Lesser bushbaby

SOCIAL ORGANIZATION AND BEHAVIOR

he social fabric of the lives of bushbabies is as varied as their habitats and for many species, incompletely known. However, the social systems of the bushbabies are best described as non-gregarious (Pullen et al. 2000). For example, G. moholi spends 70% of its activity period solitary (Bearder & Doyle 1974b; Doyle & Bearder 1977). This is not a rule however, with some populations of G. alleni being found about half of the time in association with one or several conspecifics while other populations of the same species do not show this pattern (Ambrose 2003).

The lives of bushbabies are typified by a dichotomy of daily activity; divided between solitary evening and nighttime activities alternating with morning and daytime social aggregation, especially at the sleeping site (Charles-Dominique 1972; Bearder & Martin 1980; Harcourt & Nash 1986). Individuals are not often seen together at night (Harcourt & Nash 1986). Because some of the activity period is spent solitary, social groupings are perhaps best quantified through sleeping group size, which can vary between solitary sleeping up to ten individuals per sleeping site (data compiled from the literature by Bearder 1987 & Bearder et al. 2003). Only G. demidoff does not sometimes sleep alone and has a minimum sleeping group size of two individuals (Bearder et al. 2003). Home ranges in G. zanzibaricus males show only slight overlap, as is mostly the case in females of comparable age; however females sometimes share ranges with other females who are likely kin (Nash 1984; Harcourt & Nash 1986). Males will sleep with each individual whose range overlaps his, mostly one or several adult females and immatures per night (Harcourt & Nash 1986). Adult male G. moholi never sleep together (Bearder & Martin 1980). The overlap in male ranges might be explained by low sample size, tolerance of smaller emigrating males, or the process of replacement of a resident male. Adults of G. zanzibaricus likely defend their ranges (Harcourt & Nash 1986). G. moholi females range structure is related to their age, with like-aged females showing little range overlap while those with somewhat larger age differences may overlap extensively (Bearder & Martin 1980). In both G. alleni and G. demidovii, female ranges vary in their overlap with one another and the ranges of males usually overlap at least one female range. G. alleni male home ranges are vast and may overlap the home ranges of more than 8 different females. Male ranges either do not overlap or overlap to an insignificant degree. Small areas of male overlap are sometimes deemed common areas, and may be shared by several males in G. demidovii and G. alleni (Charles-Dominique 1977a).

In G. moholi, males exhibit a linear, age-based dominance hierarchy, with only the highest ranking males defending their territories. Such individuals are usually some of the largest present in a given area and initiate agonistic contact, often resulting in the displacement of subordinate males. Regardless of rank, adult males are not tolerant of one another (Bearder & Martin 1980). There are two classes of mature G. moholi males, dominant heavier males who interact with females more often and submissive, nonterritorial adult males who weigh less and interact with females less (Bearder 1987). It is suggested that there are four classes of adult male in G. demidovii, each with particularities of home range extent and overlap relating to contacts with both female home ranges as well as with other adult males (Charles-Dominique 1977a). To show submission, individuals descend to ground-level (Bearder & Doyle 1974b; Bearder & Martin 1980). Adult male G. alleni are extremely intolerant of one another and are in pronounced competition with other males (Charles-Dominique 1977).

Female G. moholi exhibit more ambiguous social relationships, although agonistic territorial behavior is sometimes seen at the borders of home ranges (Bearder & Martin 1980). Affiliative interactions in G. senegalensis consist of urine-washing, grappling, chasing, play and grooming (Bearder & Doyle 1974b).

Females in G. zanzibaricus tend not to emigrate from their natal range while males do, usually at puberty (Nash 1984; Harcourt & Nash 1986). In some species, young pubescent males may spend time in a so-called “vagabond” phase of life, in which they are nomadic and do not spend much time in a given area (Charles-Dominique 1977a). Emigration in G. moholi is rigidly defined, with the pre-reproductive pubescent male traveling either unidirectionally either east or west over several nights, ending up several kilometers from the natal range (Bearder 1987).

In captivity, aggression is most common between members of the same sex while affinitive behaviors are mostly seen between the sexes in G. senegalensis (Nash & Flinn 1978). In the wild, confrontations are usually avoided, but rare violence can result in serious injury and/or the displacement of a formerly dominant male by a vagabond male (Charles-Dominique 1977a).

When G. moholi encounters a conspecific they will greet each other with nose-to-nose contact and smelling, which may be followed by grooming or agonistic behavior, especially when the individuals are unfamiliar with one another (Doyle 1974).

