Animal Diversity Web

Ge­o­graphic Range

Lep­ile­mur ed­wardsi is lim­ited to the west­ern trop­i­cal dry de­cid­u­ous forests of Mada­gas­car and has a range of less than 20,000 square kilo­me­ters (Mit­ter­meier et al., 2006; Andrainar­ivo et al., 2008). It is more specif­i­cally found from “north of the Bet­si­boka River to the Bay of Loza and the Mae­varano River, and in­clud­ing the Bon­golava Mas­sif” (Mit­ter­meier et al., 2006: 200). (Andrainar­ivo, et al., 2008; Mit­ter­meier, et al., 2006)

• Biogeographic Regions
• ethiopian
  • native

• Other Geographic Terms
• island endemic

Habi­tat

Lep­ile­mur ed­wardsi oc­cu­pies low­land dry de­cid­u­ous forests in west­ern Mada­gas­car (Mit­ter­meier et al., 2006). Though its avail­able habi­tat is highly frag­mented and de­creas­ing in quan­tity and qual­ity, Lep­ile­mur ed­wardsi is fre­quently en­coun­tered where found (Andrainar­ivo et al., 2008). It has an es­ti­mated den­sity of 60 in­di­vid­u­als/square kilo­me­ter within Ankarafantsika Na­tional Park (Mit­ter­meier et al., 2006). In­di­vid­u­als will use holes in dead or liv­ing trees as sleep­ing sites dur­ing the day (Ra­solo­har­i­jaona et al., 2006). (Andrainar­ivo, et al., 2008; Mit­ter­meier, et al., 2006; Ra­solo­har­i­jaona, et al., 2006)

• Habitat Regions
• tropical
• terrestrial

• Terrestrial Biomes
• forest

• Range elevation

  450 (high) m

  1476.38 (high) ft

Phys­i­cal De­scrip­tion

A mem­ber of the genus Lep­ile­mur and the fam­ily Lep­ile­muri­dae, Milne-Ed­wards' sportive lemurs are, like other species of sportive lemurs, con­sid­ered medium-sized ver­ti­cal clingers and leapers (Mit­ter­meier et al. 2006; Thal­mann 2002). A salta­tory species, L. ed­wardsi is one of the larger mem­bers of its genus and ranges from 54 to 58 cm in total length (in­clud­ing tail length) and 27 to 29 cm in head-body length; body weight ranges from ap­prox­i­mately 700 to 1000 g (Mit­ter­meier et al. 2006; War­ren & Cromp­ton 1997a). Milne-Ed­wards' sportive lemurs have white-tipped tails and gray-brown fur cov­er­ing their bod­ies and faces. The species has no­tice­ably siz­able ears and some­times has a dark stripe run­ning down the mid­dle of its back, with chest­nut-brown fur on the upper thighs, shoul­ders and fore­limbs. The front of its coat is gray with patches of cream col­ored fur (Mit­ter­meier et al. 2006). Males and fe­males are “sex­u­ally monomor­phic” and vary min­i­mally in body size, ex­cept dur­ing preg­nancy when the fe­male is con­sid­er­ably larger (Ra­solo­har­i­jaona et al., 2006; Ran­dri­anam­bin­ina et al., 2007). (Mit­ter­meier, et al., 2006; Ran­dri­anam­bin­ina, et al., 2007; Ra­solo­har­i­jaona, et al., 2006; Thal­mann, 2002; War­ren and Cromp­ton, 1997a)

