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Comparative penile morphology of East African galagos of the genus Galagoides (family Galagidae) implications for taxonomy.

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American Journal of Primatology 69:16–26 (2007)
RESEARCH ARTICLE
Comparative Penile Morphology of East African Galagos
of the Genus Galagoides (Family Galagidae):
Implications for Taxonomy
ANDREW PERKIN
Nocturnal Primate Research Group, Oxford Brookes University,
Reading, United Kingdom
Recent studies on the comparative penile morphology of galagos have
revealed complex patterns that vary among both genera and species,
and help with species identification. So far the penile morphologies of 14
galago species have been described and an identification key has been
proposed. The present study extends and revises previous work. Wild
Galagoides cocos, G. granti, G. zanzibaricus and G. rondoensis from the
Eastern Arc Mountains and coastal forests of Tanzania and Kenya were
live-trapped, and one species (G. orinus) was examined using a museum
specimen. Penile morphology was photographed, traced, and described
qualitatively. All of the mature males had penile spines. Spines were
absent or indistinct in immature males of all of the species. The penile
morphologies of G. cocos, G. granti, and G. zanzibaricus are similar in
their overall distribution and type of spines, but differ in the shape of the
glans penis. Both G. orinus and G. rondoensis have divergent penile
morphologies compared to all other galagos, and are probably phylogenetically distinct. The results support other recent morphological and
behavioral studies that consider these five galagos to be distinct species.
The evolution of the baculum and spines is thought to be linked to sexual
selection in multimale mating systems, but the mechanisms involved are
not entirely understood. Adult male G. rondoensis appear to display the
penile spines by exposing the distal section of the penis. Am. J. Primatol.
68:16–26, 2007. c 2006 Wiley-Liss, Inc.
Key words: penile morphology; galago; spines; baculum; cryptic female
choice; taxonomy
Contract grant sponsor: Oxford Brookes University; Contract grant sponsor: Primate Conservation
Inc.; Contract grant sponsor: Margot Marsh Biodiversity Fund; Contract grant sponsor: Tanzania
Forest Conservation Group; Contract grant sponsor: Africa Rainforest Conservancy.
Correspondence to: Andrew Perkin, 39A Rickman Close, Woodley, Reading, RG5 3LL UK.
E-mail: bwanakomba@yahoo.co.uk
Received 29 July 2005; revised 24 January 2006; revision accepted 13 February 2006
DOI 10.1002/ajp.20323
Published online in Wiley InterScience (www.interscience.wiley.com).
r 2006 Wiley-Liss, Inc.
Penile Morphology in Galagoides / 17
INTRODUCTION
There has been a recent increase in the number of galago species recognized,
as new techniques are enabling primatologists to better understand the extent
of the differences among these species, despite their cryptic morphology and
behavior [Bearder, 1999; Groves, 2000; Grubb et al., 2003; Masters & Bragg,
2000; Honess & Bearder, 1996; Nash et al., 1989]. The separation of species has
been aided by research on vocal repertoires, with different vocal profiles
correlating with differences in hand-pad structure, genetic diversity, reproductive
anatomy, and facial markings [Anderson et al., 2000; Bayes, 1998; Bearder et al.,
2006; Fjeldså et al., 2005; Masters et al., 2005]. The superficial similarity in
morphological characters of galagos may be due to extensive convergence
[Bearder, 1999].
Comparative studies of reproductive anatomy in primates have revealed
complex and specialized genitalia [Anderson, 1998; Dixson, 1998, 1989; Hill,
1953]. Previous studies of galago penile anatomy have shown elaborate
differences among species [Anderson, 1998, 2000; Dixson, 1995; Dixson & Van
Horn, 1977; Hill, 1953]. They vary in shape and in the presence and distribution
of hard, keratinized spines, as well as in the size, shape, and relative position
of the baculum. Three spine types have been defined: 1) simple spines with a
single point and moderate length; 2) robust spines that are single-pointed,
enlarged structures, often thickened at the base; and 3) complex spines that are
multipointed [Dixson, 1995]. The baculum can be prominently positioned and
exposed, or just visible and surrounded by tissue structures of the glans penis.
