Folia Primatologica Separatum

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Folia Primatol 199 1;57:132-146 © 199 1 S. Karger AG,

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·Patterns of Sexual Dimorphism in Body Weight among

Prosimian Primates
Peter M. Kapp eler
Department of Zoology and Duke University Primate Center, Duke University, Durham, N.C., USA

Key Word s. Sexual dimorphism · Prosimians · Body weight · Evolution · Phylogeny ·

Lemurs · Lorises ·Tarsiers

Abstract.Many primatologists believe that there is no sexual dimorphism in body size

in prosimian primates. Because this belief is based upon data that carne from only a few
species and were largely flawed in sorne aspect of sample quality, 1re-examined the extent
of sexual dimorphism in body weight, using weights of 79 1 adult prosimia ns from 34
taxa recorded over the last 17 years at the Duke Universit y Primate Center. There was no
significant sex difference in body weight in 17 species , but males were significantly
larger in Nycticebus pygma eus, Tarsius syrichta , Galago moholi, Galagoides demidovii,
Otolemur crassicaudatus and Otolemur garnettii. Moreov er, females were significantly
larger in M icrocebus murinus. Thus, the general lack of sexual dimorphism could be
confirmed , notably for lemurs, but prosimians as a group show more variability in sexual
size dimorphism than was previously thought. After including prev iously published data
obtained in the wild from 8 additional species, 1 found significant heterogeneity in the
degree of sexual dimorphism at the family level, but only the lndridae and Galagidae
were significantly different from each other. Among the prosimian infraorders, the
Lorisiformes were significantly more dimorphic than the Lemuriformes. Differences in
dimorphism between higher taxonomic groups are dis cussed in the context of
prosimian evolution, concluding that phylogenetic inertia cannot prov ide a causal
explanation for the evolution of sexual dimorphism. The relative monomor phism of most
prosimia ns may be related to allometric constraints and, especially in the
Lemuriformes, to selective forces affecting male and female behav ioral strategies.