REPRODUCTION

The mating system of bushbabies (G. moholi) is best described as dispersed and not strictly polygynous. Certain males have a better chance of mating success than others but females may still mate with more than one male (Pullen et al. 2000). Female bushbabies exhibit estrus swelling of the sex skin and the vagina is closed at all other times other than estrus (Nash 1983; Zimmermann 1989; Lipschitz et al. 2001). Estrus and the mating period lasts 1-3 days with some evidence from the wild suggesting that females do not all come into estrus at the same time (Gucwinska & Gucwinski 1968; Doyle et al. 1971; Pullen et al. 2000). G. moholi copulations in the wild last, on average, 9 minutes, but can range from 2 to 53 minutes. Copulation is often serial, with 2-5 prolonged mounts punctuated with rest and grooming (Pullen et al. 2000). Copulations between captive G. senegalensis are similarly prolonged (LT Nash, per. comm.).

In both the wild and captivity, females may mate with more than one male during a single estrus (Gucwinska & Gucwinski 1968; Pullen et al. 2000). In G. moholi, larger males have better mating success (Pullen et al. 2000). The copulation posture of bushbabies is dorso-ventral, with the male grasping the heels of the female from behind (Charles-Dominique 1977a; Lipschitz 1996b). G. demidovii copulates while suspended from a branch, while other species do not practice suspensory copulation (Charles-Dominique 1977a). The typical mating sequence in captivity of G. moholi consists of the male approaching the female, chasing her, grabbing her and mounting (Lipschitz 1996a). Nose-to-nose touching and genital sniffing may also precede copulation (Lipschitz 1996b). In the wild, all matings in this species are initiated by very persistent males and females are generally averse to their mating attempts (Pullen et al. 2000). In captivity however, females sometimes approach present her hindquarters to a male during behavioral estrus (Lipschitz 1996b).

Several species of bushbaby have two mating and birth seasons per year in the wild, including G. senegalensis, G. zanzibaricus and G. moholi, while other bushbabies of comparable size (G. gallarum) likely follow this pattern as well (Haddow & Ellice 1964; Butler 1967; Harcourt 1986a; Pullen et al. 2000; Butynski & de Jong 2004). Individual females in some species are capable of breeding twice in the same year (Harcourt 1986a; Bearder et al. 2003). Just prior to and during mating seasons, the weight of males and the volume of their testes increases (Pullen et al. 2000).

Among the bushbabies, gestation lengths can range from around 111 days to around 142 with smaller species generally having shorter gestation lengths (Gucwinska & Gucwinski 1968; Doyle et al. 1971; Bearder & Doyle 1974a; Charles-Dominique 1977a; Izard & Simons 1986; Lipschitz 1996a; Izard & Nash 1988; Nash et al. 1989; Zimmermann 1989; Nekaris & Bearder 2007). The average ovarian cycle in captive G. moholi is 38.5 days (Lipschitz 1996b). Sexual maturity is reached across bushbaby species with available data between 8 and 18 months of age (Charles-Dominique 1977a; Izard & Nash 1988; Nash 1993; data compiled by Nekaris & Bearder 2007).

Per pregnancy in most bushbaby species, usually one infant is born, with the possibility of twins and extremely rarely, triplets (Butler 1967; Doyle et al. 1971; Riordan 1971; Nash 1983; Harcourt 1986a; Izard & Simons 1986; Nash et al. 1989; Zimmermann 1989; Bearder et al. 2003). There are several species that are exceptions however. One is G. moholi, which has higher rates of twinning than most other bushbabies (Izard & Simons 1986; Izard & Nash 1988; Nash et al. 1989; Bearder et al. 2003). Bushbaby mothers can give birth to up to 4 infants per year (G. moholi) (Bearder et al. 2003). Estrus is sometimes seen for several days directly following birth and conceptions are possible during this postpartum estrus (Doyle et al. 1969; Zimmermann 1989; Nash 2003).

PARENTAL CARE

As the birth of the infant nears, G. zanzibaricus females start sleeping on their own and isolate themselves whereas before they sleep with conspecifics (Charles-Dominique 1977a; Harcourt 1986a). Births occur in nests or the hollows of trees (Bearder 1987). Pooled, multi-species data give the birth weight at about 5 to 24 g (0.2 to 0.8 oz) (Gucwinska & Gucwinski 1968; Doyle et al. 1971; Izard & Nash 1988; data compiled by Nekaris & Bearder 2007 and Zimmermann 1989 for older data). The eyes are open at birth and the pelage is ubiquitously grey and meager (Gucwinska & Gucwinski 1968; Charles-Dominique 1977a; Doyle 1979; Zimmermann 1989). Thick fur comes in between two and three weeks old (Charles-Dominique 1977a).

In captivity (G. moholi), at one day old, infants are capable of clinging to a branch and within the first several days of life, can walk quadrupedally. By two weeks of age, infants can quadrupedally run (Doyle 1979). Infants start following their mothers about between 4 and 6 weeks old (Doyle & Bearder 1977). Play begins very early, within the first week of age (Doyle 1979). Among juvenile G. senegalensis, young males play more often than females (Nash 1993).

