Synopsis of Galago species characteristics

International Journal o f Primatology, Vol. 10, No. 1, 1989

Synopsis of Galago Species Characteristics L e a n n e T. N a s h , I S i m o n K. Bearder, 2 and T o d d R . O l s o n 3 Received May 17, 1988; revised May 2, 1989

A t the time o f the symposium, "Variability Within the Galagos, " at the 1986 IPS Congress, most participants were still using Hill's (1953) classification and nomenclature o f galago species. All participants expressed some degree o f dissatisfaction with Hill's species groups. Many described how it was proving increasingly problematic and inadequate as a means to organize newly collected laboratory, museum, and field data. By the end o f the symposium, a consensus about species diversity had emerged which synthesized the current state o f knowledge about galagos. The consensus o f the participants was that the 11 species, identified by Olson (1979, 1986), most closely approximates the available data about galago species diversity. The 11 species are described. KEY WORDS: Galago species; variability;classification.

CURRENT VIEWS ON GALAGO TAXONOMY Of the several alternative treatments to Schwarz' (1931) seminal taxonomic study, only Hill's (1953) has been widely utilized. Hill's classification introduced only a few minor changes to Schwarz' original taxonomic scheme. Schwarz (1931) grouped the galagos into two genera, Galago and Euoticus, which contained a total of five spcecies: Galago crassicaudatus, G. senegalensis, G. "demidovii, " G. alleni, and Euoticus elegantulus. Hill (1953) modified this scheme by placing Demidoff's galago in a third genus, Galagoides,

tDepartment of Anthropology, Arizona State University, Tempe, Arizona 85287-2402. 2Anthropology Unit, Department of Social Science, Oxford Polytechnic, Oxford O X 3 0 B P , U.K. 3Department of Anatomy and Structural Biology, Albert Einstein College of Medicine, 1300 Morris Park Avenue, Bronx, New York 10461. 57 01~1/89/0~0-C057506.00/0 9 19~ PlenumPublishingCorporation

58

Nash, Bearder, and Olson

and recognizing Schwarz' (1930) subspecies G. s. inustus (junior synonym of Galago matschiei Lorenz, 1917) as a sixth species, which Hill reclassified in his genus Euoticus. Between the appearance of Hill's (1953) study and Olson's (1979) revision, discussion of galago classification was focused almost entirely on the grouping of species into subgenera and genera, and not on the number of species. Napier and Napier (1967) recognized Hilrs six species but reduced Hill's three genera to subgenera within a single genus, Galago. Groves (1974) suggested that Demidoff's and Matschie's galago should remain in the subgenus Galago, whereas the greater galago should be separated in its own subgenus or genus, Otolemur. Petter and Petter-Rousseaux (1979) followed the subgeneric arrangement of Napier and Napier (1967), while Jouffroy and Lessertisseur (1979), also identifying six species, classfied Alien's galago as a fourth subgenus Otolicnus, in their genus Galago. Kingdon (1971) is one of the first authorities to question the validity of Hill's six galago species. Kingdon (1971, p. 309) recognized one of Hill's subspecies of Galago senegalensis as a seventh species: Galago zanzibaricus. Although accepted by Groves (1974) and Olson (1979), this interpretation of the specific status of the Zanzibar galago was not generally accepted at the time of the symposium. Dixson and Van Horn (1977) documented that the greater galagos consist of two morphologically distinct groups which are reproductively isolated, at least in captivity. Their suggestion that two greater galago species may therefore exist in the wild was substantiated by the work of Olson (1979, 1980), Masters (1985), and others. Although many symposium participants had accepted the evidence for two rather than one greater galago species, few chose to reflect this in their nomenclatorial treatment of the group before the symposium. Olson's (1979) revised classification (Table I), which focused on the greater galagos, was based on data which he obtained from the field, literature, and the examination of 4949 specimens from 57 museums and private collections in Africa, Europe, and North America. Olson collected provenience data from 1497 localities and was the first reviewer of this group to examine all existing type specimens of the 58 available galago species group names. Analyzing the geographic distribution of both quantitative and qualitative variation, Olson concluded that Hill's (1953) six galago species encompassed 11 phenotypically distinct species as shown in Fig. 1.

Classification Above the Species Level Recent classifications have arranged the galagos above the species level into one, two, or three genera based upon the author's opinion about the

Synopsis of Galago Species Characteristics

59

Table I. Classification for the Galagos Proposed by Olson (1979, 1986): Common Name Follows Binominal Name Genus Subgenus Species

Galago E. Geoffroy, 1796 Galago E. Geoffroy, 1796 Galago senegalensis E. Geoffroy, 1796 Senegal galago

Galago gallarum Thomas, 1901 Somali galago

Galago moholi A. Smith, 1836 Mohol or South African lesser galago Subgenus Species

Euoticus Gray, 1863 Galago elegantulus (Le Conte, 1857) Elegant galago

Galago matschiei Lorenz, 1917 Matschie's galago Genus Subgenus Species

Galagoides A. Smith, 1833 Galagoides A. Smith, 1833 Galagoides deraidoff (Fischer, 1806) Demidoff's galago