• Other Physical Features
• endothermic
• bilateral symmetry

• Sexual Dimorphism
• sexes alike

• Range mass

  700 to 1000 g

  24.67 to 35.24 oz

• Range length

  54 to 58 cm

  21.26 to 22.83 in

Re­pro­duc­tion

Lep­ile­mur ed­wardsi in­di­vid­u­als form dis­persed pair-bonds that cor­re­late to ter­ri­to­ries, which in­clude sleep­ing sites and feed­ing sites. So­cial­ity in this species of lemur ap­pears very much re­lated to "sleep­ing pair-bonds" (Ra­solo­har­i­jaona et al., 2006). Pairs have been ob­served to re­strict them­selves to the usage of 1 to 4 sleep­ing sites ei­ther si­mul­ta­ne­ously or al­ter­na­tively (Ra­solo­har­i­jaona et al., 2003; Ra­solo­har­i­jaona et al., 2006). The species has been de­scribed as monog­a­mous and more gen­er­ally as pair-liv­ing (Ra­solo­har­i­jaona et al., 2003; Thal­mann, 2006). Though L. ed­wardsi is con­sid­ered spa­tially monog­a­mous, this does not pre­clude the po­ten­tial for extra pair breed­ing (Méndez-Cárde­nas & Zim­mer­mann, 2009). (Méndez-Cárde­nas and Zim­mer­mann, 2009; Ra­solo­har­i­jaona, et al., 2003; Ra­solo­har­i­jaona, et al., 2006; Thal­mann, 2006)

• Mating System
• monogamous

Lep­ile­mur ed­wardsi ex­hibits sea­sonal re­pro­duc­tion with a main mat­ing sea­son oc­cur­ring for about two months of the year be­tween May and June, cor­re­lat­ing with the be­gin­ning of this re­gion's dry sea­son in May. Fe­males go into es­trus for three months of the year, May through July, and males ex­hibit ap­par­ent sea­sonal vari­a­tion in the size of their testes (Ran­dri­anam­bin­ina et al., 2007). Neigh­bor­ing fe­males in dif­fer­ent home ranges may syn­chro­nize es­trus pe­ri­ods (Ra­solo­har­i­jaona et al., 2006). Male and fe­male body weight re­mains sim­i­lar dur­ing the mat­ing sea­son, al­though fe­males have a no­tice­ably higher body mass in Au­gust and No­vem­ber, most likely due to preg­nancy. The ob­served lit­ter size for Lep­ile­mur ed­wardsi is lim­ited to one young. Du­ra­tion of ges­ta­tion is 4 to 5 months, with off­spring typ­i­cally born in Oc­to­ber or No­vem­ber (Ran­dri­anam­bin­ina et al., 2007). Lac­ta­tion typ­i­cally be­gins at the same time as the rainy sea­son of the re­gion, which be­gins in No­vem­ber (Ran­dri­anam­bin­ina et al., 2007; Ra­solo­har­i­jaona et al., 2003). (Ran­dri­anam­bin­ina, et al., 2007; Ra­solo­har­i­jaona, et al., 2003; Ra­solo­har­i­jaona, et al., 2006)

• Key Reproductive Features
• iteroparous
• seasonal breeding
• gonochoric/gonochoristic/dioecious (sexes separate)
• sexual
• viviparous

• Breeding interval

  Milne-Edwards' sportive lemurs breed once yearly.

• Breeding season

  The breeding season occurs from May to June.

• Average number of offspring

  1

• Average gestation period

  4-5 months

• Range age at sexual or reproductive maturity (female)

  2 (low) years

• Range age at sexual or reproductive maturity (male)

  2 (low) years

There is lit­tle to no male parental in­vest­ment in Milne-Ed­wards' sportive lemurs (Ra­solo­har­i­jaona et al., 2000). For the first few days after birth, off­spring are left alone in sleep­ing trees (Ra­solo­har­i­jaona et al., 2003). Moth­ers later carry in­fants in their mouths dur­ing for­ag­ing ac­tiv­i­ties (Ran­dri­anam­bin­ina et al., 2007). They also park their in­fants while for­ag­ing (Thal­mann, 2003). (Ran­dri­anam­bin­ina, et al., 2007; Ra­solo­har­i­jaona, et al., 2003; Ra­solo­har­i­jaona, et al., 2000)

There has been one ob­served in­ci­dence of in­fan­ti­cide in this species in Ankarafantsika Na­tional Park. A mother was out for­ag­ing with her two in­fants and parked the younger one on a branch nearby. A strange, un­known male came and re­peat­edly at­tacked the younger in­fant. This oc­cur­rence of in­fan­ti­cide, how­ever, was most likely a con­se­quence of “in­ci­den­tal ag­gres­sion” as op­posed to an ac­tion out of com­pe­ti­tion for re­sources be­cause the male did not at­tack the other in­fant for­ag­ing nearby. This event sug­gests that as­so­ci­a­tion with males may ben­e­fit fe­males due to in­creased sur­vival of off­spring (Ra­solo­har­i­jaona et al., 2000; Ra­solo­har­i­jaona et al., 2003). (Ra­solo­har­i­jaona, et al., 2003; Ra­solo­har­i­jaona, et al., 2000)