While the mothers are engaging in other activities, such as foraging nearby, juveniles are parked, usually in tree forks or tangles, motionlessly hiding from potential threats (Doyle 1974; Charles-Dominique 1977a; Harcourt 1986a; Bearder 1987; Ambrose 2003). They sometimes remain parked for prolonged periods of time, up to three hours, but if the parking is prolonged, the mother will occasionally visit (Doyle 1974; Doyle & Bearder 1977). If directly disturbed or seemingly forgotten, the infant may emit a distress call and the mother comes running (Doyle et al. 1969; Charles-Dominique 1977a).

In G. senegalensis, up until 12 weeks of age in captivity, adult-type vocalizations are not heard, with infants instead producing characteristic “zek” calls (Zimmermann 1989). G. moholi infants utter three distinct types of distress vocalization which are given in increasing situations of distress to which the mother responds by coming to the infants and grooming them (Mascagni & Doyle 1993). In situations of danger, the mother will carry the infant away in her mouth (Charles-Dominique 1977a). In captivity, infants first emerge from their nest box at ten to 14 days of age, and in the wild starting between 3 and 7 days of age, the mother carries the infant out of the nest with her and starts parking it (Gucwinska & Gucwinski 1968; Doyle et al. 1969; Charles-Dominique 1977a). In the wild, independent movement and exploration begins about 2-3 weeks old (Charles-Dominique 1977a). Consumption of solid foods commences at one month of age, when infants start stealing food from the mouths of their mothers (Charles-Dominique 1977a). In the wild, juveniles usually forage alone (Bearder 1987).

The typical maternal infant carrying posture is in the mouth, often by the nape (Gucwinska & Gucwinski 1968; Doyle et al. 1969; Ward & Scott 1970; Doyle & Bearder 1977; Doyle 1979; Harcourt 1986a; Ambrose 2003; Bearder 2003; Nash 2003). Carrying with the infant clinging to the fur of the mother is rare, but has been observed in G. gallarum (Butynski & de Jong 2004). In captivity, all sexes and ages are patient with juveniles, and sometimes play, groom, and show interest towards them (Bearder & Doyle 1972). In semi-natural captive conditions, grooming frequencies between mother and offspring do not change as the infant ages, persisting until full adult separation from the mother (Doyle et al. 1969).

In captivity, both G. moholi and G. senegalensis females lactate for an average of around 100 days after the birth of their infant with weaning at 10-14 weeks of age (Doyle 1979; Izard 1987; Zimmermann 1989). However, wild G. demidoff weans its young earlier, at around 45 days old (Charles-Dominique 1977a). Females sometimes nurse infants not belonging to them and there are not profound differences in contact and nursing between sexes of infants (Nash 2003). By ten months of age, males have reached puberty and emigrate and also around this time, courting of females starts (Charles-Dominique 1977a; Bearder 1987). Females in G. moholi start showing territorial behaviors around 200 days old (Bearder 1987).

COMMUNICATION

The vocalizations of bushbabies have been roughly divided into discrete types by function. These include social cohesion and spacing calls (social contact calls), agonistic calls (threat and distress calls), and attention and alarm calls (Charles-Dominique 1977a; Zimmermann et al. 1988). Among G. moholi and G. senegalensis, there appear to be 14 types of call in common between the two species (Zimmermann et al. 1988). However, different species vary in the total numbers of vocalizations they produce. Bushbaby vocalizations are extremely variable, often grading into one another and are produced both during inhalation as well as during exhalation (Bearder et al. 1995).

Because morphological differences are not always useful for differentiation, vocalizations are considered a good way to tell bushbaby species from one another (Ambrose 2003). This is particularly true of advertising calls, which are often unique to species in primates and in bushbabies these types of calls are the most diagnostic (Zimmermann et al. 1988; Zimmermann 1990; 1995). Loud advertisement calls are often profoundly different than those of other bushbabies and are used to tell different species apart (for example Harcourt & Bearder 1989; Butynski et al. 1998; Wickings et al. 1998; Anderson et al. 2000; Perkin et al. 2002; Butynski & de Jong 2004; Ambrose 2006; Butynski et al. 2006). Vocalizations and olfactory marking are likely the best signals functioning in intra-species individual recognition by bushbabies (Ambrose 2003). Vocalizations can be so unique that bushbabies which are not distinguishable otherwise have been suggested as potential subspecies or even full species based mainly on differences in vocal structure (Perkin et al. 2002; Ambrose 2003). It is less likely that species of bushbaby can be be differentiated based on agonistic, attention, and alarm calls (Zimmermann 1990). Advertisement calls are usually heard upon emergence from the sleeping site, reconvening before sleep at the end of the activity period, and for maintaining contact during the night (Bearder et al. 1995). Further, advertisement calls might also be given in calling bouts between members of the same sex (Bearder et al. 1995).

