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Locomotor diversity in prosimian primates.

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American Journal of Primatology 13:271-281(1987)
Locomotor Diversity in Prosimian Primates
DANIEL L. GEBO
Department of Anthropology, Duke University, Durham, North Carolina
An understanding of prosimian movement is basic to many anatomical and
paleontological studies in that these studies attempt to correlate movement
with anatomy and therefore infer movement in fossil primates. Duke University has a large and diverse collection of prosimian primates, which are
housed in cages and enclosures large enough for movement studies. Extant
prosimians move in many different ways, and none are so specialized that
only one mode of travel is used. The most general locomotor patterns are
observed for the cheirogaleids, and thus theirs may best resemble the
locomotor patterns of the ancestral euprimate.
Key words: captive setting, cheirogaleids, lemurids, lorisids
INTRODUCTION
The movement patterns of prosimian primates are inadequately known at
present to provide well established correlations between anatomy and behavior.
These correlations are especially important to paleontologists, who must infer movement from anatomy and on this basis try to reconstruct the movement patterns of
extinct primates. Thus any information concerning prosimian movement activities
is helpful, even data from a captive setting. The Duke Primate Center (DPC) houses
20 species of prosimian primates, of which 17 are housed in cages with a diversity of
support types of varying diameters and large enough for movement studies. These
caging facilities have largely diminished the problems of space for movement studies. This paper provides quantitative data on prosimian locomotion gathered at the
Duke Primate Center.
MATERIALS AND METHODS
The biggest problem in studying prosimian primates is related to the access and
use of these rare materials. The DPC has a large and diverse collection of animals,
skeletons, and cadaver specimens, yet, even with the 20 living prosimian species
present at the DPC (Table 0, another 15 species exist in the wild and remain largely
unstudied.
Seventeen of the 20 species at the DPC were observed, and movement was
recorded in terms of frequency of locomotor bouts. Caging was judged t o be appropriate for movement studies after consideration of the following variables: 1) the sue of
Received July 3,1986; revision accepted April 6, 1987.
Address reprint requests to Daniel L. Gebo, Department of Cell Biology and Anatomy, "he Johns Hopkins
University School of Medicine, 725 North Wolfe Street, Baltimore, MD 21205.
0 1987 Alan R. Liss, Inc.
272 I Gebo
TABLE I. Prosimian Primates at the Duke Primate Center
Lemuridae
1. Lemur catta
2. Lemur fulvus
3. Lemur macaco
4. Lemur mongoz
5. Lemur rubriventer
6. Lemur coronatus
7. Varecia variegata
8. Hapalemur griseus
Indriidae
9. Propithecus verreauxi
Cheirogaleidae
10. Cheirogaleus medius
11. Cheirogaleus major
12. Microcebus murinus
13. Mirza coquereli
Lorisidae
14. Loris tardigradus
15. Nycticebus coucang
16. Perodicticus potto
17. Galago senegalensis
18. Galago garnettii
19. Galago crassicaudatus
20. Galago demidovii
the animal, 2) the number of animals within the cage, 3) the arrangement of
supports within the cage, and 4)the amount of space for freedom of movement. The
types of caging included two outdoor enclosures; large, silolike cages; large rectangular cages; tall indoor, triangular cages; and indoor rectangular cages. All cages
contained from fifteen to hundreds of supports varying in diameter from 2-5 mm to
over 50 cm in the outdoor enclosures. Each cage had a variety of horizontal, oblique,
or vertical supports arranged for movement diversity. Table I1 lists the cage sizes for
the species under observation.
Observations were recorded on focal animals when the animals were active and
moving, following the technique of Fleagle [1976, 19781, Fleagle and Mittermeier
[1980], and Mittermeier [1978]. Either early morning or late afternoon observation
periods were used for the diurnal prosimians, and nocturnal prosimians were observed during evenings according to their artificial daylnight light schedule. Only
adults were observed, and a minimum of 500 locomotor bouts were recorded for all
but Cheirogaleus major. A locomotor bout was defined as a movement sequence for
one type of movement. For example, if an animal climbed down a pole, leaped to
another pole, and then climbed upward, the sequence of movement would be scored
as three different locomotor bouts, two for climbing and one for leaping. A switch in
the type of movement, eg, changing from a climb to a leap, started a new bout, and
each bout continued until the animal stopped moving or changes its type of movement. If an animal stopped moving for 1 min or longer, attention was switched to
another animal. Switching was common during any one observation period. Normally, only a few animals inhabited a cage, and all were sampled many times.