Galagoides thomasi (Elliot, 1907) Thomas' galago

Galagoides zanzibaricus (Matschie, 1893) Zanzibar galago Subgenus Species

Sciurocheirus Gray, 1873 Galagoides alleni (Waterhouse, 1838) Allen's galago

Genus Species

Otolemur Coquerel, 1859 Otolemur crassicaudatus (E. Geoffroy, 1812) Large-eared greater galago

Otolemur garnettii (Ogilby, 1838) Garnett's or small-eared greater galago

phylogenetic affinities, distinctiveness, or similiarities of the different galago species. Several symposium participants expressed their preference for a classification which placed all of the galagos in a single genus to emphasize their overall similarity rather than their distinctiveness. Olson continued to support a phylogenetically based grouping of the 11 species which he recognized. Other participants also expressed their preference for a phylogenetically based classification but felt there were insufficient data to assess the evolutionary affinities of the 11 galago species recognized by Olson (1979). It was clear from the lively symposium discussions that a number of different supraspecific classifications will continue to be used. As the use of a variety of generic and subgeneric schemes will no doubt persist into the near-future, we believe that it is essential, in order to avoid confusion and to promote a better understanding of the galagos, that authors reference the source of their classificatory and nomenclatorial arrangements (e.g., Hill, 1953; Napier and Napier, 1967; Petter and Peter-Rousseaux, 1979; Olson, 1979) and, when possible, to indicate their reasons for selecting a particular scheme.

60

Nash, Bearder, and Olson

Sill

(1953)

Olson

(1979)

crassicaudatus crassicaudatus--~-----garnettii

___-senega]ensis

~

mohoJi

senega]ensis

"inuBtus" "demidovii

n

gallarum matschiei demidoff ~

thomasi sanzibaricus

alleni

alleni

elefantuJus

elefantu]us

Fig. 1. Comparison of species groups recognized by Hill (1953) and Olson (1979) showing how taxa included in Hill's six species were separated by Olson. The identity of Matschie's galago is noteworthy, as Olson found that Hill had representatives of this taxon in three of his species.

As the three authors of this paper favor different generic groupings for the 11 galago species which we all recognize, we have decided to refer to each species by the common name adopted in the following synopsis.

Subspecific Classification Hill (1953) described a total of 31 subspecies in his six species. Two of Hill's species, "Galago" alleni and "Galago inustus, " are monotypic and did not contain subspecies. Olson (1979) recognized 28 subspecies, including five yet undescribed taxa, within the 11 species identified in his revision of this group. All of Olson's species are polytypic and include at least two subspecies. A close review of the evidence for the species recognized in this synopsis indicates that the data for subspecific variation are the least convincing in the mohol and Matschie's galagos. As there remain four or five subspecies of extant galago yet to be described, the decision was made not to consider galago classification at the

Synopsis of Galago Species Characteristics

61

subspecific level in this synopsis and, instead, to focus on species which have recently been accepted by those who work on galagos in order to facilitate species identification and to provide a basis for further research. SYNOPSIS OF SPECIES CHARACTERISTICS Characteristics pertinent to the I1 species recognized by Olson (1979, 1986) are presented here to serve as a basis for the identification of both museum specimens and living animals in the field or in captivity. The synopsis also indicates some important areas where data are lacking or deficient for each species. For example, more fieldwork is needed in areas o f apparent sympatry between species, in particular, where the Senegal galago occurs with the mohol or Somali galago and where Demidoff's galago is associated with Thomas', Matschie's, or the Zanzibar galago. Further studies are necessary on chromosomes, visual and vocal repertoires, locomotor styles, and penile morphologies in order to characterize and distinguish most species more precisely. There is also a particular need for data on all aspects, including the endangered status, of Thomas' galago, Matschie's galago, and the Somali galago.

General Notes on Descriptive Categories The synopsis is designed to differentiate species within three body size classes. The sequence of species within each size grade is from largest to smallest, based on mean adult male and female body weights. "Body size" data are from wild-caught specimens only and do not reflect the larger sizes frequently associated with captive animals of the same taxon. Species characteristics listed below are drawn from this symposium, with additional data derived from most previously cited sources as well as De Boer (1973), Harcourt and Nash (1986), Nash and Harcourt (1986), Bearder (1987), and Nash, Bearder, and Olson (Personal observation). Throughout this work quotation marks are used in conjunction with binominals to indicate, when it is not contextually obvious, that the nomen was used by a cited authority. The use o f quotation marks provides a means by which genus and species group names can be used in their original association and form without implying either that the taxon is valid or that the name associated with it by the cited author is its correct name. For example, the citation "Otolicnus" garnettii followed by authority and date indicates that garnettii was attributed to the genus Otolicnus by the cited authority and that the present authors accept the validity of garnettii as a species but reject Otolicnus as its valid or appropriate generic name.

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Nash, Bearder, and Olson

DESCRIPTIVE CATEGORIES

Large (Thick-Tailed) Galagos: > 550g

Species: "Galago" crassicaudatus E. Geoffroy, 1812 Common Name: Large-eared greater galago Type Locality: Ngoye Forest, South Africa Geographic Range (Fig. 2): Range corresponds closely to the Brachystegia or miombo vegetation zone south and east of the Zaire Basin. Range in East Africa primarily in Tanzania and Kenya, limited to Kisii and the Masai Mara regions: not known from Namibia or Botswana. In South Africa limited to an area east of the Transvaal. Habitat Preferences: Occurs in subtropical and tropical forests, riverine and coastal forests, and woodland/savannah. Appears to prefer more densely forested habitats in the southern and western parts o f its range. Except for east and west of Lake Victoria, its occurrence in the northeastern portion of the range is associated primarily with open woodland/savannah habitats. Sympatric Associations: Occurs with the Garnett's galago throughout most of the eastern half of Tanzania. Ecologically separated within most of this shared range. One of us (T.R.O.) has identified in the field an area extending from the Usambara Mountains to Ngorongoro Crater in the

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Fig. 2. Range of the large-eared greater galago.

