Trachypithecus johniihooded leaf monkey

Ge­o­graphic Range

Tra­chyp­ithe­cus johnii (Nil­giri lan­gurs) are en­demic to the south­ern por­tions of the West­ern Ghats, a moun­tain range in the south of India. They are found be­tween 8°N and 12°N lat­i­tude and 76°E and 77.5°E lon­gi­tude. (Malviya, et al., 2011; Ram, 2007)

Habi­tat

Tra­chyp­ithe­cus johnii typ­i­cally in­hab­its a wide range of for­est habi­tats in the West­ern Ghats. They are found pri­mar­ily in sec­ondary moist de­cid­u­ous forests and wet ever­green to semi ever­green forests. They pre­fer lo­ca­tions that are as close to water, and as far away from hu­mans, as pos­si­ble.

Nil­giri lan­gurs are an ar­bo­real species, sleep­ing in the mid­dle or lower canopy in trees of medium height. They re­side in the sholas of the West­ern Ghats. Sholas are nar­row stretches of for­est sur­rounded by grass­lands, nes­tled in val­leys at high el­e­va­tions. Nil­giri lan­gurs live at el­e­va­tions be­tween around 300 and 2,500 me­ters, and are most com­monly found at ap­prox­i­mately 1,400 me­ters. (Malviya, et al., 2011; Poirier, 1969; Ram, 2007; Richard­son, 2005)

  • Range elevation
    300 to 2,500 m
    984.25 to ft
  • Average elevation
    1,400 m
    ft

Phys­i­cal De­scrip­tion

Tra­chyp­ithe­cus johnii is a colobine mon­key. They share many of the char­ac­ter­is­tic colobine fea­tures, such as a com­plex stom­ach, a re­duced thumb, and a long tail. Nil­giri lan­gurs have a shiny black coat with a red­dish-brown to gold head. New­borns are red­dish-brown for up to ten weeks, when they gain the hue of the adults.

Sim­i­lar to pur­ple faced lan­gurs, Nil­giri lan­gurs have dark faces and white side­burns. Fe­males have white patches on their thighs that dis­tin­guish them from males.

Nil­giri lan­gurs are sex­u­ally di­mor­phic, with males slightly larger than fe­males. The males of this species have a total body length that ranges be­tween 78 to 80 cm, weigh­ing be­tween 9.1 and 14.8 kg. The fe­males’ lengths range from 58 to 60 cm, weigh­ing be­tween 10.9 and 12 kg. The tails of both males and fe­males vary in length be­tween 68.5 and 96.5 cm. (Flea­gle, 1999; Malviya, et al., 2011)

  • Sexual Dimorphism
  • male larger
  • sexes colored or patterned differently
  • Range mass
    9.1 to 14.8 kg
    20.04 to 32.60 lb
  • Range length
    58 to 80 cm
    22.83 to 31.50 in

Re­pro­duc­tion

Tra­chyp­ithe­cus johnii in­di­vid­u­als are sex­u­ally di­mor­phic, with a male to fe­male body size ratio of 1.2 to 1.0. This in­di­cates mod­est in­tra­sex­ual com­pe­ti­tion among males for fe­male mates.

Though there is lit­tle doc­u­mented in­for­ma­tion about the Nil­giri lan­gur mat­ing sys­tem, it is as­sumed to be sim­i­lar to that of the Hanu­man lan­gur, to which it is closely re­lated. Hanu­man lan­gurs exist in uni-male or multi-male groups. The fre­quency of uni-male groups is de­pen­dent on the abil­ity of in­di­vid­ual males to de­fend and re­tain groups of fe­males. There are small num­bers of fe­males in these uni-male groups be­cause larger num­bers of fe­males above a cer­tain thresh­old are able to exert con­trol over the num­ber of males in their groups. Dis­tance be­tween groups, or pop­u­la­tion den­sity, also has an ef­fect on the pro­por­tion of uni-male to multi-male groups. When groups are more spread out, it is more ad­van­ta­geous for a male to re­main in his cur­rent group than to risk pre­da­tion while trav­el­ing alone to an­other group.