Introduction tive functions , may take several forms in

pri mates [1], but it is most commonly
described
Sexual dimorphism, or morphological reproduc-
differences between males and females
beyond tl1ose related to the basic
in terms of differences in body weight
be tween the sexes [2-4]. The extent of
sexual dimorphism in body weight among
prima tes
Sexual Size Dimorphism in Prosimians
ranges from taxa in which adult males are
nearly twice as heavy as females (e.g. Pap io,
Mandrillus and Pongo), through species with
little or no sexual dimorphism (e.g. Callice
bus ·and Hy lobates) , to those in which
adult . females are slightly larger
than males (e.g. Callithrix and
Saguinus) [5-9]. Alth ough lit tle is known
about sexual dimorphism in
prosimians, many primatologists believe
that this suborder is characterized by a gen
eral absence of sexual dimorphism [see e.g.
1, 9-1 3]. The main objectives of this paper
are to show that this conception is partly
based on a lack of reliable data for most pro
simianspecies, and that a more differen
tiated pattern emerges from a comparative
analysis based on a larger sample.
Body weight data for primates are used
not only in analyses of sexual size dimor
phism, which illuminate sorne of the
effects of natural and sexual selection on
male and female body size [5, 14-17], but
are also cen tral to many allometric [8, 14-
20] and eco
logical studies [21, 22]. New information
on prosimian body weights may theref ore
assist future studies of these relatively
little-stud ied aspects of prosimian biology.
The degree of sexual size dimorph ism of
a species can be determined with reliable
data on male and female body mass.
Despite this simple requirement , however,
'it is not uncommon for studies of sexual
dimor phism to be flawed in one or more
aspects of sample quality' [7]. This is
underscored by the nature of most
published prosimian body weights. Until
very recently, the most extensiv e list of
prosimian weights was that compiled by
Clutton-Brock and Harv ey [9]. For 5 of the
17 prosim ian species for which body
weights were given, howev er, the val ues
represented 'approximate figures taken in
sorne cases from congeneric species of
133
similar size belonging to the same ecological
category', which could not be used to estí
mate the degree of sexual dimorphism. In 2
other cases ( Loris tardigradus and Nyctice
bus coucang) the given weights did not re
present actual measurements, but midpoints
of ranges given in Napier and Napier
[23]. For Galago demidovii and G.
senegalensis, weights reported elsewhere
without distinc tion between the sexes [24,
25] were as signed to both sexes. Data for
5 more spe cies were based on personal
communica tions, so that the sample
statistics remained unknown. Although
Clutton-Brock and Har vey [2] stressed
that 'indiv idual estimates may be
unrealistic and that they should not be used
as a referen ce source', the same data were
used again by them and by other au thors
[4].
Another frequently used list of primate
weights contained only 6 prosimian species
[3, 4, 26, 27]. Although these prosimian
weights were claimed to be derived from 16
or more wild-trapped and wild-shot animals
of each species, in 4 of the 6 species they
were apparently based on measurement of
no more than 6 animals per species [28].
Both Charles-Dominique [29] and Tattersall
[30], on the other hand, published prosimian
body weights based on comparatively large
sample sizes, but , unfortunately, did not
dis tinguish between the sexes. The latest
and most complete list of primate body
weights [31], also used in [8], prov ided
neither refer ences for literature data nor
sample statistics for prev iously unpublished
v alues. The reli ability of the data for 1O
prosimian species in this sample seems at
least questionable be cause identical
figures are given for males and females.
Because the quantity and quality of pre
viously published prosimian body weights
134
are clearly insufficient to permit compre
hensiv e assessment of sexual dimorphism,
I collected body weight data for 25
prosim ian species from a total of 79 1
captive ani mals and combined them with
literature data for 105 wild animals from 8
additional species to determine sex
differences within species, and to
compare patterns of sexual size
dimorphism among higher taxonomic
units.
Materials and Methods
The Duke University Prim ate Center (DUPC) in
Durham , N.C., USA, houses the largest captive
breed ing colony of prosim ian primates in the
world. At present, more than 700 animals from ali
prosim ian families except the Lepilemuridae are
kept for captive breeding and possible
reintroduction to the wild. Dif ferent species are
housed und er different conditions, ranging from
indoor darkrooms to large outdoor en closures. All
animals can be identified indiv idually. A file is kept
for each individual, including the animal's
genealogy , medica!record and reproductive
infor ma tion. Whenever animals undergo regular
veterinary examinatíons, or are handled for other
reasons, body weight is recorded .
1 extracted data for a total of 791 animals of 34
taxa from the DUPC weight records spanning the last
17 years. Only one measurement per month was used.
When an animal was weighed more than once per
month, the mean value was used for further analysis.
A conservative estímate of age at sexual maturity was
used to define adulthood in both sexes [see e.g. 32 and
references therein]. Weights of individuals of noctur
nal species younger than 12 months old were dis
carded, as were weights for individuals of diurna!spe
cies younger than 24 months. Weights of wild-caught
animals obtained within the first 3 months following
arrival at DUPC were also discarded. In the case of
females, a conserv ative estímate of gestation length,
based on data from the literature [32 and references
therein, 33-38] or unpublished data from DUPC, was
determined for each species. Subsequently , using
the reproductiv e information in each individual file,
data from females during pregancy(ies) were also
ex cluded.
Kappeler
A total of 6, 165 values remai ned, and these were
used to calculate a mean weight for each individual.
Mean weights and other sample statistics for both
sexes of each taxon were then determ ined from these
individual means. The degree of sexual dimorphism
for each taxon was calculated as mean male weight
expressed as a percentage of female weight. An exten
sive literature search for information on prosimian
body weights was made. Data from taxa that were not
available at DUPC, and that were based on direct
measu rements , were extracted. Data from different
sources were combined in only 2 cases; in Da uben
tonia madagascariensis in order to increase sample
size, and in Lepil emur rufi caudatus because male
and female weights were not available from a single
source. Whenev er subspecies existed, the weights
from the respectiv e subspecies were combined to
cal culate mean values for the species.
The data from DUPC were subjected to the fol
lowing statistical analyses, using the BIOM
software package [39]. A one-sample Kolmogorov
-Smirnov test for norm ality was performed for each
sex of each species. The homogeneity of varian ces
between the two sexes of each species was tested
with a two-tailed F test. Student's t test was then
used to test for signif icant differences in body
weight between the sexes. All subsequent analyses
were performed at the species level, after including
8 additional species from the lit erature for which
data from at least 2 adult individu als per sex were
available. Thus, the final sample con sisted of 25
'DUPC species' and 8 species whose data carne from
the wild. The variates of the degree of sex ual
dimo1-phism were log-transformed [4] and com
bined in the respective subsamples according to the
taxonomic level of comparison following [40]
and were then subjected to the same tests of
normality and homoscedasticity. A model II nested
ANOVA with unequ al sample size, using
Satterthwaite's approxi mation, was performed to
obtain variance estimates of sexual dimorphism
within and among genera, fam ilies, and
infraorders.
A one-way ANOVA and a subsequent
comparison of means, using the GT2 method, were
performed to compare the degree of sexual
dimorphism among Lemuriformes, Lorisiformes,
and Tarsiiformes. Be cause the variates among
families were heteroscedas tic, the nonparametric
Kruskal-Wallis ANOVA by ranks [41] was used
to evaluate the homogeneity in sexual dimorphism
among families and to perform ali pairwise
comparisons. The null hypothesis in ali tests
Sexual Size Dimorphism in Prosimians 135