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60"

Synopsis of Galago Species Characteristics

63

tran.sition zone between the Masai Steppe and the montane forest where both large-eared galago species occupy the same habitat. Its range is encompassed by that of the mohol galago. See figures for additional species with overlapping ranges. Characteristics: Largest of the galagos; appearance of head dominated by large ears and large robust muzzle; dorsal pelage highly variable in color, ranging from silver-gray to dark brown (monochrome melanistic individuals are known from Kenya, Tanzania, and eastern Zaire); ventrum variable; flanks of limbs and face typically same color as body; face markings absent (Fig. 12a); distal end of nails convex without lateral points. Infants carried both dorsally and orally.

Body Size: Head and body length (mm): 255-400 (mean = 313) Tail length (mm): 300-550 (mean = 410) Ear height (mm): 48-72 (mean = 62) Hindfoot length (mm): 70-108 (mean = 93) Body weight (g): 567-1814 (mean = 1131) There is evidence to suggest a significant degree of sexual dimorphism in wild populations. In South Africa, Harcourt (1980)has reported means for male and female body weight as close to 1500 and 1250 g, respectively. Locomotion: Quadrupedal running and climbing: bipedal hopping rare; unable to land hindfeet first when leaping. Gestation Length: 135 days Litter Size: 2 or 3 normally Penile Morphology: Baculum smaller than in Garnett's galago, uni- or bidentate penile spines. Advertisement Call: Sequence of 3-17 (median, 8) drawn-out loud cries repeated at regular intervals, the last of which fades away. Total duration usually more than 4 sec. Diet: Invertebrates, fruit, and gum Karyotype: 62 chromosomes; fundamental number = 76

Species: "Otolicnus" garnettii Ogilby, 1838 Common Name: Garnett's or small-eared greater galago Taxonomic Note: Following Thomas (1917) and Olson (1979), Garnett's galago is identified as an East African greater galago.

Type Locality: Zanzibar Geographic Range (Fig. 3): Coastal regions of East Africa ranging from the Juba River area in Somalia to the Ruvuma River in Tanzania. Common throughout the Kenyan Highlands and on the islands of Pemba, Zanzibar, and Mafia. While not known from the Macua region of Mozambique, it almost certainly occurs along the coast and up larger rivers in this region.

64

Nash, Bearder, and Olson

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10"

Fig. 3. Range of Garnett's galago.

Habitat Preferences: Restricted to coast, riverine, and highland forests. Not known to inhabit woodland/savannah.

Sympatric Associations: Within the coastal and riverine forest in its range, Garnett's galago occurs with the Zanzibar galago. Appears to be ecologically separated from the Senegal galago in areas of central Kenya where their ranges overlap. Characteristics: Large size; ears small relative to head; dorsal pelage reddish to a grayish-brown in color (melanistic forms unknown); ventrum variable; flanks of limbs and face same color as body; facial markings absent (Fig. 12b); tail tip highly variable with black-, white-, and brown-tipped individuals occurring in single populations; distal end of nails concave with lateral points. Infants carried only by mouth.

Body Size: (mean = 266) Head and body length (ram): 230-338 308-440 (mean = 364) Tail length (mm): 34-55 (mean = 45) Ear height (mm): 80-103 (mean = 91) Hindfoot length (ram): 550-1040 (mean = 767) Body weight (g): There is some evidence to suggest that a significant degree of sexual dimorphism exists in wild populations. Nash and Harcourt (1986) report mean body weights in coastal Kenya as Close to 820 g for males and 720 g for females.

Synopsis of Galago Species Characteristics

65

Locomotion: Quadrupedal running and climbing and bipedal hopping; individuals are able to land hindfeet first when leaping.

Gestation length: 130 days Litter size: 1 or 2 normally Penile Morphology: Baculum large, penile spines usually tridentate, with some bidentate.

Advertisement Call: Sequence o f 6-10 brief loud cries, reaching a crescendo and trailing o f f over the last 3 or 4 units. Total duration usually less than 4 sec. Diet: Invertebrates and fruit Karyotype: 62 chromosomes; fundamental number = 90 Medium-Sized Galagos: 125-550 g

Species: "Galago" alleni Waterhouse, 1838 Common Name: Allen's galago Type Locality: Fernando P o Geographic Range (Fig. 4): Forested areas between the Niger and the Zaire Rivers around the Gulf o f Guinea and on Fernando Po. May range as far east as the Ubangui River but exact inland extent is uncertain.

Fig. 4. Range of Allen's galago.