Prosimian Locomotion I 273
TABLE 11. Cage Sizes for Species That Were Observed for Movement
Frequencies
Species
Cage size
(length x width x height)
Lemur catta
Lemur fulvus
Lemur mongoz
Lemur coronatus
Lemur macaco
Varecia variegata rubra
Varecia variegata variegata
Hapalemur griseus
Propithecus verreauxi
Cheirogaleus medius
Cheirogaleus major
Microcebus murinus
Mirza coquereli
Loris tardigradus
Nycticebus coucang
Perodicticus potto
Galago senegalensis
Galago demidovii
0.43 ha enclosure
0.43 ha enclosure
7.6 x 2.4 x 6.6m
3.04 ha enclosure
6.6 x 1.2 x 3.6 m
3.1 x 3.1 x 8.2 m
3.04 ha enclosure
7.6 x 2.4 x 8.7 m
8.7 x 5.5 x 7.6 m
2.4 x 1.5 x 2.1 m
1.8 x 1.8 x 2.1 m
2.4 x 1.5 x 2.1 m
1.8 x 1.2 x 3.6 m
1.8 x 1.2 x 3.6 m
2.4 x 1.8 x 2.1 m
2.4 x 1.8 x 2.1 m
2.1 x 1.8 x 2.1 m
1.8 x 1.2 x 3.6 m
Movement frequencies were recorded for 17 species. Definitions for the movements are as follows: Quadrupedalism A movement in which all four limbs move in
a regular pattern above a horizontal support or on the ground; walking, running,
and galloping were all recorded as a quadrupedalism; leaping: A movement in which
the hindlimbs propel an animal across a gap between two supports; dropping down
from a branch above was not scored as a leap; climbing: A movement up or down a
vertical or steeply inclined support; all four limbs move in an irregular pattern, with
variable hand and foot positions; suspensions: A movement or a posture in which
the body is held below a support by one, two, three, or four limbs in tension;
cantilevering is a special type of suspension in which two feet hold onto a vertical
support and the body is unfolded horizontally outward like a flag (Fig. 1);bridging:
A movement in which gaps are crossed by body stretching; first, the hands stretch
out and grab the distant support and then the body is stretched across; the two feet
are then released in succession, and the animal moves across to the new support;
bimanual movements: A movement in which only the hands grasp a support and are
used to pull the body up to a support from below, to hang below a support, or for
arm-swinging; bipedal movements: A movement in which only the hind feet are used
to raise an animal up, to hop, or to take a short walk.
To check the effect of differences between groups, three different silolike cages
of the same size were chosen, each cage containing a different group of Varecia
uariegata rubra. Each cage had a different arrangement of supports, but all three
cages were similar in the number, the size, and the orientation of the supports.
Table 111, which lists the movement frequencies for each cage, shows that the
greatest difference occurred in frequency of leaping; this differed by 6% from the
lowest to the highest value. Suspensory movements differed by 5% and quadrupedalism by 3%. By averaging the SD values for the four most frequent movements,
274 I Gebo
Fig. 1. Mirzu coguereli cantilevering outward from a vertical support. The arrow on the support points
upward and thus orients the support relative to the horizontally positioned Mirzu.
TABLE 111. Movement Frequencies for Vureicu vuriegutu rubm in Three Separate Silolike
Cages (1-111)
Quadrupedalism
Leaping
Climbing
Suspension
Bridging
Bimanual
Bipedal
Locomotor bouts
I (%I
I1 (%)
III (%)
Mean (%)
SD (%)
32
27
23
14
1
2
1
612
35
21
24
17
1
2
<1
607
33
27
24
12
2
<1
1
610
33.3
25.0
23.7
14.3
1.3
1.7
1.o
1.53
3.46
0.54
2.32
0.58
0.58
0.00
quadrupedalism,leaping, climbing, and suspension, a value of 1.96 was determined,
and, by doubling this value, an approximate error factor of +4% for 95% of the
variates was determined. This approximate error factor can be used as a measure of
comparison between species for the more frequent movement values listed in Table IV.
RESULTS
Lemuridae
Observations on Lemur cuttu in a 0.43 ha enclosure at the DPC revealed the
terrestrial preference of this species (Table v>. Table 4 lists L cuttu’s movement
frequencies. L. cuttu moved quadrupedally more than any other lemurid and had a
high percentage of bipedalism, consisting of hopping or walking when on the ground.