There is ev­i­dence that uni-male groups in­crease fe­male birth rates, in­di­cat­ing that less male-male com­pe­ti­tion is bet­ter for the re­pro­duc­tive suc­cess of fe­male lan­gurs. (Grueter and Van Schaik, 2009; Sri­vas­tava and Dun­bar, 1996)

There is a surge in birth rate for Nil­giri lan­gurs at two points in the year: most no­tably in the months of May and June, and sec­on­dar­ily be­tween Sep­tem­ber and No­vem­ber. This sea­son­al­ity of breed­ing may re­flect a reg­u­lar sea­sonal de­ple­tion of re­sources, which would af­fect fe­males' abil­ity to ovu­late and there­fore con­ceive. This would in­di­cate that Nil­giri lan­gurs breed year-round, but only con­ceive when re­sources are plen­ti­ful. Nil­giri lan­gurs typ­i­cally have only one off­spring at a time.

The length of the ges­ta­tion pe­riod of Nil­giri lan­gurs in the wild is not known, but is as­sumed to be com­pa­ra­ble to the closely re­lated Hanu­man lan­gur, the ges­ta­tion pe­riod for which is 200 days. In cap­tiv­ity, the ges­ta­tion pe­riod for Nil­giri lan­gurs is be­tween 140 and 220 days. At birth, in­fant Nil­giri lan­gurs weigh on av­er­age 0.5 kg. ("Nil­giri lan­gur", 2011; Koenig, et al., 1997; Malviya, et al., 2011; Richard­son, 2005)

Al­though no data exist for Nil­giri lan­gurs specif­i­cally, a re­cent study on the erup­tion of mo­lars in Tra­chyp­ithe­cus spec­i­mens in­di­cated that the erup­tion of the third molar oc­curred be­tween 3 and 5 years of age, in­di­cat­ing an in­di­vid­ual had reached sex­ual ma­tu­rity. (In­g­icco, et al., 2012)

  • Breeding interval
    Nilgiri langurs breed throughout the year, with two periods of increased breeding reflecting food availablity.
  • Breeding season
    Although limited data are available on potential breeding seasons of Nilgiri langurs, evidence suggests that mating takes place year-round.
  • Average number of offspring
    1
  • Range gestation period
    140 to 220 days
  • Average weaning age
    12 months
  • Average age at sexual or reproductive maturity (female)
    3-5 years
  • Average age at sexual or reproductive maturity (male)
    3-5 years

Tra­chyp­ithe­cus johnii does not dis­play strong bonds be­tween mother and young com­pared to sim­i­lar species, though nurs­ing lasts al­most a year. Oc­ca­sion­ally, when young in­fants are in dis­tress, their moth­ers will ig­nore the cries for help in an un­usual dis­play of ap­a­thy. One hy­poth­e­sis ex­plains this as a re­sult of the rel­a­tively low threat of preda­tors. In all other ways, how­ever, Nil­giri lan­gur moth­ers are at­ten­tive and nur­tur­ing. For ex­am­ple, off­spring cling to their moth­ers’ bel­lies while mov­ing and even jump­ing, and are also shel­tered by their moth­ers dur­ing rain­storms.

When new­borns are around 10 days old, their moth­ers will per­mit other fe­males to han­dle and care for their young. This oc­curs when the mother wants to feed. One non-ma­ter­nal fe­male could 'baby-sit' the off­spring of up to three moth­ers.

In­fants are weened by the end of one year, and at that point are al­most com­pletely in­de­pen­dent of their moth­ers. Until the end of their first year, Nil­giri lan­gur in­fants spend the ma­jor­ity of their time with their moth­ers or with other close fe­males, only rarely com­ing into con­tact with adult males. (Malviya, et al., 2011; Poirier, 1968)

  • Parental Investment
  • female parental care
  • pre-hatching/birth
    • provisioning
      • female
    • protecting
      • female
  • pre-weaning/fledging
    • provisioning
      • female
    • protecting
      • female
  • pre-independence
    • provisioning
      • female
    • protecting
      • male
      • female
  • post-independence association with parents

Lifes­pan/Longevity

There is no doc­u­mented in­for­ma­tion on the lifes­pan of Nil­giri lan­gurs in the wild, but they can live up to 29 years in cap­tiv­ity. ("Nil­giri lan­gur", 2011)

  • Range lifespan
    Status: captivity
    29 (high) years

Be­hav­ior

Tra­chyp­ithe­cus johnii is an ar­bo­real and di­ur­nal species. Dur­ing peak ac­tiv­ity, it en­gages al­ter­na­tively in feed­ing and rest­ing pe­ri­ods. There are a host of other so­cial ac­tiv­i­ties that are vis­i­ble, pri­mar­ily dur­ing rest­ing pe­ri­ods. Ex­am­ples of these are groom­ing, play­ing, chas­ing, watch­ing, fight­ing, mount­ing, ag­gres­sion, and dom­i­nance as­ser­tion.

