Lepilemur edwardsiMilne-Edwards's sportive lemur

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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)

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)

  • 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)

  • 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)

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)

  • 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)

  • 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)

  • Other Communication Modes
  • duets

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)

  • 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

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)

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.

Glossary

Ethiopian

living in sub-Saharan Africa (south of 30 degrees north) and Madagascar.

World Map

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

cryptic

having markings, coloration, shapes, or other features that cause an animal to be camouflaged in its natural environment; being difficult to see or otherwise detect.

duets

to jointly display, usually with sounds in a highly coordinated fashion, at the same time as one other individual of the same species, often a mate

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.

herbivore

An animal that eats mainly plants or parts of plants.

island endemic

animals that live only on an island or set of islands.

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).

monogamous

Having one mate at a time.

motile

having the capacity to move from one place to another.

native range

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

nocturnal

active during the night

saltatorial

specialized for leaping or bounding locomotion; jumps or hops.

seasonal breeding

breeding is confined to a particular season

sedentary

remains in the same area

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.

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.