Like all lemurs, they leaped, climbed, and suspended themselves below supports
using all possible variations of their four limbs. L cattu climbed large-diameter trees
and branches and usually leapt down from branch to branch, as Sussman 119741 has
Prosimian Locomotion I 275
TABLE IV. Movement Frequencies for Prosimian Primates at the Duke Primate Center*
&(%I
L(%) CL(%) S(%) BR(%) BIM(%) BIP(%) Bouts
Lemur catta
51
22
10
1
4
<1
7
642
Lemur fulvus
39
34
17
1
9
<1
<I
1,273
Lemur mongoz
29
37
22
2
8
1
1
2,164
Lemur coronatus
33
30
31
2
1
3
<1
842
Lemur macaco
30
31
28
1
6
4
1
525
Varecia variegata
34
25
23
2
14
1
1
1,829
rubra
Varecia variegata
35
21
31
11
1
1
1
525
variegata
Hapalemur griseus
24
56
15
7
1
1
<1
2,118
Propithecus verreauxi
6
46
30
0
5
10
3
2,149
Cheirogaleus medius
34
21
28
5
11”
<1
1
2,131
Cheirogaleus major
41
6
39
4
7a
3
<1
340
Microcebus murinus
29
38
24
3
<1
5a
<1
2,149
Mirza coquereli
20
27
24
5
1
1
23a
3,772
Loris tardigradus
15
0
31
1
30a
24
<1
2,496
Nycticebus coucang
0
21
24
23
2ga
2
1
605
Perodicticus p t t o
29
0
31
17
22a
1
<1
645
Galago senegalensis
5
63
26
<1
2
2a
<1
2,245
Galago demidovii
25
40
24
6a
4
<1
1
2,163
*Abbreviations: Q, quadrupedalism; L, leaping; CL, climbing; S, suspension;BR, bridging; BIM,bimanual;
BE’, bipedal; Bouts, locomotor bouts.
‘Species that perform cantilevering.
reported from his observations of L. catta in the wild. L catta also engaged in
bridging and bimanual movements at the DPC, although very rarely. Like all other
lemurids, they were never observed to perform a cantilever movement.
L fulvus leaped, climbed, and engaged in suspensory movements and postures
more often than did L catta (Table IV). Although very arboreal, L fulvus often
moved on the ground in the outdoor enclosures without apparent difficulty. A
comparison of arboreal movements and ground movements between L fulvus and L
catta is given in Table V. L filvus used quadrupedalism and suspension in the
arboreal setting far more often than did L catta, which tended to leap and climb
when in trees. L catta moved using bipedal hops and quadrupedal walking movements when on the ground more often than did L. fulvus.
L. mongoz leaped and climbed more than L fulvus (Table IV). Frequency of
suspensory movements was about equal for the two. L fulvus was more quadrupedal
than L. mongoz. L mongoz climbed downward tail first but would on occasion climb
downward head first. L. mongoz would also drop head first to a branch below while
hanging by two feet.
I, macaco, a generalist among the lemurs, used quadrupedalism, leaping, and
climbing in about equal proportions (Table IV). L macaco commonly leaped from
vertical support to vertical support with short climbing sequences interposed between leaps. The climbing sequences tended to be over very short distances as
compared to those of Varecia. They used their arms significantly more often than
did other lemurs. Bimanual “raise-ups” and short arm-swings were fairly common.
L coronatus was similar to L. catta in movement frequencies in that it was less
suspensory than the other lemurs and used bipedalism more often. L. coronatus was
a generalist compared to L. catta in that quadrupedalism, leaping, and climbing
276 I Gebo
TABLE V. Arboreal vs Terrestrial Movement Frequencies for Lemur cutta
and Lemur fulvus in a 0.43 ha Enclosure*
Lemur fuluus (%)
Lemur cuttu (%)
Arboreal
Q
L
CL
S
BIP
16
44
26
13
0.1
5
55
38
86
13
0
70
13
0
0
0
18
1
1
Terrestrial
Q
L
CL
S
BIP
BOUTS
1
1,273
68% arboreal
642
22%arboreal
*For explanation of the abbreviations,see footnote t o Table 4.
were used in about equal amounts (Table IV). Although L. coronatus and L macmo
were both generalists in terms of movement frequencies, L. coronatus preferred
quadrupedalism to leaping, whereas L. macaco leaped more and made more frequent
use of suspension and bimanual movements.