While usually considered non-gregarious, G. moholi emits alarm calls when predators are encountered and the alarm calling is contagious, with other bushbabies joining in and coming together, mobbing and calling for up to 30 minutes around the threat (Bearder et al. 2002).

Call frequency is affected by population density, with some species calling more over the course of the nightly activity period if more conspecifics are nearby (Courtenay & Bearder 1989).

In the wild, urine-washing is a means by which G. alleni may disperse olfactory clues throughout its home range. In this species, the soles of the feet are washed with urine, which is subsequently dispersed, especially in areas of overlap with the home ranges of conspecifics (Charles-Dominique 1977b). Urine-washing may also communicate social clues to other bushbabies (Nash 1993). There are several types of scent-marking observed in captive G. demidoff. These include urine-washing, hand-rubbing, genital-planting, cheek-/chin-rubbing, chest-rubbing, anogenital-rubbing, and substrate-biting combined with flehmen (Pitts 1988). However, urine washing may also enhance grip in some species, e.g. G. moholi (Harcourt 1981).

Displays seen in G. demidoff include defensive displays seen in both sexes, defensive anti-intruder/anti-predator displays, male-male rank demonstration displays, and dominant-female self-advertising displays. Further, it is suggested that the displays given by dominant-males in offensive situations may be species specific and are a useful tool in differentiating species of bushbaby (Pitts 1988). In the wild, aggressive postures consist of the bushbaby extending its body and tail, spreading the ears and opening the mouth (Charles-Dominique 1977a).

Content last modified: December 8, 2008

Written by Kurt Gron. Reviewed by Leanne Nash.

Cite this page as:
Gron KJ. 2008 December 8. Primate Factsheets: Lesser bushbaby (Galago) Behavior . <http://pin.primate.wisc.edu/factsheets/entry/lesser_bushbaby/behav>. Accessed 2020 July 21.

INTERNATIONAL STATUS

For individual primate species conservation status, please search the IUCN Red List.
Also search the current scientific literature for primate conservation status (overall as well as for individual species), and visit CITES (Convention on International Trade in Endangered Species of Wild Fauna and Flora).

Conservation information last updated in 2008 follows, for comparison:

Generally, bushbabies are considered adaptable and some species may be able to cope with some habitat degradation (Butynski et al. 1998; Ambrose 2006; Butynski et al. 2006). In some cases, bushbabies are still found in agricultural areas mixed with forest remnants (Butynski et al. 2006). G. demidoff, for example, is able to live in secondary habitats and near areas of human disturbance and G. gallarum is present in habitats which are significantly degraded by the overgrazing of livestock (Ambrose & Perkin 1999-2000; Butynski & de Jong 2004). Another example is G. alleni, which is sometimes found near roads and cultivation and in recently logged forests (Ambrose 2003). However, some species are extremely threatened. This is the case with the Rondo dwarf bushbaby (Galago rondoensis), which is listed as one of the world’s 25 Most Endangered Primates (Mittermeier et al. 2007).

CONSERVATION THREATS

Threat: Human-Induced Habitat Loss and Degradation

As with most primates, habitat degradation, disturbance and loss are the most serious threats to many species of bushbaby (Butynski 1996/1997; Butynski et al. 1998; Ambrose 2006; Mittermeier et al. 2007). While bushbabies in general are quite widespread, some have more restricted ranges and are correspondingly, more susceptible comparable amounts of habitat loss and degradation (Bearder 2007). In Tanzania, for example, in addition to habitat destruction occurring as a result of logging, montane forest is cleared for agriculture and lowland forest is removed for rice, sugar, and rubber agriculture (Butynski et al. 1998). One of the most endangered bushbabies, G. rondonensis is threatened by the expansion of agriculture, charcoal manufacturing and logging (Mittermeier et al. 2007). Elsewhere, species in Uganda are threatened by the clearance of forest for gardens (Ambrose 2006).

In forests that have been logged, bushbabies are found at lower densities than in primary forests (Weisenseel et al. 1993).

Threat: Harvesting (hunting/gathering)

In Tanzania, bushbabies are not actively hunted and elsewhere in central and west Africa, are only very rarely found in bushmeat markets (review by Bowen-Jones & Pendry 1999; Jørgensbye 2007). However, even one of the smallest bushbabies, G. demidoff, is reported eaten on Bioko Island, West Africa (Albrechtsen et al. 2006). This may mean that even though they are diminutive, bushbabies are not immune from threats posed by hunting and the bushmeat trade.

Threat: Accidental Mortality

Galagos have been found in traps designed to capture birds in Equatorial Guinea (Garcia & Mba 1997).