Nil­giri lan­gurs gen­er­ally live in small fe­male-cen­tric groups. These groups can con­sist of one male and sev­eral fe­males, sev­eral males and sev­eral fe­males, or one or more soli­tary males. Typ­i­cally, the uni-male, multi-fe­male model is the most com­mon. Group sizes have been doc­u­mented be­tween 1 and 27 mem­bers, with an av­er­age of 14 or 15 in­di­vid­u­als. (Malviya, et al., 2011; Oates, et al., 1994; Poirier, 1970a)

Tra­chyp­ithe­cus johnii ex­hibits sub­tle dom­i­nance hi­er­ar­chies. There are two dom­i­nance struc­tures within each group, one for males and one for fe­males. Dom­i­nance struc­tures are not com­pletely rigid, in that dom­i­nance is ex­pressed in the num­ber of times a spe­cific in­di­vid­ual is forced to act sub­or­di­nately to an­other. For ex­am­ple, an alpha fe­male with higher sta­tus may act dom­i­nantly to­wards a beta fe­male the large ma­jor­ity of the time, but would be­have sub­or­di­nately on rare oc­ca­sions. How­ever, a gamma fe­male would usu­ally be­have sub­or­di­nately to­wards a beta fe­male, ex­press­ing dom­i­nance very in­fre­quently.

So­cial sta­tus is more im­por­tant for males than for fe­males. The alpha fe­male dis­plays dom­i­nance mainly in choos­ing pre­ferred feed­ing and sleep­ing sites. The alpha male dis­plays dom­i­nance most no­tice­ably in de­ter­min­ing the trav­el­ing di­rec­tion and tim­ing of the en­tire group, but also in free­dom of choice over daily life. The alpha male can eat with or so­cial­ize with any other mem­ber of the group.

Nil­giri lan­gur males are typ­i­cally more in­volved in dom­i­nance than fe­males, though dom­i­nance be­hav­iors in gen­eral are not par­tic­u­larly fre­quent. In ad­di­tion, ag­gres­sion dur­ing these en­coun­ters is avoided. There is lit­tle com­pe­ti­tion for food or mates, and sub­or­di­nate in­di­vid­u­als are care­ful to avoid po­ten­tially harm­ful sit­u­a­tions. Nil­giri lan­gurs there­fore have a rel­a­tively re­laxed daily life. (Poirier, 1970a)

Nil­giri lan­gurs ex­hibit ter­ri­to­r­ial be­hav­iors when con­fronted by other groups of their species. This de­fense of ter­ri­tory di­rectly in­volves only one adult male of each group. Males de­fend their home areas through phys­i­cal dis­plays, vo­cal­iza­tions, and chases. Con­fronta­tions some­times es­ca­late to phys­i­cal vi­o­lence, leav­ing both de­fend­ing males and non-group males with scars on their faces and bod­ies. In ex­treme cases, male take-overs and in­fan­ti­cide are known to occur. Such ter­ri­to­r­ial in­ter­ac­tions usu­ally take place in the areas where the home ranges of two or more groups over­lap, though Nil­giri lan­gurs usu­ally avoid these over­lap areas when the mem­bers of other groups are pre­sent. (Flea­gle, 1999; Malviya, et al., 2011)

  • Average territory size
    10,000 m^2

Home Range

Tra­chyp­ithe­cus johnii has an av­er­age home range of around 10,000 square me­ters. The size of the home range fluc­tu­ates with qual­ity of habi­tat dur­ing the sea­sons: when pre­ferred food is read­ily avail­able, the range is smaller; when it is more scarce, the home range in­creases. The core area of this species in­cludes feed­ing sites and rest­ing or sleep­ing sites. These sites are cho­sen as close to water and as far away from hu­mans as pos­si­ble.