• Parental Investment
• female parental care
• pre-fertilization
  • provisioning
  • protecting
    • female
• pre-hatching/birth
  • provisioning
    • female
  • protecting
    • female
• pre-weaning/fledging
  • provisioning
    • female
  • protecting
    • female
• pre-independence
  • provisioning
    • female
  • protecting
    • female

Lifes­pan/Longevity

In­for­ma­tion on the lifes­pan and longevity of Milne-Ed­wards' sportive lemurs is lack­ing (John­son, 2008). There are no known in­stances of suc­cess­ful breed­ing in cap­tiv­ity (Ran­dri­anam­bin­ina et al., 2007). (John­son, 2008; Ran­dri­anam­bin­ina, et al., 2007)

Be­hav­ior

A noc­tur­nal species, ac­tive pe­ri­ods for L. ed­wardsi cor­re­spond to tem­po­ral changes in the length of the day. Par­tic­u­lar ac­tiv­i­ties, in­clud­ing groom­ing, for­ag­ing, and trav­el­ing, vary in fre­quency dur­ing the night (War­ren & Cromp­ton, 1997a). They al­ter­nate pe­ri­ods of ac­tiv­ity with rest and are most ac­tive for the few hours im­me­di­ately fol­low­ing dusk (Mit­ter­meier et al. 2006). Fe­males and males form dis­persed sleep­ing-pairs and mem­bers of the same sleep­ing pair will share home ranges, in­clud­ing sleep­ing and feed­ing sites (Ra­solo­har­i­jaona et al., 2003; Ra­solo­har­i­jaona et al., 2006). How­ever, the num­ber of in­di­vid­u­als found in par­tic­u­lar sleep­ing sites can vary from a soli­tary in­di­vid­ual to an ad­di­tional two or three an­i­mals, and up to mul­ti­ple in­di­vid­u­als (Ra­solo­har­i­jaona et al., 2006; Thal­mann, 2002; Mit­ter­meier et al. 2006). Though they share sleep­ing holes, Milne-Ed­wards' sportive lemurs usu­ally for­age soli­tar­ily at night (Mit­ter­meier et al. 2006). They have been ob­served ac­tive mul­ti­ple times at the open­ing of their sleep hole dur­ing the day­time, yet will quickly re­turn at any sign of a threat (War­ren & Cromp­ton, 1997a). (Mit­ter­meier, et al., 2006; Ra­solo­har­i­jaona, et al., 2003; Ra­solo­har­i­jaona, et al., 2006; Thal­mann, 2002; War­ren and Cromp­ton, 1997a)

• Key Behaviors
• arboreal
• scansorial
• saltatorial
• nocturnal
• motile
• sedentary
• territorial
• social

• Average territory size

  10000 m^2

Home Range

The av­er­age home range of Milne-Ed­wards' sportive lemurs is ap­prox­i­mately 1 hectare for both males and fe­males. They fol­low an av­er­age nightly path of 343 m (Ra­solo­har­i­jaona et al., 2006; Ran­dri­anam­bin­ina et al., 2007). In­di­vid­u­als de­fend home ranges by vo­cal­iz­ing loudly and force­fully shak­ing the branches of trees (Mit­ter­meier et al. 2006). The de­fense of ter­ri­tory may be the re­sult of in­ter- and in­tra-sex­ual re­source com­pe­ti­tion (Ra­solo­har­i­jaona et al., 2003). (Mit­ter­meier, et al., 2006; Ran­dri­anam­bin­ina, et al., 2007; Ra­solo­har­i­jaona, et al., 2003; Ra­solo­har­i­jaona, et al., 2006)