For species with spineless glans, the shape of the glans penis is diagnostic, while
for species with spines, the relative size and the position of the spines on the penis
vary among species [Anderson, 2000]. For example, studies of comparative penile
morphology have revealed little intraspecific variation within Otolemur garnettii
or O. crassicaudatus [Anderson, 1998, 2000].
Complex penile morphologies and the shape and size of the baculum in
multimale/multifemale mating systems, such as those found in the galagos, may
have evolved for two reasons: 1) to displace the sperm of other competitors, and 2)
to ‘‘encourage’’ females to take up spermatozoa by the male placing ejaculate in
the most advantageous position, or by stimulating uptake and transport of
spermatozoa within the female’s tract [Dixson, 1998]. This may lead to the penis
functioning as an ‘‘internal courtship device’’ [Eberhard, 1996]. The different
spine types found in galago penises may be related to the different roles that
each spine type may perform. Type 2 spines may have a role in mate-guarding,
while Type 1 spines may influence physiological feedback [Dixson, 1998].
While the reasons for this are not fully understood, it is speculated that penile
morphological variation may be the result of speciation events influenced by
sexual selection via cryptic female choice [Dixson, 1998].
The penile morphologies of 14 galago species have been described, including
the East African dwarf galago species Galagoides zanzibaricus, G. udzungwensis,
G. granti, and G. rondoensis [Anderson, 2000]. However, since Anderson’s [2000]
study was published, taxonomic changes affecting these species have been made.
G. cocos has been recognized as the correct name for the Kenya coast galago (often
mistakenly referred to as G. zanzibaricus) [Butynski et al., in press], and G.
udzungwensis has been subsumed into the taxonomy of G. zanzibaricus [Grubb
et al., 2003]. This study provides a detailed comparison of five species: Galagoides
cocos, G. zanzibaricus, G. granti, G. orinus, and G. rondoensis. These species
occupy the globally important biodiversity hotspots of the Eastern Arc Mountains
Am. J. Primatol. DOI 10.1002/ajp
18 / Perkin
and the coastal forest mosaic of Tanzania and Kenya [Burgess & Clarke, 2000;
Burgess et al., 2004].
MATERIALS AND METHODS
Twenty-five galagos were photographed and measured to examine the details
of their penile morphology. The study subjects included 21 live animals that were
trapped from various localities in the Eastern Arc Mountains and coastal forests
of Tanzania and Kenya, and four museum specimens. From this group, mature
males (based on spine and testes development) were selected for analysis
(Table I). Most of the galagos were trapped with the use of a Chardonneret trap
[Charles-Dominique, 1977], and others were captured opportunistically with mist
nets set for bats. Live galagos can be processed without the need for anesthetic if
they are handled in a quiet environment. In the case of museum specimens, both
the baculum and, in many cases, the penile spines survived wet and dry
preservation. The testes and scrotum survive only wet preservation.
Tracings were taken from each photograph and drawn to scale for direct
comparison within and among species. The delineation of regions of the penis is
adapted from Anderson [2000], who divided the penis into three equal sections:
distal, mid, and proximal. The qualitative characters for each species were
described by section and included the 1) relative shape and size of the glans penis,
2) spine type (see Introduction), 3) distribution of spines, and 4) relative position
of the baculum. Measurements of penis length were taken and in some cases the
dorsoventral and lateral penis widths were taken from points on the proximal,
mid-point, and distal parts of the penis. However, in contrast to Anderson’s
[2000] study, in which digital scanning technology enabled detailed quantitative
calculations of glans and spine area to be made, here I describe the penile
morphology qualitatively. When the galagos were captured, the amount by which
the distal part of the penis protruded beyond the prepuce was also noted. Dixson
[1998] compared the baculum and glans penis length with body mass. In this
study I determined a penis length to body mass ratio by dividing the species’
mean greatest penis length by the species’ mean body mass. The sample sizes
were too small to conduct statistical analyses. I assessed ontogenetic development
by comparing penile spine development with body size and testes development.