was that there was no difference in sexual
dimor phism between the respective subgrou ps. The
signifi cance level in ali tests and the
experimentwise error rate in ali multiple
comparisons were set at alpha = 0.05.
Results
Sexual dimorphism in body weight
among prosimians ranged from species with
significantly larger males through
species
with no significant weight difference be
tween the sexes, to one with significantly
larger females. The majority of species,
nota bly lemurs, exhibited no significant
degree of sexual size dimorphism, howev er.
The mean weights and selected sample
statistics for males and females of 34 taxa,
based on the DUPC records, are presented
in table 1. Data on body weights for 18
additional taxa extracted from primary
references are sum marized in table 2.
None of the Kolmogo-

Table l. Body weights (g) 34 prosim ian at DUPC L

of taxa
Taxon Sex n Mean SE DSD

Lorisiformes
Lorisidae
Loris tardigradus malabaricus m 10 192 7 177 207 99.5
f 8 193 8 175 21 1
Nycticebus pygmaeus m 7 462 17 420 503 122.9
f 5 376 18 325 427
Nycticebus coucang2 m 14 1,207 54 1,091 1,323 1o1.0
f 15 1, 195 61 1,064 1,325
Perodicticus pot to m 5 948 56 793 1,104 95.9
f 6 989 71 804 1, 173

Galagidae
Otolemur crassica udatus m 40 1,495 40 1,415 1,576 120.4
f 44 1,242 34 1,172 1,3 1 1
O. crassicauda tus argenta tus m 9 1,750 47 1,642 1,858 116.9
f 14 1,497 39 1,413 1,581
O. crassicaudatus crassicauda tus m 10 1,226 56 1,099 1,354 109.0
f 8 l , 125 59 986 1,263
O. crassicaudatits monteiri m 21 1,515 41 1,430 1,600 135.0
f 22 1,122 30 1,060 1,184
Otolemur garnettii m 17 1,2 12 48 1,1 1o 1,314 117.9
f 19 1,028 28 968 1,087
Galagoides demidovii m 4 81 5 64 98 1 17.4
f 8 69 2 64 73
Galago moholi m 25 181 4 172 190 1 16.8
f 20 155 4 147 163

(Table 1 continued nex t page.)