66

Nash, Bearder, and Olson

Habitat Preferences: Prefers understory of primary rain forest. Sympatric Associations: Found with both Demidoff's and the elegant galago over almost entire range. North-south range may be slightly more extensive than that of the elegant galago. May also occur with Thomas' galago on Mount Cameroun. Characteristics: Moderate size; head long and narrow with pointed muzzle; dorsal pelage very dark brown to almost black; ventrum light in color; flanks of limbs brightly colored; face with dark circumocular rings and lacking prominent interorbital strip (Fig. 12c); taft bushy with or without darker tip; nails not pointed.

Body Size: Head and body length (ram): 155-240 (mean = 199) Tail length (mm): 205-300 (mean = 261) Ear height (ram): 30-45 (mean = 37) Hindfoot length (ram): 55-77 (mean = 69) Body weight (g): 200-445 (mean = 314) Locomotion: Leaping between vertical supports. Hindfeet are not brought forward to make initial contact on landing. Gestation Length: 133 days Litter Size: 1 normally, 2 maximum Penile Morphology: Penile spines unidentate Advertisement Call: Loud, low-pitched croaks emitted with increasing intensity, usually in a sequence of 10-30 units (median 12) separated by approximately 2-sec intervals ["croaking" call (Charles-Dominique, 1977)]. Diet: Mainly fallen fruit; invertebrates Karyotype: 40 chromosomes; fundamental number = ?

Species: "Microcebus" elegantulus Le Conte, 1857 Common Name: Elegant or needle-clawed galago Type Locality: Njola, Ogowe River, Gabon Geographic Range (Fig. 5): Forested areas between the Niger and the Zaire Rivers around the Gulf of Guinea and on Fernando Po. May range as far east as the Ubangui River but exact inland extent is uncertain. Habitat Preferences: Primary/secondary rain forest Sympatric Associations: Found with both Demidoff's and Allen's galago over almost entire range. May also occur with Thomas' galago on Mount Cameroun. Characteristics: Moderate size; head dominated by large eyes and short muzzle; dorsal body grayish- to reddish-brown; ventrum gray; flanks of limbs lightly colored; tail bushy with or without tip; face not prominently colored (Fig. 12d); nails of all digits except first keeled dorsally and pointed terminally.

67

Synopsis of Galago Species Characteristics

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Fig. 5. Ranges of the elegant and Matschie's galagos.

Body Size: Head and body length (ram): 106-270 (mean = 188) Tail length (mm): 194-337 (mean = 283) Ear height (mm): 24-40 (mean = 30) Hindfoot length (mm): 34-67 (mean = 57) Body weight (g): 223-350 (mean = 293) Locomotion: Leaping and running. Minimal body rotation during leaps; landing normally done on all. four extremities simultaneously. Gestation Length: ? Litter Size: 1(?) normally Penile Morphology: Characteristic frills around urethral opening, penile spines unidentate. Advertisement Call: A short, high-pitched sound which is usually uttered at the end of the night without any organization into definite sequences ["tsic" call (Charles-Dominique, 1977)]. Diet: Mainly gum and invertebrates Karyotype: Chromosomes ? fundamental number = ? Species: "Galago" matschiei Lorenz, 1917 Common Name: Matschie's galago Taxonomic Note: Lorenz (1917) was the first to recognize the unique identity of the gaiago which Schwarz subsequently named "Galago senegalensis

Nash, Bearder, and Olson

68

inustus" in 1930. Accordingly, the name "G." matschiei, given to this species by Lorenz, has priority and is the correct name for this taxon. Reviewers prior to Olson (1979) mistakenly considered Lorenz's type specimen to be a junior synonym of Thomas' galago. Type Locality: Mbau, Kivu Provence, Zaire Geographic Range (Fig. 5): Known only from the Kivu region of eastern Zaire, the Western Province and Mt. Moroto in Uganda. Given its distribution in Uganda, it would not be surprising to find this species in the few sites in Buganda Province that are still densely forested. Habitat Preferences: Found in relic tropical forest which is now restricted to the highlands east, south, and west of the Zaire Basin. Kingdon (1971, p. 325) reported that "the animal is found in medium altitude forest where Parinari excelsa is the dominant tree, it also occurs along the forest margins" and that it "moves freely between the canopy and the thick tangles at lower levels." Sympatric Associations: Occurs with Thomas' galago throughout its range with the possible exception of Mt. Moroto. Characteristics: Moderate size; dorsal pelage dark "burnt" brown; flanks of limbs similar to dorsum except over shoulders, where it may have a yellowish tinge; face with very dark orbital rings and distinct interorbital strip; eyes and orbits relatively larger than in all except the elegant galago (Fig. 12e); tail not bushy; nails keeled and pointed as in the elegant galago but keeling less pronounced.

Body Size: Head and body length (mm): Tail length (mm): Ear height (mm): Hindfoot length (mm): Body weight (g):

147-184 240-279 37-42 63-70 196-225

(mean (mean (mean (mean (mean

= 166) = 255) = 39) = 68) = 210)

Locomotion: ? Gestation Length: ? Litter Size: 1 (Kindgon, 1971) Penile Morphology: ? Advertisement Call: ? Diet: Invertebrates, fruit, and gum, which Kingdon (1971) suggested were consumed according to seasonal preference or availability.