In some cases the testes length and widths were also measured.
RESULTS
In all of the mature male galagos examined, the penises had various
arrangements and types of backward-pointing spines; however, in immature
males the spines were absent or only just visible (Table I, Figs. 1,2,3). When data
were available, spine development was linked to the size and development of the
testes. Species accounts are given below.
Galagoides cocos
The penile morphology of this species has not previously been described.
Four specimens from three sites were examined, and only two of these were
mature (Table I). The glans penis is cylindrical but enlarges slightly in the middle
more dorsoventrally then laterally (Fig. 1). The mean lateral width at the penis
base, mid-section, and tip was 2.0, 2.3, and 2.2 mm, respectively (n 5 2). The
baculum protruded very little (o1 mm) from the glans penis in one specimen
examined, but remained equal to the level of the glans penis in the other mature
Am. J. Primatol. DOI 10.1002/ajp
G. zanzibaricusa
G. zanzibaricus
G. zanzibaricusa
G. zanzibaricus
G. granti,a
FMNH/ KMH
25830
G. granti,a
FMNH/ KMH
25799
G. zanzibaricusa
G. cocos
G. cocos,a NMK,
MR14
G. cocos
Galagoides cocosa
Species, museum,
and specimen
number given
where relevant
Diani FR, Kenya
(coordinates: 0419S
3933E)
Mgambo FR, Kenya
(coordinates: 0445S
3848E )
Mgambo FR, Tanzania
Mrima Hill FR, Kenya
(coordinates: 0428S
3915E)
Kichi Hills FR, Tanzania
(coordinates: 0429S,
3916E)
Lulanda FR, Tanzania
(coordinates: 0707S
3623E)
Pugu FR, Tanzania
(coordinates: 0642S
3905E)
Pugu FR, Tanzania
Kazimzumbwi FR,
Tanzania (coordinates:
0643S 3902E)
Kazimzumbwi FR,
Tanzania
Pande GR, Tanzania
(coordinates: 0642S
3905E)
Locality
21.0
16.0
23.0
9.0
133.5
136.0
147.0
122.0
145.0
160.0
18.6
22.0
140.0
108.0
165.0
69.0
177.0
Mass (g)
21.0
20.0
23.0
18.0
21.5
Greatest penis
length (mm)
10/9
5/5
NM
TND
18/15
NM
NM
TND
NM
TND
NM
Scrotum greatest
length/width (mm)
Partly developed
Not present
Fully developed
Not present
Fully developed
Fully developed
Fully developed
Not present
Fully developed
Not present
Fully developed
Penile spine
development
Mature
Immature
Mature
Immature
Mature
Mature
Mature
Immature
Mature
Immature
Mature
Assessment of
sexual maturity
TABLE I. The Localities, Greatest Penis Length, Greatest Width, and Length of Scrotum, Body Mass and Presence of Spines of the
Galagos Examined
Penile Morphology in Galagoides / 19
Am. J. Primatol. DOI 10.1002/ajp
Am. J. Primatol. DOI 10.1002/ajp
zanzibaricusa
rondoensis
rondoensis
rondoensisa
rondoensisa
rondoensisa
rondoensisa
rondoensis
rondoensis
Pande GR, Tanzania
Mkungwe FR, Tanzania
(coordinates: 0652S
3734E)
Pugu FR, Tanzania
Pugu FR, Tanzania
Pugu FR, Tanzania
Pugu FR, Tanzania
Pugu FR, Tanzania
Pugu FR, Tanzania
Pugu FR, Tanzania
Pugu FR, Tanzania
Kazimzumbwi FR,
Tanzania
Kazimzumbwi FR,
Tanzania
Pande GR, Tanzania
New Dabaga FR, Tanzania
(coordinates: 0805S
3554E)
Locality
25.0
12.8
26.0
18.2
11.0
11.0
24.0
26.0
22.0
20.0
9.0
12.0
21.5
8.0
11.0
Greatest penis
length (mm)
80.0
97.0
65.0
150.0
71.0
52.0
62.0
72.0
61.0
81.0
54.0
57.0
177.0
60.0
111.0
Mass (g)
Fully developed
Fully developed
Partly developed
Fully developed
Partly developed
Not present
Fully developed
Fully developed
Fully developed
Fully developed
Partly developed
Partly developed
Fully developed
Not present
Not present
Penile spine
development
Mature
Mature
Mature
Mature
Immature
Immature
Mature
Mature
Mature
Mature
Immature
Immature
Mature
Immature
Immature
Assessment of
sexual maturity
Animals compared in this study.