Com­mu­ni­ca­tion and Per­cep­tion

Lep­ile­mur ed­wardsi lacks "spe­cial­ized glands” for mark­ing ter­ri­tory and there­fore does not use scent or urine to es­tab­lish ter­ri­to­ries. In­stead, these lemurs use loud calls to com­mu­ni­cate ter­ri­to­ri­al­ity. Calls occur both in the morn­ings and evenings and are sex-spe­cific. Nine types of calls have been iden­ti­fied with 3 used ex­clu­sively by fe­males, 5 used ex­clu­sively by males, and 1 used by both sexes. Most calls are re­lated to sleep­ing or feed­ing sites, and they are hy­poth­e­sized to be re­lated to mate de­fense and at­trac­tion as well as re­source de­fense. It is thought that loud vo­cal­iza­tions also serve in "reg­u­lat­ing spac­ing and co­he­sion” (Ra­solo­har­i­jaona et al., 2006: 598). (Ra­solo­har­i­jaona, et al., 2006)

Mem­bers of pairs "duet" with one an­other; both males and fe­males par­tic­i­pate equally in duet­ting, and this be­hav­ior is wit­nessed more fre­quently dur­ing the pe­riod of off­spring care and at feed­ing sites as op­posed to sleep­ing sites. Duet­ting causes the "syn­chro­niza­tion" of dif­fer­ent types of be­hav­ior, es­pe­cially move­ment. Be­cause this vocal syn­chro­niza­tion be­hav­ior most often oc­curs fol­low­ing birth of off­spring and when the fe­male is lac­tat­ing, Méndez-Cárde­nas and Zim­mer­mann (2009) pro­posed that duet­ting serves to lessen the risks of in­fan­ti­cide and to pro­tect a pair's ter­ri­tory, par­tic­u­larly their food re­sources. (Méndez-Cárde­nas and Zim­mer­mann, 2009)

• Communication Channels
• acoustic

• Other Communication Modes
• duets

• Perception Channels
• visual
• tactile
• acoustic
• chemical

Food Habits

In­di­vid­u­als of both sexes for­age soli­tar­ily at night, but a sleep­ing pair will search for food within the same range (Ra­solo­har­i­jaona et al., 2006; Thal­mann, 2001). This species is pri­mar­ily fo­liv­o­rous, how­ever, they will con­sume plump seeds, flow­ers, and fruits (Mit­ter­meier et al., 2006). They are rel­a­tively undis­crim­i­nat­ing and will se­lect from com­mon species of trees. Milne-Ed­wards' sportive lemurs have a larger food re­source base when com­pared to the sym­patric Avahi oc­ci­den­talis (Thal­mann, 2002). Though there is lit­tle over­lap be­tween these two species with re­gards to food re­sources, com­pe­ti­tion with Avahi oc­ci­den­talis may ex­plain the ten­dency of Lep­ile­mur ed­wardsi to eat leaves of poorer nu­tri­tive value in Ankarafantsika com­pared to other habi­tats in which it oc­curs (Thal­mann, 2006; Mit­ter­meier et al. 2006). (Mit­ter­meier, et al., 2006; Ra­solo­har­i­jaona, et al., 2006; Thal­mann, 2001; Thal­mann, 2002; Thal­mann, 2006)

• Primary Diet
• herbivore
  • folivore

• Plant Foods
• leaves
• seeds, grains, and nuts
• fruit
• flowers

Pre­da­tion

Fos­sas (Cryp­to­procta ferox) are main preda­tors of Lep­ile­mur ed­wardsi (Mit­ter­meier et al. 2006). While being pur­sued by a fossa, a Milne-Ed­wards' sportive lemur was ob­served “jump­ing rapidly from tree to tree into the vicin­ity of its sleep­ing site emit­ting loud bark call se­quences” (Scheumann et al., 2007: 110). The va­ri­ety of calls seemed to change and be­came more in­sis­tent in fre­quency and vol­ume when the fossa came close to cap­tur­ing L. ed­wardsi. Shriek­ing out of fear has been shown to draw other sportive lemurs in the area to gather and pro­duce alarm calls as well (Scheumann et al., 2007). (Mit­ter­meier, et al., 2006; Scheumann, et al., 2007)

Birds of prey, rep­tiles, such as boas, and other car­ni­vores (apart from fos­sas) prey on lemurs (Scheumann et al., 2007; Ra­solo­har­i­jaona et al., 2003). (Ra­solo­har­i­jaona, et al., 2003; Scheumann, et al., 2007)

• Anti-predator Adaptations
• cryptic

• Known Predators

  • fossas (Cryptoprocta ferox)
  • owls (Strigiformes)
  • diurnal raptors (Accipitriformes)
  • Madagascar tree boa (Sanzinia madagascariensis)
  • Dumeril's boa (Acrantophis dumerili)
  • Madagascar ground boa (Acrantophis madagascariensis)