136 Kappeler

Table 1(continued)

Taxon Sex n Mean SE DSD

Lemuriformes

Cheirogaleidae
Cheirogaleu majo r m 3 575 73 260 889 129.8
s f 3 443 36 289 598
Cheirogaleus medius m 23 283 8 268 299 100.4
f 25 282 8 266 298
M irza coquereli m 10 307 6 294 320 101.7
f 9 302 14 27 1 333
M icrocebus murinus m 33 90 3 85 96 82.6
f 27 109 4 10 1 117

Lemuridae
H apalemur griseus griseus m 10 939 44 839 1,040 105.3
f 9 892 43 794 990
Lemur catta m 24 2,705 70 2, 560 2,849 1o1 .0
f 16 2,678 71 2, 526 2,830
Lemitr corona tus m 11 1,7 12 80 1,534 1,89 1 101.5
f 8 1,687 88 1,478 1,895
Lemur fulvus m 60 2,39 5 37 2,32 1 2,470 99.9
f 47 2,397 46 2,304 2,489
Lemur fu lvus albifrons m 13 2,390 96 2, 18 1 2, 598 104.8
f 11 2,281 37 2,200 2, 362
Lemur fulvus collaris m 13 2,459 61 2, 327 2, 59 1 98.6
f 4 2,49 5 191 1,885 3, 104
Lemur fu lvus ful vus m 16 2,429 77 2,265 2,593 95.3
f 9 2,550 126 2,259 2,840
Lemur fulvus rufus m 11 2,450 61 2,322 2,597 99.8
f 17 2,456 81 2,284 2,627
Lemur fulvus sanfordi m 7 2, 1 1o 93 1,883 2,337 98.3
f 6 2,147 83 1,934 2,359
Lemur macaco m 23 2,403 61 2,276 2,529 96.6
f 20 2,487 56 2,369 2,604
Lemur macaco fla vifrons m 3 2, 32 1 1 1o 1,850 2,792 101.4
f 5 2,290 92 2,033 2, 546
Lemur macaco macaco m 20 2,41 5 69 2,271 2,559 94.6
f 15 2,552 60 2,423 2,68 1
Lemur mongoz m 14 1,682 73 1,525 1,839 101 .
f 13 1,658 65 1,517 1,800 4
Lemur rubri venter m 4 2,267 124 1,870 2,662 106.0
f 4 2, 139 1 16 1,769 2,510
Sexual Size Dimorphism in Prosimians 137

Table 1 (continued)

Taxon Sex n Mean SE

DSD Lemuridae (continued)

Varecia variega ta m 46 3,47 1 48 3,374 3,568 98.8
f 35 3,512 59 3,393 3,631
Varecia variegata rubra m 20 3,546 73 3,393 3,699 102.1
f 17 3,473 95 3,272 3,675
Varecia variegata variegata m 26 3,414 63 3,284 3,544 96.2
f 18 3,548 72 3,397 3,700

Indridae
Prop ithecus verreauxi m 10 3,637 122 3,362 3,9 13 98.4
f 12 3,696 138 3,391 4,000
Prop ithecus verreauxi coquereli m 8 3,703 134 3,386 4,019 98.6
f 10 3,757 160 3,395 4, 118
Propith ecus verreauxi maj ori m 2 3,376 284 6,978 99.6
f 2 3,389 53 2,7 16 4,063
Prop ithecus tattersalli m 4 3,039 204 2,388 3,689 96.0
f 5 3,167 272 2,412 3,922

Daubentonidae
Daubentonia madagascariensis m 2 2,758 149 865 4, 541 107.2
f 2 2,573 227 5,457

Tarsiiformes
Tarsidae
Tarsius bancanus m 3 128 6 101 155 100.8
f 3 127 5 104 150
Tarsius syrichta m 10 134 3 126 142 114.5
f 17 1 17 2 113 121

