M. coquereli, found well to the south at Kirindy, in central western Madagascar.

Goodman’s mouse lemur
Kappeler et al. (2005) also included the description, this time authored by Roos and Kappeler, of a new species of mouse lemur named after Steven M. Goodman: Microcebus lehilahytsara Roos and Kappeler, 2005. The description was a based on a type series of nine live animals housed in the Zürich Zoo, Switzerland, obtained by Samuel Furrer and Robert Zingg in March 2005 from Andasibe, Province of Toamasina, eastern central Madagascar, 18º55’S, 48º25’E. Goodman’s mouse lemur (lehilahy is Malagasy for “man” and tsara means “good”) is one of the smallest members of its genus, measuring about 9 cm head and body length (similar to M. berthae), and is distinctive in its fur color, external morphology (for example, short round ears) and genetics (analysis of mitochondrial cytochrome b gene).

Cleese’s woolly lemur
Urs Thallmann and Thomas Geismann (both of the Anthropological Institute of the University of Zürich), described a new species of woolly lemur or avahi: Cleese’s woolly lemur or the Bemeraha woolly lemur, Avahi cleesei Thalmann and Geismann, 2005. It was named in honor of the British comedian and actor John Cleese for his promotion, in films, talks and documentaries, of conservation issues concerning lemurs. The species was first mentioned in the description of Avahi unicolor (Thalmann and Geissman , 2000, but it was not described due to the lack of a type specimen. The authors still lack a conventional type specimen, but took courage from a publication by Wakeham-Dawson et al. (2002), who argued that it was not necessary. The species is described as such by hair samples, photographs, and audiotape recordings stored in the Anthropological Institute of the University of Zürich. It occurs in central western Madagascar, in the Tsingy de Bemaraha region north of the Manambolo River; the type locality is given as approximately 3 km east-northeast of the village of Ambalarano, western Madagascar, 18º59’S, 44º45’E. The holotype, not sacrificed for ethical reasons, was captured in Bemaraha on 3 October 1991. It is distinguishable from A. occidentalis by its lack of a white facial mask and broad dark eye rings, and from both A. occidentalis and A. unicolor by the presence of a dark chevron pattern on the forehead.

The first edition of the Lemurs of Madagascar field guide (Mittermeier et al., 1994) listed 50 taxa. With the publication of these three species, the number has now reached 68; a second edition of the lemur field guide, which will soon go to press, lists 71 (including three still unnamed mouse lemurs), and many more lemurs remain to be described in the coming years.

Two New Cercopithecine Species – a Mangabey and a Macaque

The highland mangabey, from the southern highlands in Tanzania—(Lophocebus kipunji Ehardt, Butynski, Jones and Davenport, in Jones et al., 2005)—was described in May 2005 in Science (Volume 308), and the Arunachal macaque from the far northeast of India, in the state of Arunachal Pradesh—(Macaca munzala Sinha, Datta, Madhusudan and Mishra, 2005)—was described in August 2005 in the International Journal of Primatology (Volume 26).