LINKS TO MORE ABOUT CONSERVATION

CONSERVATION INFORMATION

• No current links for Galago
• Links for all species

CONSERVATION NEWS

• Strange endangered primates you may have never heard of  (BBC; January 25, 2012)
• Exotic Animals Found in Tanzanian Mountains  (LiveScience; June 23, 2006)
• Links for all species

ORGANIZATIONS INVOLVED IN Galago CONSERVATION

• Makerere University Biological Field Station
• Monkeyland Primate Sanctuary
• Projet Conservation de la Foret de Nyungwe (P.C.F.N.)

Content last modified: December 8, 2008

Written by Kurt Gron. Reviewed by Leanne Nash.

Cite this page as:
Gron KJ. 2008 December 8. Primate Factsheets: Lesser bushbaby (Galago) Conservation . <http://pin.primate.wisc.edu/factsheets/entry/lesser_bushbaby/cons>. Accessed 2020 July 21.

The following references were used in the writing of this factsheet. To find current references for Galago, search PrimateLit.

REFERENCES

Albrechtsen L, Fa JE, Barry B, Macdonald DW. 2006. Contrasts in availability and consumption of animal protein in Bioko Island, West Africa: the role of bushmeat. Environ Conserv 32(4):340-8.

Ambrose L, Perkin AW. 1999-2000. A survey of nocturnal prosimians at Moca on Bioko Island, Equatorial Guinea. Afr Prim 4(1-2):4-10.

Ambrose L. 2006. A survey of prosimians in the national parks and forest reserves of Uganda. In: Newton-Fisher NE, Notman H, Paterson JD, Reynolds V, editors. New York: Springer p 329-43.

Ambrose L. 2003. Three acoustic forms of Allen’s galagos (Primates; Galagonidae) in the central African region. Primates 44(1):25-39.

Anderson MJ, Ambrose L, Bearder SK, Dixson AF, Pullen S. 2000. Intraspecific variation in the vocalizations and hand pad morphology of southern lesser bush babies (Galago moholi): a comparison with G. senegalensis. Intl J Primatol 21(3):537-55.

Anderson MJ. 2001. The use of hair morphology in the classification of galagos (Primates, subfamily Galagoninae). Primates 42(2):113-21.

Anderson MJ. 1999. The use of hand morphology in the taxonomy of galagos. Primates 40(3):469-78.

Ankel-Simons F. 2007. Primate Anatomy: an introduction, 3rd Edition. San Diego: Elsevier Acad Pr. 724 p.

Bearder SK, Martin RD. 1980. Acacia gum and its use by bushbabies, Galago senegalensis (Primates: Lorisidae). Intl J Primatol 1:103-28.

Bearder SK. 2007. Afterward. Am J Primatol 69(1):131-4.

Bearder SK, Nekaris KAI, Buzzell CA. 2002. Dangers in the night: are some nocturnal primates afraid of the dark? In: Miller LE, editor. Eat or be eaten: predator sensitive foraging among primates. Cambridge: Cambridge U Pr. p 21-43.

Bearder SK, Doyle GA. 1974a. Ecology of bushbabies Galago senegalensis and Galago crassicaudatus, with some notes on their behaviour in the field. In: Martin RD, Doyle GA, Walker AC, editors. Prosimian biology. Pittsburgh: U Pittsburgh Pr. p 109-30.

Bearder SK, Doyle GA. 1974b. Field and laboratory studies of social organization in bushbabies (Galago senegalensis). J Hum Evol 3:37-50.

Bearder SK. 1987. Lorises, bushbabies, and tarsiers: diverse societies in solitary foragers. In: Smuts BB, Cheney DL, Seyfarth RM, Wrangham RW, Struhsaker TT, editors. Primate societies. Chicago: U Chicago Pr. p 11-24.

Bearder SK. 1999. Physical and social diversity among nocturnal primates: a new view based on long term research. Primates 40(1):267-82.

Bearder SK, Martin RD. 1980. The social organization of a nocturnal primate revealed by radio tracking. In: Amlaner Jr., CJ, Macdonald DW, editors. A handbook on biotelemetry and radio tracking. Oxford: Pergamon Pr. p 633-48.

Bearder SK, Honess PE, Ambrose L. 1995. Species diversity among galagos with special reference to mate recognition. In: Alterman L, Doyle GA, Izard MK, editors. Creatures of the dark: the nocturnal prosimians. New York: Plenum Pr. p 331-52.

Bearder SK, Ambrose L, Harcourt C, Honess P, Perkin A, Pimley E, Pullen S, Svoboda N. 2003. Species-typical patterns of infant contact, sleeping site use and social cohesion among nocturnal primates in Africa. Folia Primatol 74(5-6):337-54.