1
Arithmetic means for each sex of each taxon were calculated from n individual means. The standard error of
the mean (SE), and the lower (Lt ) and upper (L2) 95 º/o confidence limits around the mean are presented as
well. The degree of sexual dimorphism (DSD) is expressed as the ratio of mean male weight/female weight X
1OO. The values of the members of respective subspecies were combined to obtain means for each sex of a
species if it contained two or more subspecies. The statistics for D. madagascariensis females are based
upon the mean female weight obtained from my own measurements and one datum in the literature [77].
Deviations from Kappeler [42] are due to increased sample size and/or rounding.
2
Combined data for two unidentified subspecies and their hybrids.
1 38 Kappeler

rov -Smirnov tests was significant, i.e. the
variates in ali samples did not depart
signifi cantly from a normal distribution.
More over, the variances of male and
female weights were not significantly
different from each other except in one
species (M irza co quereli: F = 5.28; p
0.05). In Ny cticebus pygmaeus , Tarsius
syrichta and all Galagi dae, males were
significantly larger than fe males. In
M icrocebus murinus, on the other hand,
females were the significantly larger sex.
In all other species, sex differences in
bod y weight were statistically insignificant
(table 3). The average degree of sexual di
morphism was significantly different
among lemurs, lorises, and tarsiers (F =
4.39; p =
0.02). The overall heterogeneity was due to
a significant difference between the
Lemuri formes and Lorisiformes
(MSD 3,3o = 2.92; p 0.05). There was also
significant hetero geneity in the degree of
sexual dimorphism among all eight
prosimian families (KW = 16.0; p 0.05),
but only the difference be tween two of
them (Galagidae and Indridae) was
statistically significant (KW = 20.3; p
0.05; fig. 1). The nested ANOVA dis
closed that most of the interspecific
variation occurred within genera (54.4 °/o)
and among infraorders (26.4 º/o), but very
little among genera ( 15.6 °/o) and families
(6.6 °/o). How ever, this analysis did not
rev ea!significant added variance at any
taxonomic leve!.

Table 2. Summary of selected previously publish ed prosi mian weigh ts 1

Taxon Sex n Mean Range DSD Ref.

Lorisidae
Arctocebus calabarensis m 2 318 1 06.7 78
f 9 298 -
Galagidae
Galago elegantulus 39 300 270-360 - 29
293 223-350 79
Galago matschiei - 210 1 96-225 79
Galago senegalensis - 206 1 12-300 79
Galagoides alleni 17 260 1 88-340 29
- - 314 200-445 79
Galagoides thomasi - 99 55-1 49 - 79
Galagoides zanzibaricus m 12 1 59 1 50-167 1 1 80
f 14 1 36 128-145 6.9
Cheirogaleidae
M icrocebus ruf us m 23 50 35-70 102.0 74
f 5 49 41-63
Phaner f urcif er - 3 460 350-600 81
Allocebus trichotis2 m 2 92 75-98 l 82
f 2 85 78-90 08.2
Sexual Size Dimorphism in Prosimians 139

Table 2 ( continued)

Taxon Sex n Mean Range DSD Ref.

Lemuridae
H apalemur aureus3 m 1 1,245 l 05.9 83
f 1 1,175
m 3 1,613 1,540-1,660 107.5 84
f 1 1,500
Hap alemur simus m 2 2,1 50 165.3 77, 83, 85
f 1 1,300

Indridae
A vahi laniger laniger4 m 4 1,033 83 78.5 84
f 4 1,316 95
Avahi laniger occidentalis m 3 853 97.5 77, 85
f 1 875
Indri indri m 1 4,600 (sick) 73.6 77, 84
f 1 6,250
Propithecus diadema edwardsi 5 m 8 5,633 123 95.5 84
f 6 5,895 152

Daubentonidae
Da ubentonia madagascariensis f 1 2,800 - 77, 85

Lepilemuridae
Lepilemur mustelinus6 m 2 617 103.9 86
f 5 594
Lep ilemur rufica udatus ro 2 764 600-900 90.4 77, 81,
f 4 845 607-995 85

Tarsidae
Tarsius bancanus m 21 128 109.4 87
f 16 117

1
Mean body weights, the lowest and highest reported values (range), and the sample size for these 19 pro
simian species were extracted from the literature. Whenev er data for both sexes were available, the degree
of sexual dimorph ism (DSD) was calculated as in table 1.
2
Not included in analyses in Kappeler [42].
3
Due to the discrepancy between available weights, these 2 samples were not combined, and this species
was not included in the interspecific comparisons, although the DSD is similar in both samples.
4 The value of the eastern subspecies was used in the interspecific comparisons. The standard error of the
mean is presented instead of the range.
5 The standard error of the mean is presented instead of the range.
6 Males and females were confused in Kappeler [42].
140 Kappeler