The highland mangabey
The highland mangabey was discovered independently at two localities in southern Tanzania—Ndundulu in the Udzungwa Mountains, and the Mount Rungwe Forest Reserve and Livingstone Forest in the Southern Highlands (Beckman, 2005; Boy, 2005; Jones et al., 2005). Although seen first in the Mount Rungwe forest in May 2003, they were first observed clearly only in Livingstone in December 2003. The same species was then seen by a second group of researchers in the Udzungwa Mountains in July 2004. The two groups found out about each other’s discoveries in October 2004. Due to its rarity, it was decided that none should be killed in order to provide a holotype. The description was instead based on a photograph and observations of an adult male from the Rungwe-Livingstone population. The type locality was recorded as “Rungwe-Livingstone (09º07’S to 09º11’S and 33º40’E to 33º55’E), Southern Highlands, Tanzania.” (This resulted in some controversy; see below.) A second photograph of an adult (sex unknown) from the Ndundulu Forest Reserve in the Udzungwa Mountains was registered as a paratype. The diagnostic features of this mangabey are as follows: the pelage of the dorsum is light to medium brown; the center of the ventrum and distal half of the tail are off-white; the crown has a very long, broad, erect crest of hair; the eyelids are black, not contrasting with the color of the face; the adults emit a distinctive, loud, low-pitched “honk-bark” (Jones et al. [2005] provide an illustrative sonogram of a series of honk-barks, prepared and interpreted by Andrew Perkin); it is found at altitudes of 1300 to 2450 m above sea level. The Kinyakyusa name for this mangabey in Rungwe-Livingstone is kipunji. The habitat of the populations known to date is pristine submontane forest in Ndundulu and degraded montane and upper montane forest in Rungwe-Livingstone. At the time of its description, 10 groups had been located in Rungwe-Livingstone, indicating a total population of 250-500 animals. They are, on the other hand, known from only 3 km² of forest in Udzungwa, although Jones et al. (2005) estimated 3-50 km² as their total range there and a population of less than 500 animals overall. Forest degradation, fragmentation, logging, charcoal-making, poaching and unmanaged resource extraction are severe threats to the species, and Jones et al. (2005) predict that its status will be Critically Endangered once a full assessment is made.

The Arunachal macaque
The fact that the higher elevations of the state of Arunachal Pradesh (>3000 m) have been so poorly inventoried was given as the reason that the distinctive Macaca munzala had never been previously described. In the local dialect of Dirang Monpa, mun means deep forest and zala is the word for monkey. It was described from photographs of an adult male taken by M. D. Madhusudan in a group of 17 individuals in Zemithang (27º42’N, 91º43’E) in Tawang District, Arunachal Pradesh, at an altitude of 2180 m above sea level. Sinha et al. (2005) recorded it from 13 localities in the district of Tawang, and one (Nawrok) in the district of West Kameng. All but one of the localities were at elevations of 2000-3000 m (the exception was 1650 m), and Sinha et al. point out that the species occurs at the highest altitude of any of the macaques of the Indian sub-continent. Its known range is about 1200 km², but further surveys may find it in other parts of Arunachal Pradesh, and even across the border in adjoining areas of Tibet and Bhutan. They are seen around villages and often go into cultivated areas, but otherwise their typical habitat is the subtropical broadleaved forests which are the predominant vegetation of the region. It is classified as a member of the Macaca sinica species group (see Fooden, 1980) due to its penile morphology. The adult pelage of this macaque is varying shades of dark brown to dark chocolate. Its chief characteristics include its relative tail length (a little less than half the length of its back) which distinguishes it from M. sinica, M. radiata, M. assamensis pelops and M. thibetana, and distinctive facial features which distinguish it from M. assamensis assamensis, which occurs in Arunachal Pradesh further to the east.

Questions of Description

The publication of the description and name of Lophocebus kipunji was based entirely on sightings and two photographs, and resulted in a letter being published in Science (vol. 309, pp.2163-2164, 2005) by Robert M. Timm and Rob Roy Ramey II on behalf of the Nomenclature Committee of the American Society of Mammalogists, which argued that the species’ description is not valid: “…the photographs do not function as name-bearing types [reference to Article 16.4 of the International Code of Zoological Nomenclature of the International Commission on Zoological Nomenclature (ICZN), London, ed. 4, 1999]. Thus, Lophocebus kipunji Ehardt, Butynski, Jones, and Davenport is not an available name and has no formal standing in zoology…The photographs are not valid substitutes for a type specimen.” (p.2163). Stuart O. Landry (2005) also published a letter in the same issue of Science arguing that the name kipunji was a nomen nudum because it was not based on a properly designated type specimen.