Bowen-Jones E, Pendry S. 1999. The threat to primates and other mammals from the bushmeat trade in Africa, and how this threat could be diminished. Oryx 33(3):233-46.

Butler H. 1967. Seasonal breeding of the Senegal galago (Galago senegalensis senegalensis) in the Nuba Mountains, Republic of the Sudan. Folia Primatol 5:165-75.

Butynski TM. 1996/1997. African primate conservation-the species and the IUCN/SSC primate specialist group network. Prim Conserv 17:87-100.

Butynski TM. 1982. Blue monkey (Cercopithecus mitis stuhlmanni) predation on galagos. Primates 23(4):563-6.

Butynski TM, de Jong. 2004. Natural history of the Somali lesser galago (Galago gallarum). J East Afr Nat Hist 93(1-2):23-38.

Butynski TM, Ehardt CL, Struhsaker TT. 1998. Notes on two dwarf galagos (Galagoides udzungwensis and Galagoides orinus) in the Udzungwa Mountains, Tanzania. Prim Conserv 18:69-75.

Butynski TM, de Jong YA, Perkin AW, Bearder SK, Honess PE. 2006. Taxonomy, distribution, and conservation status of three species of dwarf galagos (Galagoides) in eastern Africa. Prim Conserv 21:63-79.

Byrne RW. 2007. Animal cognition: bring me my spear. Curr Biol 17(5):R164-5.

Charles-Dominique P. 1972. Ecologie et vie sociale de Galago demidovii (Fisher 1808; Prosimii). Fortschritte Der Verhal Tensforschung 9:7-41.

Charles-Dominique P. 1977a. Ecology and behaviour of nocturnal primates: prosimians of equatorial West Africa. New York: Colombia U Pr. 277p.

Charles-Dominique P. 1974. Ecology and feeding behaviour of five sympatric lorisids in Gabon. In: Martin RD, Doyle GA, Walker AC, editors. Prosimian biology. London: Duckworth Co. Ltd. p 131-50.

Charles-Dominique P, Bearder SK. 1979. Field studies of Lorisid behavior: methodological aspects. In: Doyle GA, Martin RD, editors. The study of prosimian behavior. New York: Academic Pr. p 567-629.

Charles-Dominique P. 1977b. Urine marking and territoriality in Galago alleni (Waterhouse, 1837-Lorisoidea, Primates)-a field study by radio-telemetry. Z Tierpsychol 43:113-38.

Courtenay DO, Bearder SK. 1989. The taxonomic status and distribution of bushbabies in Malawi with emphasis on the significance of vocalizations. Intl J Primatol 10(1):17-34.

Crompton RW. 1983. Age differences in locomotion of two subtropical Galaginae. Primates 24(2): 241-59.

Crompton RH. 1984. Foraging, habitat structure, and locomotion in two species of Galago. In: Rodman PS, Cant JGH, editors. Adaptations for foraging in nonhuman primates: contributions to an organismal biology of prosimians, monkeys, and apes. New York: Columbia U Pr. p 73-111.

Doyle GA. 1974. The behaviour of the lesser bushbaby. In: Martin RD, Doyle GA, Walker AC, editors. Prosimian biology. Pittsburgh: U Pittsburgh Pr. p 213-31.

Doyle GA. 1979. Development of behavior in Prosimians with special reference to the lesser bushbaby, Galago senegalensis moholi. In: Doyle GA, Martin RD, editors. The study of prosimian behavior. New York: Academic Pr. p 157-206.

Doyle GA, Bearder SK. 1977. The Galagines of South Africa. In: Rainier III (Grimaldi) Prince of Monaco, Bourne GH, editors. Primate conservation. New York: Academic Pr. p 1-35.

Doyle GA, Andersson A, Bearder SK. 1969. Maternal behaviour in the lesser bushbaby (Galago senegalensis moholi) under semi-natural conditions. Folia Primatol 11:215-38.

Doyle GA, Andersson A, Bearder SK. 1971. Reproduction in the lesser bushbaby (Galago senegalensis moholi) under semi-natural conditions. Folia Primatol 14:15-22.

Garcia JE, Mba J. 1997. Distribution, status and conservation of primates in Monte Alen National Park, Equatorial Guinea. Oryx 31(1):67-76.

Gonzalez-Kirchner JP. 1995. The diet of sympatric prosimians in Equatorial Guinea. Folia Zool 44(1):13-8.

Groves C. 2005. Order primates. In: Wilson DE, Reeder DM, editors. Mammal species of the world: a taxonomic and geographic reference, third edition, volume 1. Baltimore (MD): Johns Hopkins U Pr. p 111-84.

Gucwinska H, Gucwinski A. 1968. Breeding the Zanzibar galago Galago senegalensis zanzibaricus. Intl Zoo Ybk 8:111-4.