Varecia variegata, the largest lemurid, leaped with the same frequency as L.
catta but less often than the other lemurs (Table IV). Varecia engaged in suspensory
movements more often than the other lemurids. Hanging by two feet was common.
Varecia, without hesitation, would let go while hanging by two feet and drop to the
branch below. Varecia most frequently engaged in above-branch quadrupedalism
(Table IV).
Observations on the black-and-white ruffed lemur, Varecia uariegata uariegata,
in the 3.04 ha enclosure showed movement frequencies that were similar to those of
the red ruffed lemur, Varecia uariegata rubra, which was observed in three separate
silolike cages. All movement frequencies for these two subspecies were within 4% of
one another, with the exception of climbing. The ruffed lemurs in the outdoor
enclosure climbed more frequently than the ones in the silolike cages. The much
taller trees in the enclosure are the likely explanation for this difference in climbing
frequencies.
Hapalemur griseus was far less quadrupedal than the other lemurs and leaped
far more frequently (Table IV).Hapalemur climbed and engaged in suspension about
as often as L. fulvus. Hapalemur clung vertically; hopped bipedally; dropped downward head first while hanging from two feet; suspended themselves by any combination of four or fewer limbs, including one hand, although rarely; and occasionally
bridged across two supports. Hapalemur hopped or made short leaps upward or
downward to a branch more often than they climbed to that spot. They climbed
downward tail first, as do all lemurids, and would on occasion make bouncy leaps
with all four limbs, perform bipedal hops, or rise up on two legs.
Indriidae
For Propithecus verreauxi, leaping was the most frequent locomotor movement
observed at the DPC (Table IV). These indriids never engaged in bridging move-
Prosimian Locomotion I 277
ments, nor did they use a hanging rope at the DPC. They were observed to support
themselves between two vertical supports with one hand and one foot on different
supports. I! verreauxi showed a high frequency of climbing, as do the large-bodied
lorisines (Table IV).I! verreauxi climbed downward tail first like lemurids, however,
rather than descending head first as has been observed in lorisines. Quadrupedalism
and suspension were not common. I! verreauxi did engage in a significant amount
of bimanual movements, including brachiation, although for short periods of time.
The more frequent use of bimanual movements by I! verreuuxi was very unusual
compared to other prosimians in this study.
Cheirogalidae
Cheirogaleus medius moved in a scurrying quadrupedal manner, with its body
held close to the support. Quadrupedalism and climbing were the two most frequent
movements (Table IV).C. medius preferred horizontal supports and performed short,
cautious leaps. They often leaped up to a support, grabbed on using a four-limb
suspensory posture, and pulled themselves up on top of the support. Cheirogaleids
were, in general, more suspensory in their movement preferences, climbed more
often, and bridged more frequently than lemurids (Table IV). Cheirogaleids climbed
downward head first. Suspensory movements included all possible hand and foot
combinations. All cheirogaleids at the DPC performed the cantilevering movement
(Fig. 1). To perform this suspensory movement, cheirogaleids move their feet to the
same side of the vertical support and spread their legs outward for a wider base of
support. The body is fully extended horizontally, including the knees. "his leaves
only the feet to hold onto the vertical support and leaves the hands free to grab
insects. The whole movement of unfolding the body outward is similar to an accordian in the pattern of movement. This insect-catching movement was first noted by
Martin [1972] for Microcebus.
C. major, which is larger than C. medius, preferred quadrupedalism and climbing over all other movements (Table IV). C. major moved using quadrupedalism
more frequently than any other cheirogaleid. C. major was less suspensory and
leaped less frequently than C. medius (Table N).
Mirza coquereli was the most suspensory of the cheirogaleids, being just below
the lorisines in frequency (Table IV). Mirzu was the most generalized, movementwise, of the cheirogaleid species, with quadrupedalism, leaping, climbing, and suspension frequencies being very similar (Table IV).Mirzu preferred short leaps and
would size-up large leaps before committing. Bipedal raise-ups were performed with
bent knees.