These arguments are, of course, pertinent to the description of Macaca munzala—also based on photographs. In addition, Cleese’s woolly lemur, Avahi cleesei Thalmann and Geismann, 2005, was based not on a type specimen but on hair samples, photographs, and audiotape recordings. Indeed, Thalmann and Geissman mentioned the Bemeraha form when describing Avahi unicolor in 2000, but made no formal description then due to the lack of a type specimen.

Andrew Polaszek (Executive Secretary of the International Commission on Zoological Nomenclature), Peter Grubb, Colin Groves, Carolyn L. Ehardt and Thomas M. Butynski (2005) provided a reply to Timm et al. and Landry in the same issue of Science. They reassured them that, in describing Lophocebus kipunji, Jones and his co-authors had undertaken a series of consultations with the ICZN Secretariat and several eminent taxonomists regarding the validity of the use of photographs instead of a prepared specimen. They pointed out that Article 16.4.2 of the International Code of Zoological Nomenclature demands that authors of new taxa must publish a statement of intent that extant types will [italics added here] be deposited in a collection, while Article 73.1.4 allows for the description of new taxa without the need to provide dead type specimens. This situation was discussed by Wakeham-Dawson et al. (2002), who were cited for this reason by Thalmann and Geismann (2005) in their description of Avahi cleesei. Polaszek et al. argued that “the allowance under the code for designation of surviving specimens as holotypes needs to be more widely recognized, given contemporary concerns for the conservation of threatened species” (p.2165), and insisted that dead animal specimens should not be understood to be essential to the process of establishing new taxa.

Landry (2005) also pointed out that identifying the responsibility for the description of Lophocebus kipunji as “Ehardt, Butynski, Jones, and Davenport” is not valid because they are only four of the seven authors of the publication (Jones et al., 2005) and “the purpose of the citation is to identify the paper, not to assign credit, and all the authors should be cited.” (p.2164). This observation is relevant also to the description of the two lemurs in the article of Kappeler et al. (2005). Five people authored the article, but just two were given as authors of the species’ descriptions: Kappeler and Roos of the northern giant dwarf lemur, Mirza zaza, and Roos and Kappeler of Goodman’s mouse lemur, Microcebus lehilahytsara. This procedure is, however, valid and even recommended by the ICZN (1999). Chapter 11 of the ICZN deals with “Authorship”, and Recommendation 50A reads as follows:

“Multiple authors. When a name is proposed in a multi-authored work, but only one (or some) of the authors is (are) directly responsible for the name and satisfying the criteria that make the name available, then the author(s) directly responsible should be identified explicitly. Co-authors of the whole work who have not had such direct responsibility for the name should not automatically be included as authors of the name. See Recommendation 51(E) for citing the names of such authors.” (p.52).

For the two lemurs and Lophocebus kipunji, it is necessary to cite both the authors of the species names and the authors of the articles in which the descriptions were published, as recommended in ICZN (1999; Recommendation 51E, pp.54–55). The proper citation for the highland mangabey is, therefore, Lophocebus kipunji Ehardt, Butynski, Jones and Davenport in Jones, Ehardt, Butynski, Davenport, Mpunga, Machaga and De Luca, 2005. Those for the two lemurs are Mirza zaza Kappeler and Roos in Kappeler, Rasoloarison, Razafimanantsoa, Walter and Roos, 2005, and Microcebus lehilahytsara Roos and Kappeler in Kappeler, Rasoloarison, Razafimanantsoa, Walter and Roos, 2005. This as such gives the authorship of the species and the scientific reference where it was published. Cumbersome, but there we are. Note, however, that Recommendation 51C of ICZN (1999) allows for the use of the term “et al.” following the name of the first author for the citation of three or more authors, provided that all authors of the name are cited in full elsewhere in the same work, either in the text or in a bibliographic reference. The authorship of Lophocebus kipunji could, therefore, be written as “Lophocebus kipunji Ehardt, Butynski, Jones and Davenport in Jones et al., 2005” if Jones et al. (2005) is in the bibliographic references, or if the authorship of the species has also been written in full previously or is noted with the bibliographic reference: “Lophocebus kipunji Ehardt et al. in Jones et al., 2005”.