FMNH, Field Museum of Natural History (Chicago); NMH, National Museums of Kenya; KMH, Collection number of Prof. K.M. Howell; FR, forest reserve; GR, game reserve;
NM, not measured; TND, testes not descended (therefore not measured).
a
NM
NM
NM
NM
13/10
TND
10/24
15/18
NM
NM
TND
TND
NM
TND
3/3
Scrotum greatest
length/width (mm)
A level of sexual maturity is assessed on the basis of the presence of penile spines and testes development.
G. rondoensisa
G. orinus,a
FMNH/ KMH
22496
G. rondoensisa
G.
G.
G.
G.
G.
G.
G.
G.
G.
Galagoides cocosa
G. zanzibaricus
G. zanzibaricus
Species, museum,
and specimen
number given
where relevant
TABLE I. Continued
20 / Perkin
Penile Morphology in Galagoides / 21
Fig. 1. Schematic drawings and relative sizes of the penile morphologies of 1) G. cocos, 2) G. granti,
3) G. zanzibaricus, 4) G. rondoensis, and 5) G. orinus (specimen FMNH/KMH 22496). A: Ventral
view. B: Dorsal view. C: Lateral view (dorsum right). Scale is indicated on the right.
specimen. The mean length of the penis is similar to G. zanzibaricus and G.
granti, with a relative penis length index of 0.130 (n 5 2; Table II). There are no
spines on the tip of the penis, but simple Type 1 spines start to emerge in the
proximal half of the distal section. These develop into downward-pointing robust
Type 2 spines that are found in the mid-section, and in the proximal section they
reduce in size to Type 1 spines (Fig. 1). The spines are geometrically arranged in
diagonal lines on the penis. The immature animals examined (n 5 2) had low body
mass and no penile spines. One had a relatively long penis (20 mm) but no
descended testes (Table I). Prior to examination it was noted that only the distal
tip of the penis protruded from the foreskin.
Am. J. Primatol. DOI 10.1002/ajp
22 / Perkin
Fig. 2. Examples of an immature G. zanzibaricus with undeveloped spines (top: lateral profile), and
a mature G. zanzibaricus penis with fully developed spines (bottom: dorsal profile). The baculum
protrudes more prominently in mature males. A mature G. rondoensis (right) held after it was
extracted from the trap, showing the prominent distal penile morphology display. Photos not
to scale. [Color figure can be viewed in the online issue, which is available at www.interscience.
wiley.com.]
Fig. 3. Photographs of the penile morphology of G. rondoensis: (a) an immature male (lateral
profile), (b) a mature male (lateral view), and (c) a mature male (dorsal view). Photos not to scale.
[Color figure can be viewed in the online issue, which is available at www.interscience.wiley.com.]