Karyotype: Chromosomes ? fundamental number = ? Species: "Galago" senegalensis E. Geoffroy, 1796 Common Name: Senegal galago Type Locality: Senegal. Geographic Range (Fig. 6): Ranging across Africa from Senegal to the Gulf of Aden in the area between the Sahara and coastal forest in the

69

Synopsis of Galago Species Characteristics

Fig. 6. Rangeof the Senegal galago. West and the Zaire Basin in Central Africa. In East Africa, it occurs throughout Uganda, Kenya, and Tanzania. Habitat Preferences: Open woodlands over most of its range but not atypically in closed forests, particularly in the area east of Lake Victoria, where it occupies the most densely forested part of its range. Sympatric Associations: The only galago found within its habitat in West and Central Africa. Occurs with a variety of galagos in East Africa--the Somali galago in southern Ethiopia and along the Tana River, the mohol galago in Tanzania, the large-eared greater galago in Tanzania and and around Lake Victoria, and possibly Matschie's galago in southern Uganda. Characteristics: Moderate size; head broad with short muzzle and relatively large eyes and small ears, dorsal pelage gray to brownish-gray; flanks of limbs variable but always distinctly yellow in color; interocular strip and dark circumocular ring prominent (Fig. 12f); tall not bushy; nails not pointed.

Body Size: Head and body length (mm): Tail length (mm): Ear height (mm): Hindfoot length (mm): Body weight (g):

132-210 195-303 21-57 52-78 112-300

(mean (mean (mean (mean (mean

= = = = =

165) 261) 37) 69) 206)

Nash, Bearder, and Olson

70

There is strong evidence from wild-caught specimens to suggest a significant degree of sexual dimorphism. Data on wild-caught specimens analyzed by Olson (personal observation) indicate a mean male body weight of close to 225 g, with the female mean closer to 200 g. Locomotion: Bipedal hopping and active leaping, usually landing hindfeet first. Gestation Length: 142 days Litter Size: 1 normally, 2 maximum

Penile Morphology: ? Advertisement Call: Single-unit, low-pitched calls repeated at regular intervals for up to 1 hr.

Diet: Gums, invertebrates, and possibly fruit Karyotype: 36 chromosomes; fundamental number = ? Species: "Galago" gallarum Thomas, 1901 Common Name: Somali galago Type Locality: Dawa River in the vicinity of the common border of Somalia, Kenya, and Ethiopia

Taxonomic Note: After its original description, universally identified as a subspecies of the Senegal galago prior to Olson (1979), who found that it was sympatric with the Sengal galago (sensu strictu) in southern Ethiopia and along the Tana River. Geographic Range (Fig. 7): Occurs within the area bounded to the northwest by the Ethiopian Rift Valley, to the northeast by the Shebele River, to the southwest by the Tana River, and to the southeast by the Somali coastal zone. Habitat Preferences: There are no contemporary field accounts of this species. L6nnberg (1913, p. 45) reported that it is "not uncommon in the dry thornbush country." Drake-Brockman (1910, p. 8) states, "It is always found living in the tall acacia trees so common along the large riverbeds." From its range, it would appear that this is the most xerically adapted of all the galagos. Sympatric Associations: Occurs with the Senegal galago along the north and southeast margins of its range. May be found with the Zanzibar and Garnett's galagos along riverine forest margins of the lower reaches of the Tana and Giuba Rivers, but this has yet to be established. Characteristics: Moderate size, being slightly smaller than the Senegal galago; head broad with short muzzle and relatively large eyes; dorsal pelage buff to sandy-brown; ventrum white to light gray; flanks of limbs brightly colored; face lighter in color than ventrum; interocular strip prominent because of circumocular rings, brown and usually incomplete laterally (Fig. 12g); tail not bushy, being black over much of its length; nails not pointed.

Synopsis of Galago Species Characteristics

71

0

:

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Fig. 7. Range of the Somali galago.

Body Size: Head and body length (mm): 130-200 (mean = 167) Tall length (ram): 205-293 (mean = 252) Ear height (ram): 30-40 (mean = 35) Hindfoot length (ram): 57-75 (mean = 62) Body weight (g): Unknown Locomotion: Drake-Brockman (1910, p. 7) observed it frequently on the ground feeding and "jumping on its hindlegs like a jerboa." Lfnnberg (1913) reported them leaping with great dexterity in acacia bush. Gestation Length: ? Litter Size: ? Penile Morphology: ? Advertisement Call: ? Diet: Drake-Brockman (1910) states that its "food consists of the seeds of the 'gurha' tree, insects, and probably the fruit of the 'gob' tree." He also speculates that there are seasonal dietary patterns based on availability. Karyotype: Chromosomes ? fundamental number = ? Species Name: "Galago" moholi A. Smith, 1836 Common Name: Mohol or South African lesser galago Type Locality: Near Limpopo River between points of confluence with the Marico and Notwani Rivers.

72

Nash, Bearder, and Olson

........./ ......

\ Ill

Fig. g. Range of the mohol galago.