Polaszek et al. (2005) argued that supplementary evidence (such as from genetic analyses, vocalizations and hair samples) is important but that the description of new species using photographs is open to abuse, and that an obvious measure to avoid this would be to introduce a registration system for animal names in order to alert zoologists to the appearance of new species and to ensure that the names are “code-compliant.” Incorporating the power of bioinformatics in this sense, and in order to clarify ambiguities and avoid difficulties in interpretation, Polaszek et al. suggested that a revision of the 1999 ICZN would already be timely.

Anthony B. Rylands (Center for Applied Biodiversity Science at Conservation International) and Russell A. Mittermeier (President, Conservation International), IUCN/SSC Primate Specialist Group, c/o Conservation International, 1919 M Street NW, Suite 600, Washington, DC 20036.


Beckman, M. 2005. Biologists find new species of African monkey. Science 308(5725): 1103.
Boy, G. 2005. 'Parallel' discovery. Swara 28(3): 28-32.
Fooden, J. 1980. Classification and distribution of living macaques (Macaca Lacépède, 1799). In: The Macaques: Studies in Ecology, Behavior and Evolution, D. G. Lindburg (ed.), pp.1-9. Van Nostrand Reinhold, New York.
ICZN. 1999. International Code of Zoological Nomenclature. 4th Edition. International Commission on Zoological Nomenclature (ICZN), c/o The Natural History Museum, London.
Jones, T., Ehardt, C. L., Butynski, T. M., Davenport, T. R. B., Mpunga, N. E., Machaga, S. J. and De Luca, D. W. 2005. The highland mangabey Lophocebus kipunji : A new species of African monkey. Science 308(5725): 1161–1164.
Kappeler, P. M., Rasoloarison, R. M., Razafimanantsoa, L., Walter, L. and Roos, C. 2005. Morphology, behavior and molecular evolution of giant mouse lemurs (Mirza spp.) Gray, 1870, with description of a new species. Primate Report (71): 3-26.
Landry, S. O. 2005. Untitled letter. Science 309(5744): 2164.
Mittermeier, R. A., Tattersall, I., Konstant, W. R., Meyers, D. M. and Mast, R. B. 1994. Lemurs of Madagascar Illustrations by Stephen D. Nash. Conservation International Tropical Field Guide Series, Conservation International, Washington, DC.
Polaszek, A., Grubb, P., Groves, C., Ehardt, C. L. and Butynski, T. M. 2005. Response. Science 309(5744): 2164-2165.
Sinha, A., Datta, A., Madhusudan, M. D. and Mishra, C. 2005. Macaca munzala: A new species from western Arunachal Pradesh, northeastern India. Int. J. Primatol. 26(4): 977-989.
Thalmann, U. and Geismann, T. 2000. Distribution and geographic variation in the western woolly lemur (Avahi occidentalis) with description of a new species (A. unicolor). Int. J. Primatol. 21: 915-941.
Thalmann, U. and Geismann, T. 2005. New species of woolly lemur Avahi (Primates, Lemuriformes) in Bemaraha (Central Western Madagascar). Am. J. Primatol. 67(3): 371-376.
Timm, R. M., Ramy II, R. R. and the Nomenclature Committee of the American Society of Mammalogists. 2005. What constitutes a proper description? Science 309(5744): 2163-2164.
Wakeham-Dawson, A., Morris, S. and Tubbs, P. 2002. Type specimens: Dead or alive? Bull. Zool. Nomenclat. 59: 282-284.

Suggested citation:

Rylands, A. B. and Mittermeier, R. A. 2006. New primates discovered in 2005. Unpublished report, IUCN/SSC Primate Specialist Group and Conservation International, Washington, DC.