An ob­ser­va­tional study found that the ter­ri­tory size is es­sen­tially equiv­a­lent to the home range. (Hor­wich, 1972; Malviya, et al., 2011)

Com­mu­ni­ca­tion and Per­cep­tion

Nil­giri lan­gurs com­mu­ni­cate mainly through vo­cal­iza­tion. Vo­cal­iza­tions have many dif­fer­ent func­tions, with var­i­ous calls re­lated to dif­fer­ent so­cial sit­u­a­tions. Vocal com­mu­ni­ca­tions are ob­served dur­ing the main­te­nance of the so­cial hi­er­ar­chy, dur­ing ter­ri­to­r­ial dis­putes, as a part of within-group fe­male-fe­male con­flicts, and fi­nally for male in­ter­ven­tion into such con­flicts.

Nil­giri lan­gurs also com­mu­ni­cate with ol­fac­tory, tac­tile, and vi­sual sig­nals. They are known to en­gage in mouth sniff­ing as a part of feed­ing. This species also ex­hibits nu­mer­ous ex­pres­sions to sig­nal to other species mem­bers. Some ex­am­ples of ex­pres­sions in­clude a threat­en­ing open mouth ges­ture, a sub­mis­sive head shak­ing ges­ture, a friendly play in­vi­ta­tion, and a com­fort­ing out­stretched hand ges­ture di­rected from moth­ers to­wards in­fants. In­stances of tac­tile com­mu­ni­ca­tion in­volve bit­ing, lick­ing, em­brac­ing, stroking, and slap­ping. (Malviya, et al., 2011)

Food Habits

A fo­liv­o­rous species, Nil­giri lan­gurs eat many kinds of leaves, which make up the ma­jor­ity of their diet. They also take in nu­tri­ents from fruits, seeds, nuts, stems, bark, flow­ers, and soil. The only car­niv­o­rous as­pect of their diet is the oc­ca­sional con­sump­tion of in­sects.

Nil­giri lan­gurs pre­fer the leaves of teak trees, Tec­tona gran­dis, and con­sume these larger leaves more fre­quently than oth­ers that oc­ca­sion­ally make up their diet.

Nil­giri lan­gurs are re­ported to choose foods based on di­gestibil­ity; they have a diet that is low in both fiber and veg­etable tan­nin. It is hy­poth­e­sized that they use the soil they con­sume as an antacid, which sta­bi­lizes the pH level of the stom­ach. (Hor­wich, 1972; Malviya, et al., 2011; Oates, et al., 1980)

Nil­giri lan­gurs spend al­most half of their wak­ing hours feed­ing. When eat­ing leaves, they first eat the tips, then they rip off the sides to ex­pose the midrib, which they also con­sume. This is a del­i­cate and de­lib­er­ate process, with one side of each leaf pealed back at a time to ex­pose the midrib of the leaf. Each leaf takes ap­prox­i­mately 30 sec­onds to eat. Dur­ing eat­ing pe­ri­ods, in which Nil­giri lan­gurs take and con­sume leaves al­most con­tin­u­ously, about 5-10 sec­onds are taken be­tween leaves. (Hor­wich, 1972; Malviya, et al., 2011)

  • Animal Foods
  • insects
  • Plant Foods
  • leaves
  • wood, bark, or stems
  • seeds, grains, and nuts
  • fruit
  • flowers

Pre­da­tion

Nil­giri lan­gurs have vo­cal­iza­tions for dis­tress, warn­ing, and alarm, each with its own sound and sig­nif­i­cance. They emit high-pitched barks as alarm calls to nearby in­di­vid­u­als, which can be ei­ther short in quick in­ter­vals or longer and con­tin­u­ous. The con­tin­u­ous alarm call usu­ally rep­re­sents a more press­ing haz­ard. In re­sponse to hear­ing ei­ther of the bark­ing alarm calls, Nil­giri lan­gurs will dis­play signs of fear often ac­com­pa­nied by flight.

Nil­giri lan­gurs are rel­a­tively free from pre­da­tion, al­though one doc­u­mented preda­tor is the In­dian wild dog. Hu­mans also hunt these lan­gurs. (Poirier, 1968; Poirier, 1970b; Singh, et al., 2008)

Ecosys­tem Roles

Of all the pri­mates en­demic to the West­ern Ghats in India, Nil­giri lan­gurs feed on the largest num­ber of plant species. They con­sume over 115 dif­fer­ent species, in­clud­ing at least 58 tree species, 6 shrubs, 13 non-woody plants, 32 vines, and 6 par­a­sitic plants. Fruits, nuts, and seeds are also parts of their di­verse diets, and they func­tion in their ecosys­tem as seed dis­persers. (Malviya, et al., 2011)