Microcebus murinus was the most frequent leaper of the cheirogaleids (Table
W. Microcebus made quick, agile movements, and quadrupedalism and climbing
were relatively frequent. Suspensory movements and postures were less frequent
for Microcebus than for Mirza. Microcebus also contrasted with Mirzu in that quadrupedalism and leaping frequencies were higher (Table IV).On the ground, Micrccebus moved by quadrupedalism or by hopping with all four limbs. Microcebus
bridged to other supports, especially to branches above themselves. Microcebus
dropped to lower branches if the distance was short. Microcebus, Mirza, and Cheiroguleus all clung vertically and leaped between vertical supports.
Lorisinae
Lorisines moved with more deliberate movements than did the cheirogaleids.
They bridged more often and were the most suspensory of all prosimians (Table IV).
Loris turdigradus, a small-bodied lorisine, preferred climbing and suspensory movements over quadrupedalism and bridging. Like the other lorisines, it can cantilever.
278 I Gebo
Lorisines, however, do not always place their feet on the same side of the vertical
support as do cheirogaleids. Loris was capable of very rapid movements when
moving quadrupedally. In this study, Loris was similar to Perodicticus in climbing
frequency but was most similar to Nycticebus in suspension and bridging frequencies
(Table IV). Loris sometimes climbed vertical supports backward while holding the
body vertical and while grasping the support with only two feet. These animals
occasionally moved on the ground, but this rarely occurred.
Of all lorisines at the DPC, Perodicticus was quadrupedal most frequently (Table
IV). Perodicticus moved quadrupedally and climbed more frequently than did the
other large-bodied lorisine, Nycticebus. Perodicticus bridged between supports in the
small branch periphery. When on the ground, Perodicticus used quadrupedalism
that was of a serpentine type and had its feet placed in an everted position.
Nycticebus was more suspensory than Perodicticus (Table IV).The most frequent
movements for Nycticebus were suspension and quadrupedalism, followed by bridging and climbing, all of which ranked between 20% and 30% (Table IV). Like
Perodicticus, Nycticebus moved using a serpentine style of quadrupedalism when on
the ground, lifting each foot in a stereotypic sequence that contrasted with the more
fluid motion of Perodicticus. First, the foot was lifted off the ground and quickly
inverted as it moved forward; it was then everted and abducted just before it was
placed on the ground. Nycticebus and Perodicticus moved on the ground rarely but
more frequently than did Loris.
GaIaginae
Table IV shows that Galago demidovii was predominantly a leaping species,
with quadrupedalism and climbing being the next two most frequent movements.
Quick rather than deliberate quadrupedal movements were made by G. demidovii.
G. demidovii was more suspensory than G. senegalensis (Table IV). G. demidovii
climbed downward head first, performed the cantilevering suspensory movement,
and moved on the ground using quadrupedalism or by bipedal hops.
Long, fast leaps from vertical supports were the most common movement for G.
senegalensis. Climbing was the only other movement. G. senegalensis performed
with any frequency above 5% (Table N).
G. senegalensis normally climbed downward head first but would on occasion climb downward tail first. G. senegalensis
preferred to hop bipedally on the ground. Suspensory movements were not common,
although G. senegalensis did cantilever.
DISCUSSION
Historically, primate movements have been categorized into locomotor types
[Mollison, 1910; Erickson, 1963; Ashton & Oxnard, 1964; Walker, 1967; Napier &
Walker, 19671. Stern and Oxnard [19731 reviewed the implications of categorization,
suggesting that categories should be an aid to thinking, not a hindrance, and noting
that field research has established that greater variation exists for a variety of
movements among primates than was previously thought. Stern and Oxnard [1973]
concluded that categories no longer serve a useful purpose. General locomotor
categories are important to paleontologists, however, who must infer broad movement patterns from anatomy alone.
Further field research has suggested that prosimian body postures and movements follow broad phylogenetic patterns but with some interesting exceptions. For
example, most lemurs move in ways that distinguish them from cheirogaleids or
indriids, yet the frequency of specific movements is related not only to which lemur
is being considered but also to forest structure, where plant types and density,
support diameters, and support angles may vary. This is especially true for support
Prosimian Locomotion I 279
diameters and angles, since foot and body size largely determine the size of arboreal
supports a prosimian will use, with some qualifications. In some prosimians, nails
have been modified into “c1aws7’[Charles-Dominique, 19771 or the big toe has been
shifted into a more abducted position (ie, lorisines); such modifications have allowed
prosimians to use larger-diameter supports relative to foot size.