Taxonomic Note: Until recently interpreted as a subspecies of the Senegal galago. Olson (1979) elevated it to separate-species status based on distinct morphology and sympatric occurrence with the Senegal galago (sensu strictu) in Tanzania. Geographic Range (Fig. 8): Most of southern Africa, known only from the eastern part of South Africa, northern Mozambique, particularly western and southern Tanzania, and easternmost Zaire. Habitat Preferences: Savannah, woodland, riverine bush, and forest fringes Sympatric Associations: Shares a similar range with the large-eared greater galago but extending into drier, more open country; occurs in the same habitat as the Senegal galago in parts of Tanzania and is either sympatric or parapatric with Thomas' galago in the Kivu Region of Zaire and with the Zanzibar galago in parts of Malawi, Mozambique, and Zimbabwe. Characteristics: Moderate size but smaller than the Senegal galago; head broad with short muzzle and relatively large eyes; ears proportionately larger than in either the Senegal or the Somali galagos; dorsal pelage brownish-gray to light brown; flanks of limbs variable but always colored a distinct yellow; interocular and circumocular marking present (Fig. 12h); tail not bushy; nails not poim.ed, Schwartz (1974) reported that the mohol, Somali, and Senegal galagos possess slightly different dental eruption sequences.

Synopsis of Galago Species Characteristics

73

Body Size: Head and body length (mm): 88-205 (mean = 150) Tail length (mm): 113-279 (mean = 228) Ear height (mm): 23-50 (mean = 39) Hindfoot length (mm): 37-78 (mean = 59) Body weight (g): 95-244 (mean = 158) There is some evidence to suggest a significant degree of sexual dimorphism in wild populations and that size may vary geographically. Harcourt and Bearder (personal communication) studied two adjacent populations of the mohol galago in the Transvaal of South Africa and found mean male body weights to be approximately 210 and 195 g, while the female means in these respective samples were close to 185 and 165 g. Locomotion: Bipedal hopping and active leaping, usually landing hindfeet first. Gestation Length: 125 days Litter Size: 2 normally, 2 maximum Penile Morphology: ? Advertisement Call: Double-unit, high pitched, calls mixed with single- and triple-unit calls in a series lasting for up to 1 hr. Diet: Invertebrates and gums Karyotype: 38 chromosomes; fundamental number = 70 Species: "Galago" zanzibaricus Matschie, 1893 Common Name: Zanzibar galago Type Locality: Jambiani, Zanzibar Geographic Range (Fig. 9): Zanzibar but absent on Pemba and Mafia Islands; coastal Tanzania, Kenya, and Somalia; throughout Mozambique and Malawi; southest Zimbabwe. Habitat Preferences: Coastal and evergreen forests Sympatric Associations: Found with Garnett's galago in coastal areas of Tanzania, Kenya, and Somalia. Range overlaps with that of the largeeared greater galago and the mohol galago in Mozambique, Zimbabwe, and Malawi where they almost always occupy different habitats. Characteristics: Moderate size, being only slightly smaller than the mohol galago; muzzle long; pointed and concave dorsally; dorsal pelage cinnamon reddish-brown; ventrum bright yellow; flanks of limbs same or slightly lighter in color than dorsum; interocular and circumocular markings prominent (Fig. 12i); tail moderate to bushy and with a dark tip of variable size. Body Size: Head and body length (ram): 120-190 (mean = 153) Tail length (ram): 170-265 (mean = 220)

74

Nash, Bearder, and Olson

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Fig. 9. Range of the Zanzibar galago.

Ear height (mm): 27-46 (mean = 35) Hindfoot length (mm): 47-70 (mean = 69) Body weight (g): 104-203 (mean = 145) Harcourt and Nash (1986) found in coastal Kenya that the mean male body weight o f 160 g was significantly higher than the mean for females of 135 g. Locomotion: Quadrupedal running, no bipedal hopping or landing hindfeet first. Gestation Length: 120 days Litter Size: 1 normally, 2 maximum

Penile Morphology: ? Advertisement Call: Sequence of up to 18 double or triple units usually lasting a total o f 3-6 see, reaching a crescendo over the first few units and frequently trailing into a rapid series of staccato notes of lower frequency. Diet: Invertebrates and fruit; not known to eat gum Karyotype: 36 chromosomes; fundamental number = 68 Small Galagos: < 125 g

Species: "Galago" thomasi Elliot, 1907 Common Name: Thomas' galago

75

Synopsis of Galago Species Characteristics

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Fig. 10. Range of Thomas' galago.

Type Locality: Fort Beni, Semliki River, Zaire Taxonomic Note: Since its description in 1907, this taxon has been almost universally interpretated as a subspecies of Demidoff's galago. Olson (1979) concluded that Thomas' galago is a separate species based on its morphology, larger size, and sympatric occurrence with Demidoff's galago at a number of circum Zaire Basin localities. Geographic Range (Fig. 10): Extremely disjunctive, restricted to montane and highland regions around the Zaire Basin. Know to occur at Mr. Cameroun, in the Loanda Highlands of Angola, in the region between Dilolo and Kolwezi in southern Zaire, in the Kivu and Ituri regions in eastern Zaire, and in southwest Uganda; possibly at Mt. Marsabit. Habitat Preferences: Relic primary forest Sympatric Associations: Found with Demidoff's galago over the entire range except the Angolan Highlands; almost certainly sympatric with Allen's and the elegant galago on Mt. Cameroun, with Matschie's galago in western Zaire and Uganda, and along forest margins, with the mohol and large-eared greater galago in the Kivu region of Zaire. Characteristics: Small size but larger than Demidoff's galago; head narrow with pointed muzzle; projecting premaxillae; dorsal pelage blackish-brown; ventrum yellowish; flanks of limbs similar to dorsum; both interocular and circumocular markings prominent (Fig. 12j); tail not bushy.