  • Ecosystem Impact
  • disperses seeds

Eco­nomic Im­por­tance for Hu­mans: Pos­i­tive

Tra­chyp­ithe­cus johnii in­di­vid­u­als are poached for var­i­ous pur­poses. Their body parts are sold for use in tra­di­tional med­i­cine and are even ad­ver­tised as com­po­nents of recipes for aphro­disi­acs. In ad­di­tion, they are hunted for their skins, which are used as drum heads. Nil­giri lan­gurs are also cap­tured and sold as pets. While these may be pos­i­tive eco­nomic ef­fects for hu­mans, they have dev­as­tat­ing neg­a­tive con­se­quences for the pop­u­la­tions of Nil­giri lan­gurs that re­main in de­cline. (Malviya, et al., 2011; Singh, et al., 2008)

Eco­nomic Im­por­tance for Hu­mans: Neg­a­tive

Nil­giri lan­gurs are known as crop pests to cau­li­flower, potato, car­damom, and gar­den poppy farm­ers. How­ever, this neg­a­tive ef­fect on hu­mans may be ex­ac­er­bated by the fact that peo­ple are con­stantly en­croach­ing on the nat­ural habi­tat of Nil­giri lan­gurs by plant­ing crops, min­ing, and build­ing homes. (Malviya, et al., 2011; Singh, et al., 2008)

  • Negative Impacts
  • crop pest

Con­ser­va­tion Sta­tus

Only 50% of this species’ ter­ri­tory is within the pro­tected area in the West­ern Ghats. Their pop­u­la­tion is es­ti­mated be­tween 5,000 and 15,000 in­di­vid­u­als and less than 10,000 ma­ture in­di­vid­u­als. The two main threats to Nil­giri lan­gurs in­clude the de­struc­tion of their nat­ural habi­tat and poach­ing for var­i­ous pur­poses.

Be­cause of the rel­a­tively small num­ber of ma­ture adult Nil­giri lan­gurs in the world, and be­cause of the de­clin­ing trend in pop­u­la­tion size, they are placed on the IUCN Red List as Vul­ner­a­ble. In ad­di­tion, they have been pro­tected under the In­dian Wildlife Pro­tec­tion Act since 1972. (Malviya, et al., 2011; Singh, et al., 2008)

There is ev­i­dence that the level of human dis­tur­bance, or habi­tat sat­u­ra­tion, cor­re­lates pos­i­tively with in­stances of in­fan­ti­cide in lan­gurs. In ef­fect, human dis­tur­bance low­ers in­stances of fe­male dis­per­sal rel­a­tive to male takeover. When males in­vade and take over a group, lev­els of in­fan­ti­cide are much higher than if fe­males are able to more peace­fully leave their pre­vi­ous groups, which only hap­pens when lan­gurs have not reached the hold­ing ca­pac­ity of their en­vi­ron­ment. This shows the neg­a­tive ef­fects of hu­mans en­croach­ing on the lan­gurs' ter­ri­tory. (Sterck, 1998)

Other Com­ments

There is some de­bate over the clas­si­fi­ca­tion of this species, giv­ing rise to con­flict­ing views on its tax­o­nomic place­ment. Some tax­on­o­mists place Nil­giri lan­gurs with Hanu­man lan­gurs, while oth­ers con­sider them a species of Tra­chyp­ithe­cus. Still oth­ers place them in their own genus known as Kasi. Nil­giri lan­gurs are there­fore named Tra­chyp­ithe­cus johnii, Semno­p­ithe­cus johnii, Pres­bytis johnii, and Kasi johnii in var­i­ous sci­en­tific lit­er­a­ture.

This species' var­i­ous com­mon names in­clude: black leaf mon­key, hooded leaf mon­key, In­dian hooded leaf mon­key, John's lan­gur, Nil­giri black lan­gur, and Nil­giri leaf mon­key. (Flea­gle, 1999; Richard­son, 2005)

Con­trib­u­tors

Claire Solomon (au­thor), Yale Uni­ver­sity, Eric Sar­gis (ed­i­tor), Yale Uni­ver­sity.