Table 3. Intraspecific of male and female body weights 1

comparison
Taxon DSD t d.f. p

Lorisidae
Loris tardigradus 99.5 0.10 16 NS
Nycticebus pygmaeus 122.9 3.40 10 < 0.01
Nyticeb us coucang 101.0 0.15 27 NS
Perodicticus p otto 95.9 0.50 9 NS

Galagidae
Galago moholi 116.8 3.74 43 < 0.001
Galagoides demidovi i 1 17.4 2.79 10 < 0.05
Otolemur crassicaudatus 120.4 4.86 82 <
Otolemur ga rnettii 117.9 3.40 34 0.001
< 0.01

Cheirogaleidae
Cheirogaleus maj ar 129.8 2.1 6 4 NS
Cheirogaleus medius 100.4 0.38 46 NS
Mirza coqueref i2 101.7 0.33 17 NS
M icrocebus murinus 82.6 4.06 58 <
0.001
Lemuridae
Hap alemur griseus 105.3 0.76 17 NS
Lemur catta 101 .0 0.04 38 NS
Lemur corona tus 1o1 .5 0.2 1 17 NS
Lemur ful vus 99.9 0.02 105 NS
Lemur macaco 96.6 1.00 41 NS
Lemur mongoz 101.4 0.24 25 NS
Lemur rubriventer 106.0 0.74 6 NS
Varecia variega ta 98.8 0.53 79 NS

Indrid ae
Propithecus verreauxi 98.4 0.30 20 NS
Prop ithecus tattersalli 96.0 0.36 7 NS
Propithecus diadema edwardsi3 95.5 1.35 12 NS
Ava hi laniger laniger4 78.5 2.25 6 NS

Tarsidae
Tarsius bancanus 100.8 0. 14 4 NS
Tarsius syrichta 1 14.5 4.89 25 <
0.001