Ecosys­tem Roles

In Ankarafantsika, Lep­ile­mur ed­wardsi is sym­patric with Avahi oc­ci­den­talis, Pro­p­ithe­cus ver­reauxi co­quereli, Eu­le­mur ful­vus, Eu­le­mur macaco, Mi­cro­ce­bus mur­i­nus, and Cheirogaleus medius, and may com­pete with Pro­p­ithe­cus ver­reauxi co­quereli, Eu­le­mur ful­vus, and Eu­le­mur macaco (War­ren & Cromp­ton, 1997b). Sportive lemurs serve as prey for nu­mer­ous taxa (Ra­solo­har­i­jaona et al., 2003). (Ra­solo­har­i­jaona, et al., 2003; War­ren and Cromp­ton, 1997b)

Eco­nomic Im­por­tance for Hu­mans: Pos­i­tive

Hu­mans will oc­ca­sion­ally con­sume Lep­ile­mur ed­wardsi (Mit­ter­meier et al., 2006). (Mit­ter­meier, et al., 2006)

• Positive Impacts
• food

Eco­nomic Im­por­tance for Hu­mans: Neg­a­tive

There are no known ad­verse ef­fects of Lep­ile­mur ed­wardsi on hu­mans.

Con­ser­va­tion Sta­tus

Habi­tat de­struc­tion due to clear­ing of for­est with fires for agri­cul­ture and live­stock graz­ing pas­ture, as well as the species' role as a food source for hu­mans, pose threats to Lep­ile­mur ed­wardsi (Andrainar­ivo et al., 2008; Mit­ter­meier et al., 2006; Ran­dri­anam­bin­ina et al., 2007). Its small num­ber of off­spring each year puts this species at higher risk (Ran­dri­anam­bin­ina et al., 2007). In ad­di­tion, the species does not ex­hibit an in­nate fear of new and po­ten­tially threat­en­ing or­gan­isms such as hu­mans even in areas with poach­ing (Ran­dri­anam­bin­ina et al., 2007). Ankarafantsika Na­tional Park is cur­rently the sin­gle pro­tected area in which Lep­ile­mur ed­wardsi is found, but they are also pre­sent in a 50,300 hectare pro­posed pro­tected area on the Bon­golava Mas­sif (Mit­ter­meier et al. 2006). (Andrainar­ivo, et al., 2008; Mit­ter­meier, et al., 2006; Ran­dri­anam­bin­ina, et al., 2007)

• IUCN Red List

  Vulnerable

• US Federal List

  Endangered

• CITES

  Appendix I

Other Com­ments

This species may be mis­taken for the sym­patric species of lemur, Avahi oc­ci­den­talis, which has a sim­i­lar stature (Mit­ter­meier et al. 2006; Thal­mann 2001). Milne-Ed­wards' sportive lemurs can be dif­fer­en­ti­ated by their “darker color, pointed face and more promi­nent ears” (Mit­ter­meier et al., 2006: 200). (Mit­ter­meier, et al., 2006; Thal­mann, 2001)

The num­ber of species rec­og­nized as mem­bers of the genus Lep­ile­mur and the clas­si­fi­ca­tion of this taxon has been de­bated. Ac­cord­ing to Louis, Jr. et al. (2006), eleven for­merly un­rec­og­nized sportive lemurs should be clas­si­fied in the same genus and fam­ily with L. ed­wardsi, which was most closely re­lated to Lep­ile­mur grew­cocki (Lep­ile­mur grew­cock­o­rum) in their analy­sis. A study by An­dri­a­holini­rina et al. (2006), on the other hand, rec­og­nized three new species, with L. ed­wardsi most closely re­lated to Lep­ile­mur mi­crodon. There is also dis­agree­ment in the lit­er­a­ture as to whether Lep­ile­mur ed­wardsi, and sportive lemurs in gen­eral, should be clas­si­fied as mem­bers of the fam­ily Lep­ile­muri­dae or as part of the fam­ily Lep­ile­muri­dae, which in­cludes the ex­tinct genus of giant lemur Mega­l­adapis (Mit­ter­meier et al., 2006). (An­dri­a­holini­rina, et al., 2006; Louis, Jr., et al., 2006; Mit­ter­meier, et al., 2006)