Glossary

acoustic

uses sound to communicate

arboreal

Referring to an animal that lives in trees; tree-climbing.

bilateral symmetry

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.

chemical

uses smells or other chemicals to communicate

detritus

particles of organic material from dead and decomposing organisms. Detritus is the result of the activity of decomposers (organisms that decompose organic material).

diurnal
  1. active during the day, 2. lasting for one day.
dominance hierarchies

ranking system or pecking order among members of a long-term social group, where dominance status affects access to resources or mates

drug

a substance used for the diagnosis, cure, mitigation, treatment, or prevention of disease

endothermic

animals that use metabolically generated heat to regulate body temperature independently of ambient temperature. Endothermy is a synapomorphy of the Mammalia, although it may have arisen in a (now extinct) synapsid ancestor; the fossil record does not distinguish these possibilities. Convergent in birds.

female parental care

parental care is carried out by females

folivore

an animal that mainly eats leaves.

food

A substance that provides both nutrients and energy to a living thing.

forest

forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.

frugivore

an animal that mainly eats fruit

granivore

an animal that mainly eats seeds

herbivore

An animal that eats mainly plants or parts of plants.

iteroparous

offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).

motile

having the capacity to move from one place to another.

mountains

This terrestrial biome includes summits of high mountains, either without vegetation or covered by low, tundra-like vegetation.

native range

the area in which the animal is naturally found, the region in which it is endemic.

nomadic

generally wanders from place to place, usually within a well-defined range.

oriental

found in the oriental region of the world. In other words, India and southeast Asia.

World Map

pet trade

the business of buying and selling animals for people to keep in their homes as pets.

polygynandrous

the kind of polygamy in which a female pairs with several males, each of which also pairs with several different females.

polygynous

having more than one female as a mate at one time

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

social

associates with others of its species; forms social groups.

tactile

uses touch to communicate

terrestrial

Living on the ground.

territorial

defends an area within the home range, occupied by a single animals or group of animals of the same species and held through overt defense, display, or advertisement

tropical

the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.

visual

uses sight to communicate

viviparous

reproduction in which fertilization and development take place within the female body and the developing embryo derives nourishment from the female.

year-round breeding

breeding takes place throughout the year

Ref­er­ences

2011. "Nil­giri lan­gur" (On-line). Arig­nar Anna Zo­o­log­i­cal Park - Nil­giri lan­gur. Ac­cessed April 28, 2013 at http://​www.​aazoopark.​in/​Nilgiri-Langur.​php.

Flea­gle, J. 1999. Pri­mate Adap­ta­tion and Evo­lu­tion. San Diego: Aca­d­e­mic Press; An Im­print of El­se­vier.

Grueter, C., C. Van Schaik. 2009. Sex­ual Size Di­mor­phism in Asian Colobines Re­vis­ited. Amer­i­can Jour­nal of Pri­ma­tol­ogy, Vol­ume 71, Issue 7: 609-616. Ac­cessed April 23, 2013 at http://​onlinelibrary.​wiley.​com/​doi/​10.​1002/​ajp.​20695/​abstract.

Hor­wich, R. 1972. Home Range and Food Habits of the Nil­giri Lan­gur, Pres­bytis johnii. Jour­nal of the Bom­bay Nat­ural His­tory So­ci­ety, Vol­ume 69, Issue 2: 255-267. Ac­cessed April 06, 2013 at https://​docs.​google.​com/​viewer?​a=v&​q=cache:​4czjuOlrqTEJ:​www.​communityconservation.​org/​publications/​jbnhs69.​2..​pdf+Presbytis+johnii&​hl=en&​gl=us&​pid=bl&​srcid=ADGEESjbZUp-U6S8ISn6YU2jbcx340kuQ3WYMCPJWuyP4oGOX1-unBo-hl1qCZYLBsFk1tcaYDoUittWxqeLzPTIlsSZFQI4FnsacDtnf6Y270HuL1iXJvWI9ijLORviYz3V8w3SlUbn&​sig=AHIEtbQK4SakPGiB_​lxSRWT7HPgAesiTkw.

In­g­icco, T., A. Moigne, D. Gom­mery. 2012. A de­cid­u­ous and per­ma­nent den­tal wear stage sys­tem for as­sess­ing the age of Tra­chyp­ithe­cus sp. spec­i­mens (Colobi­nae, Pri­mates). Jour­nal of Ar­chae­o­log­i­cal Sci­ence, Vol­ume 39, Issue 2: 421-427. Ac­cessed April 27, 2013 at http://​www.​sciencedirect.​com/​science/​article/​pii/​S0305440311003578.

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