Haddow AJ, Ellice JM. 1964. Studies on bush-babies (Galago spp.) with special reference to the epidemiology of yellow fever. Trans Royal Soc Trop Med Hyg 58(6):521-38.

Harcourt CS, Bearder SK. 1989. A comparison of Galago moholi in South Africa with Galago zanzibaricus in Kenya. Intl J Primatol 10(1):35-45.

Harcourt CS. 1981. An examination of the function of urine washing in Galago senegalensis. Z Tierpsychol 55:119-28.

Harcourt C. 1986a. Galago zanzibaricus: birth seasonality, litter size and perinatal behaviour of females. J Zool Lond 210(3):451-7.

Harcourt C. 1986b. Seasonal variation in the diet of South African galagos. Intl J Primatol 7(5):491-506.

Harcourt CS, Nash LT. 1986. Species differences in substrate use and diet between sympatric galagos in two Kenyan coastal forests. Primates 27(1):41-52.

Harvey PH, Clutton-Brock TH. 1985. Life history variation in primates. Evolution 39(3):559-81.

Honess PE, Bearder S. 1996. Descriptions of the dwarf galago species of Tanzania. Afr Prim 2(2):75-9

Izard MK, Nash LT. 1988. Contrasting reproductive parameters in Galago senegalensis braccatus and G. s. moholi. Intl J Primatol 9(6):519-27.

Izard MK, Simons EL. 1986. Infant survival and litter size in primigravid and multigravid galagos. J Med Primatol 15(1):27-35.

Izard MK. 1987. Lactation length in three species of Galago. Am J Primatol 13(1):73-6.

Jørgensbye H. 2007. Mountain galago Galagoides orinus in the evergreen forest fragments on the Dabaga plateau, Tanzania. Nat East Afr: EANHS Bull 35(2):7-12.

Kingdon J. 2004. The Kingdon pocket guide to African mammals. Princeton(NJ):Princeton U Pr. 272 p.

Lipschitz DL. 1996a. Male copulatory patterns in the lesser bushbaby (Galago moholi) in captivity. Intl J Primatol 17(6):987-1000.

Lipschitz DL. 1996b. A preliminary investigation of the relationship between ovarian steroids, LH, reproductive behaviour and vaginal changes in lesser bushbabies (Galago moholi). J Reprod Fert 107(2):167-74.

Lipschitz DL, Galpin JS, Meyer D. 2001. Reproductive behavioral changes during the ovarian cycle of lesser bushbabies (Galago moholi) in captivity. Am J Primatol 55(2):101-15.

Llorente M, Pi JS, Houle A. 2003. Association between Galago thomasi and Pan troglodytes schweinfurthii in the Kibale National Park, Uganda. Folia Primatol 74(2):80-4.

Mascagni O, Doyle GA. 1993. Infant distress vocalizations in the southern African lesser bushbaby (Galago moholi). Intl J Primatol 14(1):41-60.

Milikazi N, Masters JC, Lovegrove BG. 2006. Lack of torpor in free-ranging southern lesser galagos, Galago moholi: ecological and physiological considerations. Folia Primatol 77(6):465-76.

Mittermeier RA, Ratsimbazafy J, Rylands AB, Williamson L, Oates JF, Mbora D, Ganzhorn JU, Rodríguez-Luna E, Palacios E, Heymann EW, Kierulff MCM, Yongcheng L, Supriatna J, Roos C, Walker S, Aguiar JM. 2007. Primates in peril: the world’s 25 most endangered primates, 2006-2008. Prim Conserv 22:1-40.

Molez N. 1976. Adaptation alimentaire du galago d’Allen aux milieux forestiers secondaires. Terre Et La Vie 30:210-28.

Nash LT, Flinn L. 1978. Group formation in captive lesser galagos (Galago senegalensis). Primates 19(3):493-503.

Nash LT. 1986. Influence of moonlight level on traveling and calling patterns in two sympatric species of Galago in Kenya. In: Taub DM, King FA, editors. Current perspectives in primate social dynamics. New York: Van Nostrand Reinhold Co. p 357-67.

Nash LT. 1993. Juveniles in nongregarious primates. In: Pereira ME, Fairbanks LA, editors. Juvenile primates: life history, development, and behavior. New York: Oxford U Pr. p 119-37.

Nash LT, Whitten PL. 1989. Preliminary observations on the role of Acacia gum chemistry in Acacia utilization by Galago senegalensis in Kenya. Am J Primatol 17(1):27-39.

Nash LT. 1983. Reproductive patterns in galagos (Galago zanzibaricus and Galago garnettii) in relation to climatic variability. Am J Primatol 5(3):181-96.

Nash LT. 2003. Sex differences in the behavior and the social interactions of immature Galago senegalensis braccatus. Folia Primatol 74(5-6):285-300.