This species is also known as: Milne-Ed­wards’ weasel lemur, Lépile­mur de Milne-Ed­wards (French), Lemur Co­madreja De Ed­wards (Span­ish), Ed­wards Wiesel­maki (Ger­man), and Repa­haka/Boeng/Bo­engy/Kitronto Ki­tanta (Mala­gasy) (Mit­ter­meier et al., 2006; Andrainar­ivo et al., 2008). (Andrainar­ivo, et al., 2008; Mit­ter­meier, et al., 2006)

Con­trib­u­tors

Julia Os­ter­man (au­thor), Yale Uni­ver­sity, Eric Sar­gis (ed­i­tor), Yale Uni­ver­sity, Rachel Raci­cot (ed­i­tor), Yale Uni­ver­sity, Tanya Dewey (ed­i­tor), Uni­ver­sity of Michi­gan-Ann Arbor.

Ref­er­ences

Andrainar­ivo, C., V. An­dri­a­holini­rina, A. Feist­ner, T. Felix, J. Ganzhorn, N. Gar­butt, C. Golden, B. Kon­stant, E. Louis Jr., D. Mey­ers, R. Mit­ter­meier, A. Perieras, F. Princee, J. Rabariv­ola, B. Rako­to­sami­manana, H. Rasami­manana, J. Rat­sim­bazafy, G. Rav­eloari­noro, A. Razafi­manantsoa, Y. Rum­pler, C. Schwitzer, U. Thal­mann, L. Wilmé, P. Wright. 2008. "Lep­ile­mur ed­wardsi" (On-line). IUCN Red List of Threat­ened Species. Ac­cessed April 12, 2012 at http://​www.​iucnredlist.​org/​apps/​redlist/​details/​11617/​0.

An­dri­a­holini­rina, N., J. Fausser, C. Roos, I. Ravoa­ri­manana, J. Ganzhorn, B. Meier, R. Hil­gar­t­ner, L. Wal­ter, A. Zaramody, C. Langer, T. Hahn, E. Zim­mer­mann, U. Rade­spiel, M. Craul, J. Tomiuk, I. Tat­ter­sall, Y. Rum­pler. 2006. Mol­e­c­u­lar phy­logeny and tax­o­nomic re­vi­sion of the sportive lemurs (Lep­ile­mur, Pri­mates). BMC Evo­lu­tion­ary Bi­ol­ogy, 6/17. Ac­cessed May 01, 2012 at http://​www.​biomedcentral.​com/​1471-2148/​6/​17.

John­son, D. 2008. "The Life Spans of Non­hu­man Pri­mates" (On-line). Pri­mate Info Net. Ac­cessed May 01, 2012 at http://​pin.​primate.​wisc.​edu/​aboutp/​phys/​lifespan.​html.

Louis, Jr., E., S. En­g­berg, R. Lei, H. Geng, J. Som­mer, R. Ran­dria­mam­pi­onona, J. Ran­dria­manana, J. Za­onar­iv­elo, R. An­driantom­po­ha­vana, G. Ran­dria, Pros­per, B. Ra­maromi­lanto, G. Rako­toarisoa, A. Rooney, R. Bren­ne­man. 2006. Mol­e­c­u­lar and Mor­pho­log­i­cal Analy­ses of the Sportive Lemurs (Fam­ily Mega­l­adap­i­dae: Genus Lep­ile­mur) Re­veals 11 Pre­vi­ously Un­rec­og­nized Species. Spe­cial Pub­li­ca­tions: Mu­seum of Texas Tech Uni­ver­sity, 49: 1-46.

Mit­ter­meier, R., W. Kon­stant, F. Hawkins, E. Louis, O. Lan­grand, J. Rat­sim­bazafy, R. Ra­soloari­son, J. Ganzhorn, S. Ra­jao­belina, I. Tat­ter­sall, D. Mey­ers. 2006. Con­ser­va­tion In­ter­na­tional Trop­i­cal Field Guide Se­ries: Lemurs of Mada­gas­car. Wash­ing­ton, D.C.: Con­ser­va­tion In­ter­na­tional.