76

Nash, Bearder, and Olson

demidoff

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Fig. 11. Range of Demidoff's galago.

Body Size: Heat and body length (mm): 123-166 (mean = 146) 150-233 (mean = 261) Tall length (mm): Ear height (ram): 23-33 (mean = 37) Hindfoot length (mm): 39-58 (mean = 69) Body weight (g): 55-149 (mean = 99) Locomotion: Running, leaping, bipedal hopping (Walker, 1979) Gestation Length: ? Litter Size: ? Penile Morphology: ? Advertisement Call: ? Diet: ? Karyotype: Chromosomes ? fundamental number = ? Species: "Galago" demidoff Fischer, 1806 Common Name: Demidoff's or dwarf galago Type Locality: West Africa Taxonomic Note: Originally given the name Galago demidoff by Fischer in 1806. Two years later, Fischer redescribed his type specimen as "Galago demidovii." Many authorities have recognized the primacy of Fischer's 1806 work, which accounts for the frequent identification of

Synopsis of Galago Species Characteristics

77

this species as "demidoffi"or "demidoffii."Nomenclatorial rules, however, are clear that the original 1806 spelling is correct (Olson, 1986) and that demidoff is the senior available name for this taxon. Geographic Range (Fig. 11): West and Central Africa; F e r n a n d o Po. In East Africa it occurs in Uganda northwest o f Lake Victoria and in Tanzania in the Uluguru Mountains. Habitat Preferences: Primary and secondary forest Sympatric Associations: On Fernando P o and most o f the Camerouns and Gabon it is found with Allen's and the elegant galago; on Mt. Cameroun, with Thomas' galago. Along the eastern edge o f its range, it occurs with

Fig. 12. Facial views of (a) the large-eared greater galago, (b) Garnett's galago, (c) Allen's galago, (d) the elegant galago, (e) Matschie's galago, (f) the Senegal galago, (g) the Somali galago, (h) the mohol galago, (i) the Zanzibar galago, (j) Thomas galago, and (k) Demidoff's galago. All drawn to the same scale. Figure prepared by D. Eden.

78

Nash, Bearder, and Olson

Matschie's and Thomas' galago and possibly with the m o h o l galago in forest margins south o f Kivu Province in Zaire. Characteristics: Smallest galago; head narrow with pointed upturned muzzle; projecting premaxillae; ears small; dorsal pelage rufus to reddish-brown; ventrum yellow; flanks o f limbs similar to dorsum; interocular strip prominent; circumocular markings variable (Fig. 12k); tail not bushy. B o d y Size: (mean = 129) Head and body length (mm): 73-155 (mean = 179) Tail length (ram): 110-215 (mean --- 24) Ear height (ram): 14-35 (mean = 46) H i n d f o o t length (ram): 35-60 (mean = 70) Body weight (g): 44-97 L o c o m o t i o n : Quadrupedal running and jumping; no bepedal hopping; unable to land hindfeet first when jumping. G e s t a t i o n L e n g t h : 111-114 days L i t t e r Size: 1 normally, 2 maximum P e n i l e M o r p h o l o g y : Penile spines unidentate A d v e r t i s e m e n t Call: A sequence o f 15-20 units organized as a crescendo which usually lasts 2-4 sec. The units increase gradually in pitch and speed o f repetition, culminating in a very rapid, high-pitched, final section ["gathering" call (Charles-Dominique, 1977)]. D i e t : Invertebrates, fruits, and gum K a r y o t y p e : 58 chromosomes; fundamental n u m b e r = ?

REFERENCES Bearder, S. K. (1987). Lorises, bushbabies and tarsiers: Diverse societies in solitary foragers.

In Smuts, B. R., Cheney, D. L., Seyfarth, R. M., Wrangham, R. W., and Struhsaker, T. T. (eds.), Primate Societies, University of Chicago Press, Chicago, pp. II-24. Charles-Dominique, P. (1977). Ecology and Behaviour o f Nocturnal Primates, Duckworth, London. Coquerel, C. (1859). Notes de mammalogie. Rev. Mag. Zool. (2)11: 457-460. De Boer, L. E. M. (1973). Cytotaxonomyof the Lorisoidea (Primates: Prosimii). I. Chromosome studies and karyological relationships in the Galagidae. Genetica 44: 155-193. Dixson, A. F., and Van Horn, R. N. (1977). Comparativestudies of morphologyand reproduction in two subspecies of the greater bushbaby, Galago crassicaudatus r and G. c. argentatus. J. Zool. Lond. 183: 517-526. Drake-Brockman, R. E. (1910). The Mammals o f Somaliland, Hurst and Blackett, London. Elliot, D. G. (1907). Descriptions of apparently new species and subspecies of mammals belonging to the families Lemuridae, Cebidae, Callithrichidae and Cercopithecidae in the collection of the Natural History Museum. Ann. Mag. Nat. Hist. 20: 185-196. Fischer yon Waldheim, O. (1806). Nouvelle esp~ces d'animaux qui se trouvent au Mus6um Imperial d'Historia Naturelle. Mdm. Soc. Nat. Moscow 1(2): 23-26. Fischer yon Waldheim, G. (1808). Galago Demidovii, nova species Quadrimanorum, observatis anatomicis IUustrata. Comment. Sor Phys.-Med. Moscow 1: 57-79.