1 After testing for normality and homoscedasticity, a Student's t test was performed to compare the mean
weights of males and females in each species. The t value, the number of degrees of freedom and the
significance level are presented along with the degree of sexual dimorphism (DSD).
2
Results of a modified t test for unequal variances [39].
3
Raw data from Glander et al. [84]; based on measurements of wild animals. N ote that , compared to
Kappeler [42], sex difference is no longer significant.
4
Raw data from Glander et al. [84]; based on measureme nts of wild animals.
Sexual Size Dimorphism in Prosimians
E 120
(/')
·.!::.
•· . .-:
110 .• . :
- more data become available. This point is
..,_ . .•.-•: .. .
...,,, ' ·' ···· '
- •'. ·•
(ij •
;:'.) 100 '
X
-o
Q)
C/)
Q)
9 sample indicated that females of this species
0
O>
Q)
o
Fig. l. Mean degree of sexual dimorphism in body
weight (male weight/female weight X 100), based on
the respectiv e species means, along with the standard
error of the means for each prosimian family. Ind:
Indridae, Lep: Lepilemuridae, Lem: Lemurid ae,
Che: Cheirogaleidae, Lor: Lorisidae, Dau :
Daubentonidae, Tar: Tarsiidae, Gal: Galagidae.
Discussion
Prosimian primates can now be seen to
exhibit more variety in sexual dimorphism
than was prev iously thought. Recognition of
this is largely due to an increase in sample
size, with respect to both the number of spe
cies and the number of indiv iduals within
species included in this study. Moreover, the
use of carefully sifted data obtained predo m
inantly from a single source has probably
also contribu ted to a more reliable picture of
sexual dimorphism in prosimians.
Although the data presented in this study
carne predominantly from captive animals,
and therefore exceed the weights of wild
individuals to different extents , the degree of
sexual dimorphism is not significantly af
fected by this [Kappeler, in prep.], suggesting
that males and females do not respond dif
ferently to conditions in captivity. Small
sample size for certain species reflect their
141
rarity , both in captivity and in the wild, and
sorne estimates of sexual dimorphism will
hence become more accurate in the future
as
well illustrated by P. diadema edwardsi.
Whereas the preliminary analysis of a small
are significantly larger [42], the sex differ
ence in the present, somewhat larger sample,
is no longer significant.
Although prosimian and anthropoid pri
mates exhibit similar overall variety in the
direction of sexual dimorphism, anthropoids
as a group are nev ertheless significantly
more dimorphic on average [42]. Only in
this sense are prosimians a group with rela
tively little dimorphism. This difference be
tween the two primate suborders may be a
correlate of the marked difference in average
size between them. A positive correlation
between body size and sexual dimorphism
has been documented in many groups of
birds and mammals [16, 17], including an
thropoid primates [2, 3]. This so-called al
lometric effect explains interspecific dif
ferences in sexual dimorphism as a non
selected, correlated response to evolutionary
changes in absolute body size [7, 27]. Al
though body size in prosimia ns is not corre
lated with sexual dimorphism [42], it may
nev ertheless constrain its expression to sorne
extent, because anthropoids in the prosim
ian size range fail to exhibit an allometric
effect as well, and are not significantly more
dimorphic than prosimians [42]. However ,
the recently extinct subfossil lemurs, which
included gorilla-sized species, apparently <l id
not exhibit marked size dimorphism either
[43, 44], suggesting that the constraint on
dimorphism in extant prosimians may be of
a phylogenetic, rather than allometric na
ture.
142
The differences in sexual dimorphism
among higher taxonomic units are particu
larly interesting in light of the evolutionary
history of prosimians and may illuminate
such possible phylogenetic constrain ts of di
morphism. There is now convincing ev i
dence that the Lemuriformes and Lorisi
farmes together constitute a monophyletic
group [45-48], but the exact phylogenetic
relationships among the members of these
two groups is still subject to heated debate
[45, 48-53]. Irrespective of their ancestral
relationships , the average degree of sexual
size has necessarily diverged in the two
groups. lf the ancestral species was sexually
dimorphic, or if the lemuriform radiation
represents the descendants of a colonization
of Madagascar by galago-like ancestors,
either average male size decreased and/or
average female size increased among the Le
muriformes. The Adapidae , a group of sex
ually dimorphic Eocene primates [1 1], have
been suggested by sorne authors as a group
containing the possible an.cestors of the ex
tant lemurs and lorises [46, 54], although
sorne authors disagree [55, 56]. This could
indicate that the common ancestor of Lemu
riformes and Lorisiformes was sexually di
morphic. If, on the other hand, the ancestral
species was monomorphic , or the mainland
lorises descended from a cheirogaleid-like
ancestor, the degree of sexual dimorphism of
the former group must have increased over
time. The choice between these two possibil
ities remains open.
Although inform ation about patterns of
sexual dimorphism above the species lev el
is potentially instructive in identifying di
rectionality of evolutionary change, and an
important prerequisite for statistically
sound comparative analyses [57], so-called
phylogenetic inertia cannot prov ide a con-