Nash LT. 1984. Social organization of two sympatric galagos at Gedi, Kenya. In: Else JG, Lee PC, editors. Primate ecology and conservation. Cambridge: Cambridge U Pr. p 125-31.

Nash LT, Bearder SK, Olson TR. 1989. Synopsis of Galago species characteristics. Intl J Primatol 10(1):57-80.

Nekaris A, Bearder SK. 2007. The lorisiform primates of Asia and mainland Africa: diversity shrouded in darkness. In: Campbell CJ, Fuentes A, MacKinnon KC, Panger M, Bearder SK, editors. Primates in perspective. New York: Oxford U Pr. p 24-45.

Olson TR, Nash LT. 2002-2003. Galago (Galagidae) body measurements and museum collections data. Afr Prim 6(1-2):50-3.

Off EC, Gebo DL. 2005. Galago locomotion in Kibale National Park, Uganda. Am J Primatol 66(2):189-95.

Perkin A. 2007. Comparative penile morphology of east African galagos of the genus Galagoides (Family Galagidae): implications for taxonomy. Am J Primatol 69(1):16-26.

Perkin A, Bearder S, Butynski TM, Agwanda B, Bytebier B. 2002. The Taita Mountain dwarf galago Galagoides sp: a new primate for Kenya. J East Afr Nat Hist 91(1-2):1-13.

Perkin A. 2001. The taxonomic status and distribution of bushbabies (Galagos) in the Uluguru Mountains. Miombo 23:11-3.

Pimley ER, Bearder SK, Dixon AF. 2005. Home range analysis of Perodicticus potto edwardsi and Sciurocheirus cameronensis. Intl J Primatol 26(1):191-206.

Pitts RS. 1988. Classical heinrothian intimidation displays exhibited by Galago demidoff demidoff: a paradigm for pinpointing Galagine species diversity. Intl J Primatol 9(6):529-56.

Poulsen JR, Clark CJ. 2001. Predation on mammals by the grey-cheeked mangabey Lophocebus albigena. Primates 42(4):391-4.

Pruetz JD, Bertolani P. 2007. Savanna chimpanzees, Pan troglodytes verus, hunt with tools. Curr Biol 17(5):412-7.

Pullen SL, Bearder SK, Dixson AF. 2000. Preliminary observations on sexual behavior and the mating system in free-ranging lesser galagos (Galago moholi). Am J Primatol 51(1):79-88.

Riordan DV. 1971. Reproductive variation in Galago senegalensis sub-species (lesser bush baby). Ann Rep Jersey Wildl Preserv Trust 8:15-8.

Ross C. 1988. The intrinsic rate of natural increase and reproductive effort in primates. J Zool London 214(2):199-219.

Struhsaker TT. 1970. Notes on Galagoides demidovii in Cameroon. Mammalia 34:207-11.

Vinyard CJ. 2007. Interspecific analysis of covariance structure in the masticatory apparatus of galagos. Am J Primatol 69(1):46-58.

Ward JP, Scott S. 1970. Observations of two infant bushbabies, Galago senegalensis. Lab Prim News 9(4):3-6.

Weisenseel K, Chapman CA, Chapman LJ. 1993. Nocturnal primates of Kibale forest: effects of selective logging on prosimian densities. Primates 34(4):445-50.

Wickings EJ, Ambrose L, Bearder SK. 1998. Sympatric populations of Galagoides demidoff and Galagoides thomasi in the Haut-Ogooué region of Gabon. Folia Primatol 69(suppl 1):389-93.

Zimmermann E. 1989. Aspects of reproduction and behavioral and vocal development in Senegal bushbabies (Galago senegalensis). Intl J Primatol 10(1):1-16.

Zimmermann E. 1990. Differentiation of vocalizations in bushbabies (Galaginae, Prosimiae, Primates) and the significance for assessing phylogenetic relationships. Z Zool Syst Evol Forsch 28(3):217-39.

Zimmermann E. 1995. Loud calls in nocturnal prosimians: structure, evolution and ontogeny. In: Zimmermann E, Newman JD, Jürgens U, editors. Current topics in primate vocal communication. New York: Plenum Pr. p 47-72.

Zimmermann E, Bearder SK, Doyle GA, Andersson AB. 1988. Variations in vocal patterns of Senegal and South African lesser bushbabies and their implications for taxonomic relationships. Folia Primatol 51(2-3):87-105.

Content last modified: December 8, 2008

IMAGES

	Galago moholi Photo: Gerald Doyle
	Galago moholi Photo: Gerald Doyle
	Galago moholi Photo: Gerald Doyle
	Galago moholi Photo: Gerald Doyle
	Galago moholi Photo: Gerald Doyle
	Galago moholi Photo: Gerald Doyle
	Galago senegalensis braccatus Photo: R. A. Barnes