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Geoffroy Saint-Hilaire, E. (1796). M~moire sur les rapports naturals des makis Lemur, L. et description d'une esp~ce nouvelle de Mamrnifbres. Magasin Encycl. 1(2): 20-50. Geoffroy Salnt-Hilaire, E. (1812). Note sur trois dessins de commer~on, representant des quadrumanes d'un genre inconnu. Ann. Mus. Hist. Nat. Paris 19: 171-175. Gray, J. A. (1863). Revision of the species of lemuroid animals, with the description of some new species. Proc. zool. Soc. Lond. 129-152. Gray, J. A. (1873). Notes on Propithecus, Indris and other lemurs (Lemurina) in the British Museum. Proc. zool. Soc. Lond. 846-860. Groves, C. P. (1974). Taxonomy and phylogeny of prosimians. In Martin, R, D., Doyle, G. A., and Walker, A. C. (eds.), Prosimian Biology, Academic Press, London, pp. 449-473. Harcourt, C. S. (1980). Behavioural Adaptations in South African Galagos, M. S. dissertation, University of the Witwatersrand, Johannesburg. Harcourt, C. S., and Nash, L. T. (1986). Social organization of galagos in Kenyan coastal forest. L Galago zanzibaricus. Am. J. PrimatoL 10: 339-355. Hill, W. C. O. (1953). Primates: Comparative Anatomy and Taxonomy, VoL L Strepshirhini, University of Edinburgh Press, Edinburgh. Jouffroy, F. K., and Lessertisseur, J. (1979). Relationships between limb morphology and locomotor adaptations among prosimians: An osteometric study. In Morbeck, M. E., Preuschoft, H., and Gomberg, N. (eds.), Environment, Behavior and Morphology, Gustav Fischer, New York, pp. 143-181. Kingdon, J. (1971). East African Mammals, Vol. I, Academic Press, London. Le Conte, J. 1857. Description of several new mammals from Western Africa. Proc. Acad. Nat. Sci. Phil. 9: 10-11. L6nnberg, E. (1913). Mammals collected by the Swedish Zoological Expedition to East Africa. K. Svenska Vetensk. Akad. Handl. 48(5): 1-188. Lorenz, yon Liburnau, L. (1917). Beitrag zur Kenntnis der Affen und Halbaffen yon Zentralafrika. Ann. Nat. g u s . Wien 31: 169-241. Masters, J. C. (1985). Species Within the Taxon Galago crassieaudatus E. Geoffroy, Ph.D. thesis, Univeristy of the Witwatersrand, Johannesburg. Matschie, P. (1892). Uber anscheinend neue africanische Saugethiere. Sber. Ges. Naturf. Freunde Berl. 4: 107-114. Napier, J. R., and Napier, P. H. (1967). A Handbook o f Living Primates, Academic Press, London. Nash, L. T., and Harcourt, C. S. (1986). Social organization of galagos in Kenyan coastal forest. If. Galago garnettii. Am. J. Primatol. 10: 357-369. Ogilby, W. (1838). On a new species of galago. Proc. Zool. Soc. Lond. 846-860. Olson, T. R. (1979). Studies on Aspects o f the Morphology and Systematics o f the Genus Otolemur, 1859 (Primates: Galagidae), Ph.D. thesis, University of London, London (University Microfilms No. 79-70, 038, Ann Arbor, Mich.). Olson, T. R. (1980). The identity of Galago crassicaudatus E. Geoffroy, 1812: A proposal for the designation of a neotype. Bull. ZooL Nora. 37: 176-185. Olson, T. R. (1986). Species diversity and zoogeography in the Galagidae. PrimateRep. 14: 213. Petter, J. J., and Petter-Rousseaux, A. (1979). Classification of the Prosimians. In Doyle, G. A., and Martin, R. D. (eds.), The Study o f Prosimian Behavior, Academic Press, New York, pp. 1-44. Schwartz, J. (1974). Dental Development and Eruption in the Prosimians and Its Bearing on Their Evolution, Ph.D. thesis, Columbia University, New York. Schwarz, E. (1930). Ein neuer Galago yore Albert-See. Rev. ZooL Bot. Air. 19: 391-392. Schwarz, E. (1931). On the African long-tailed lemurs or galagos. Ann. Mag. Nat. Hist. 7: 41-66. Smith, A. (1833). African zoology. 1. Mammalia. S. Aft. Q. J. 2: 1-136. Smith, A. (1836). Report Expedition for Exploring Central Africa, from Cape o f Good Hope, June 23, 1834, Under the Superintendance o f Dr. A. Smith, Government Gazette Office, Cape Town. Thomas, M. R. Oldfield (1901). Some new African bats (including one from the Azores) and a new galago. Ann. Mag. Nat. Hist. 8: 27-34.

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Thomas, M. R. Oldfield (1917). The geographical races of Galago crassicaudatus. Ann. Mag. Nat. Hist. 20: 47-50. Walker, A. C. (1969). Prosimian locomotor behavior. In Doyle, G. A., and Martin, R. D. (eds.), The Study of Prosimian Behavior, Academic Press, New York, pp. 543-565. Waterhouse, G. R. (1838). Characters of a new galago (Galago alleni) and a new Pteromys (P. horsfieldii), in the Society's collection. Proc. zooL Soc. Lond. 87-88.