The angle of a support on which a prosimian chooses to move is governed by a
different set of conditions related to forest structure. These conditions are the type
and sizes of trees in a forest, the distribution of food resources in the trees, and the
size of the animal relative to the branches. Some species (eg, Lemur catta) have been
documented to use forest structure differently in different forests [Jolly, 1966; Sussman, 1972; 1974; Budnitz & Dainis, 1975; Tattersall, 1977; Richard, 19781.
The quantitative results of this captive study allow several simple observations
to be made concerning prosimian locomotion. First, all prosimians climb. Second, all
prosimians move using quadrupedalism, although this movement was rare in G.
senegalensis and I? uerreauxi. Third, lorisines never leap. Fourth, only cheirogaleids
and lorisids perform cantilevering movements. Last, all prosimians use the posture
of vertical clinging, which in reality is nothing more than a stopping position in a
vertical climbing sequence. Of course, certain prosimians (eg, G. senegalensis) cling
on vertical supports more frequently than others and can maintain this position for
long periods of time. Thus the posture vertical clinging differs from the movement
vertical climbing only in the amount of time delegated toward maintaining this
position.
The most important result of this study is that when prosimians are given
adequate space and support diversity, data from captive locomotor studies can agree
quite well with field observations and thus add to our understanding of locomotor
diversity (see, eg, the locomotor descriptions provided by Walker [19741, CharlesDominique [1977], and Tattersall [1982]). When quantitative data are known, for
example, in galagines studied by Walker [1979] and Crompton [1984], these data
also compare well, although some discrepancies are apparent (Table Vn.
CONCLUSIONS
1. Species of extant prosimians move in many different ways, but only a few
movements are preferred; these distinctions tend to be highly informative in a
comparative approach. Although data from captive locomotor studies are not the
final word toward understanding prosimian locomotor adaptations, they agree quite
well with results from previous field studies and should be of value to comparative
anatomists and paleontologists.
2. Lemurid species move in many different ways, whereas indriid species are
less diverse.
3. Cheirogaleids are generalists, but there are interspecific differences in the
amount of leaping and suspensory movements.
4.Lorisines are like indriids in that all species move in similar ways, although
quadrupedalism, suspension, and bridging frequencies do show species-specific
usages.
5. Galagine species differ along two movement gradients, one for leaping and
the other for quadrupedalism.
6. The most generalized locomotor patterns are observed for the cheirogaleids,
which are active in all four major locomotor categories: quadrupedalism, leaping,
climbing, and suspensory movements. Thus the locomotor patterns of cheirogaleids
like Mirza coquereli or Cheirogaleus medius, which are the most generalized, movementwise, of all cheirogaleids, might best resemble the locomotor patterns of the
ancestral euprimate.
25
40
24
6
4
<1
1
5
63
26
2
<1
<1
2
3.41
-
24.11
45.30
25.86
1.82
Walker
(G. demidovii)
21
53
22
4
1
5
40
23
34
2
7
-
Crompton
G. senegalensis
G. crassicaudatus
*Walker’s[1979]and Crompton’s [1984] data have been modified to be comparablewith my own observations.
Quadrupedalism
Leaping
Climbing
Suspension
Bridging
Bimanual
Bipedal
This study
G. demidovii
G. senegalensis
TABLE VI. Movement Freauencies (%) for Several Soecies of Guluao*
Prosimian Locomotion I 281
ACKNOWLEDGMENTS
A special thanks must go to the Director of the Duke Primate Center, Dr. E.L.
Simons, and to the staff at the Duke Primate Center for their help throughout this
project. I would also like to thank D.T. Rasmussen for his drawing of Mirza coquereli
and for his comments on the manuscript.
REFERENCES
Ashton, E.H.; Oxnard C.E. Locomotor patterns in primates. PROCEEDINGS OF THE
ZOOLOGICAL SOCIETY OF LONDON
14211-28, 1964.
Budnitz, N.; Dainis K. Lemur catta: Ecology
and behavior. Pp. 219-235 in LEMUR BIOLOGY. I. Tattersall and R.W. Sussman
eds. New York, Plenum Press, 1975.
Charles-Dominique, P. ECOLOGY AND BEHAVIOR OF NOCTURNAL PRIMATES.
New York: Columbia, 1977.
Crompton, R.H. Foraging habitat structure,
and locomotion in two species of Galago. Pp.
73-111, in ADAPTATIONS FOR FORAGING IN NONHUMAN PRIMATES. P.S.
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