Méndez-Cárde­nas, M., E. Zim­mer­mann. 2009. Duet­ting- A Mech­a­nism to Strengthen Pair Bonds in a Dis­persed Pair-Liv­ing Pri­mate (Lep­ile­mur ed­wardsi)?. Amer­i­can Jour­nal of Phys­i­cal An­thro­pol­ogy, 139: 523-532.

Ran­dri­anam­bin­ina, B., S. Mbo­ti­zafy, S. Ra­solo­har­i­jaona, R. Ravoa­hangi­malala, E. Zim­mer­mann. 2007. Sea­son­al­ity in Re­pro­duc­tion of Lep­ile­mur ed­wardsi. In­ter­na­tional Jour­nal of Pri­ma­tol­ogy, 28: 783-790.

Ra­solo­har­i­jaona, S., B. Rako­to­sami­manana, B. Ran­dri­anam­bin­ina, E. Zim­mer­mann. 2003. Pair-spe­cific usage of sleep­ing sites and their im­pli­ca­tions for so­cial or­ga­ni­za­tion in a noc­tur­nal Mala­gasy pri­mate, the Milne Ed­wards' sportive lemur (Lep­ile­mur ed­wardsi). Amer­i­can Jour­nal of Phys­i­cal An­thro­pol­ogy, 122: 251-258.

Ra­solo­har­i­jaona, S., B. Rako­to­sami­manana, E. Zim­mer­mann. 2000. In­fan­ti­cide by a Male Milne-Ed­wards' Sportive Lemur (Lep­ile­mur ed­wardsi) in Ampi­joroa, NW-Mada­gas­car. In­ter­na­tional Jour­nal of Pri­ma­tol­ogy, 21/1: 41-45.

Ra­solo­har­i­jaona, S., B. Ran­dri­anam­bin­ina, P. Braune, E. Zim­mer­mann. 2006. Loud call­ing spac­ing and co­he­sive­ness in a noc­tur­nal pri­mate, the Milne-Ed­wards' sportive lemur. Amer­i­can Jour­nal of Phys­i­cal An­thro­pol­ogy, 129: 591-600.

Scheumann, M., A. Rabesan­dratana, E. Zim­mer­mann. 2007. Pre­da­tion, Com­mu­ni­ca­tion, and Cog­ni­tion in Lemurs. Pp. 100-126 in S Gursky, K Nekaris, eds. Pri­mate Anti-Preda­tor Strate­gies. On­line: Springer US. Ac­cessed April 11, 2012 at http://​dx.​doi.​org/​10.​1007/​978-0-387-34810-0_​5.

Thal­mann, U. 2002. Con­trasts Be­tween Two Noc­tur­nal Leaf-Eat­ing Lemurs. Evo­lu­tion­ary An­thro­pol­ogy, 11: 105-107.

Thal­mann, U. 2001. Food re­source char­ac­ter­is­tics in two noc­tur­nal lemurs with dif­fer­ent so­cial be­hav­ior: Avahi oc­ci­den­talis and Lep­ile­mur ed­wardsi. In­ter­na­tional Jour­nal of Pri­ma­tol­ogy, 22: 287-323.

Thal­mann, U. 2006. Be­hav­ioral and Eco­log­i­cal Adap­ta­tions in Two Small Fo­liv­o­rous Lemurs with Dif­fer­ent So­cial Or­ga­ni­za­tion: Avahi and Lep­ile­mur. Pp. 327-352 in L Gould, M Sauther, eds. Lemurs: Ecol­ogy and Adap­ta­tion. USA: Springer.

War­ren, R., R. Cromp­ton. 1997. A com­par­a­tive study of rang­ing be­hav­iour, ac­tiv­ity rhythms and so­cial­ity of Lep­ile­mur ed­wardsi (Pri­mates, Lep­ile­muri­dae) and Avahi oc­ci­den­talis (Pri­mates, In­dri­idae) at Ampi­joroa, Mada­gas­car. Jour­nal of Zo­ol­ogy, 243: 397-415.

War­ren, R., R. Cromp­ton. 1997. Lo­co­mo­tor ecol­ogy of Lep­ile­mur ed­wardsi and Avahi oc­ci­den­talis. Amer­i­can Jour­nal of Phys­i­cal An­thro­pol­ogy, 104: 471-486.