Kappeler
vincing explan ation for the evolution of
sexual dimorphism. Cheverud et al. [58]
and Leutenegger and Cheverud [59] hav e
argued that phylogenetic constraints are
primarily responsible far the degree of sex
ual dimorphism in extant primates . The ar
gument that a modern primate is dimor
phic because its ancestor was dimorphic
<loes not prov ide a causal explanation of
the origins and functions of sexual dimor
phism, but represen ts merely a description
of the same problem at a different level.
Morover, Leutenegger and Cheverud 's con
clusions may hav e been affected by meth
odological problems , including arbitrary as
sumptions about the magnitude of phyloge
netic distances among hierarchically nested
taxa [60-62].
The large amount of intrageneric
v ariance in body weight dimorphism
found in pro simians indicates that there
is no significant influence of phylogenetic
inertia on sexual dimorphism. This <l oes
not mean that phylo genetic inertia <loes
not exist, but rather that sexual dimorphism
may not be a good exam ple to illustrate
this problem, because it may not be a direct
target of selection. More like ly, sexual
dimorphism is the result of natural and
sexual selection acting on body size in
both sexes separately [5, 63]. However,
inter specific differences in mean body size
among prosimian species were also most
pro nounced among genera and not
among higher taxonomic levels, as would
be ex pected under the 'phylogenetic
inertia hy pothesis' [42].
Among prosimia n species with signifi
cant weight dimorphism , one is of particu
lar interest . In grey mouse lemurs (M . mu
rinus) females were significantly larger than
males. The size difference between the
sexes in M icrocebus has recently been cor-
Sexual Size Dimorphism in Prosimians
roborated independently, using skull length
as size variate [64]. Such 'reversed sexual
size dimorphism' is relatively unusual for
mammals [5], and has not previously been
demonstrated statistically for any primate
species [but see 65]. Larger body size in
female mammals continues to be a puzzle
to evolutionary biologists [5, 15, 66], and it
cannot be explained easily for this species
either. Most explanations of significantly
larger female body size in mammals have
centered around an advantage in fecundity
that larger females are supposed to obtain
through natural selection [5, 15, 66-68],
but a size reduction of males, favored for
example by intrasexual selection [69],
would provide an equally likely explanation
[42, 63].
The lack of male-biased sexual dimor
phism among Malagasy primates is associ
ated with the phenomenon of female social
dominance, suggesting a relation to selective
forces affecting the behav ioral and physio
logical strategies of males and females [12,
13, 70-72]. Howev er, other factors besid
es size are obviously important in
determining sex effects on dominance. In
the monomor phic ringtailed lemur (L.
eatta) , for example, females invariably
dominate males [70], whereas in the
equally monomorphic red fronted lemur (L.
fulvu s rufus) sex has no consistent effect
on the outcome of agonistic interactions
[71]. Detailed comparisons of the
socioecology and physiology of closely
related species of similar average body size
but with different degrees of sexual dimor
phism might help to identify sorne of the rel
evant selective forces that shaped the evolu
tion of sexual size dimorphism and sex
dif ferences in behavior. M . murinus, with
sig nificantly larger females, G. demidovii,
with significantly larger males, and the
mono-
143
morphic M . rufus might together provide a
good test case [73-75].
Significantly larger males were found
only among the non-Malagasy prosimians.
Bushbabies (Galagidae) were the most di
morphic group in this respect. Significant
weight dimorphism in favor of males is
usually explained as a result of intrasexual
selection [2, 14, 17]. The intensity of intra
sexual selection is assumed to increase with
increasing v ariance in lifetime reproductive
success among males. If intrasexual selection
was indeed the prime cause for the evolution
of larger male size, then male bushbabie s
should experience a greater v ariance in life
time reproductive success than lemur males.
This line of reasoning assumes that male
reproductive success is positively related to
body size. Alternatively, the costs of large
body size, such as increased risk of starva
tion and heat loss for example, m ay be eval
uated differently by natural selection in the
two groups. Howev er, far too little is
presently known about the mating systems
and other factors determining lifetime repro
ductive success of these prim ates to test this
explanation [13, 76].
Acknowledgmen ts
This study would hav e been impossible without
the num erous primate tech nicians at the DUPC who
collected the majority of the presented data over the
last 17 years. 1am also indebted to Drs. Ken Glander,
Kay Izard and Pat Wright for contributing sorne of
their own unpublished data. Special thanks to Drs.
Jorg Ganzhorn and Carel van Schaik for many stimu
lating discussions, and to Drs. Peter Klopfer, Robert
Martín, Louise Roth and Henry Wilbur for many
helpful comments on the manuscript. This work was
supported by a Harry Frank Guggenheim Foundation
Dissertation Award. This is Duke University Primate
Center publication No. 507.
144
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Received: Jun e 7, 1989
Accepted: May 29, 1991
Peter M. Kappeler
Department of Zoology
Duke University
Durham , NC 27706 (USA)