Galagoides granti
Two mature individuals from different sites were examined (Table I). In both
cases the penis was larger dorsoventrally (mean width at mid-section 5 5.52 mm)
than laterally (mean width at mid-section 5 2.54 mm), giving a thin pointed
appearance overall (Fig. 1). The baculum protrudes at a point ‘‘off center’’ toward
the dorsal side of the penis. The mean length of the penis is similar to but shorter
than that of G. cocos and G. zanzibaricus, and the relative penis length index
is 0.133 (n 5 2; Table II). There are no spines around the penis tip, but Type 1
spines start to emerge in the proximal half of the distal section. Downwardpointing robust Type 2 spines cover the mid-section, with the highest densities on
Am. J. Primatol. DOI 10.1002/ajp
Penile Morphology in Galagoides / 23
TABLE II. Mean Penis Length and Body Mass of the Mature Galagos Examined
and the Penis Length to Body Mass Ratios Compared for the Five Galagos
Species Studied
Species
G.
G.
G.
G.
G.
cocos
granti
zanzibaricus
rondoensis
orinus
Mean
greatest penis
length (mm)
Range
(mm)
Mean
mass (g)
Range
(g)
n
Relative
penis lengtha
22.25
20.50
21.05
23.60
12.80
21.5–23.0
18.6–21.5
18.2–23.0
20.0–26.0
12.8
171.00
150.00
143.90
70.16
97.00
165.0–177.0
140.0–160.0
133.5–150.0
61.0–80.0
97.0
2
2
4
6
1
0.130
0.133
0.146
0.342
0.132
G. rondoensis has a high penis length to body mass index.
a
Species mean greatest penis length/species mean body mass (see text and Table I).
the lateral flanks and the dorsal section, and few on the ventral section. The
spines continue partially into the proximal third and then disappear. The spines
are geometrically arranged in lines on the penis. Prior to examination it was
noted that only the distal tip of the penis protruded from the foreskin.
Galagoides zanzibaricus
Seven individuals from four different sites were examined, and five of these
were considered mature (Table I). The penises of the mature individuals were
cylindrical in shape, but expanded greatly laterally in the mid-portion of the penis
(Figs. 1 and 2). The mean lateral width at the penis base, mid-section, and tip was
2.2, 3.5, and 2.1 mm respectively (n 5 3). Immature individuals were similar in
shape, but expansion of the mid-section of the penis was less exaggerated. The
baculum protrudes at a point off-center toward the dorsal side of the penis. The
mean length of the penis is similar to G. cocos and G. granti, and the relative
penis length index is 0.146 (n 5 5; Table II). There are no spines around the tip,
but Type 1 simple spines start to emerge in the proximal half of the distal section.
Downward-pointing robust Type 2 spines cover the mid-section completely. The
spines on the dorsal side of the penis are arranged geometrically and longitudinally, and point slightly inward to a central line or crease down the middle of
the penis, mostly in the mid-section (Figs. 1 and 2). The spine cover continues
down into the proximal third of the penis on the dorsal surface, but it starts to
reduce in size and spines are absent on the ventral and lateral surfaces. Immature
animals of G. zanzibaricus with undescended or relatively small testes either have
no spines, or they have a lower body mass and less-developed spines (Table I,
Fig. 2). However, the gross shape of the glans penis remains consistently similar
between immature and mature males. Prior to examination it was noted that only
the distal tip of the penis protruded from the foreskin.
Galagoides rondoensis
Ten individuals from three sites were examined, and six of these were
considered mature (Table I). G. rondoensis has a large and highly differentiated
penis. The penis has a long narrow shaft in the proximal region (mean
dorsoventral width 5 2 mm, n 5 6) and mid-sections (mean dorsoventral
width 5 2 mm, n 5 6) before it expands and becomes ‘‘flute shaped’’ in the distal
Am. J. Primatol. DOI 10.1002/ajp
24 / Perkin
region (mean dorsoventral width 5 6.5 mm, n 5 6). The baculum is exposed at a
point off-center toward the dorsal side of the penis of the glans penis, but is
‘‘nested’’ inside so that it does not protrude above the level of the glans. In
comparison to the other species examined here, G. rondoensis has a remarkably
large penis, resulting in a high relative penis length index of 0.342 (n 5 6;
Table II). The penis has a cluster of Type 1 and 2 spines around the ventral and
lateral portions of the tip. Two of the largest spines, one on each side, appear to
protrude markedly at approximately 451 to the penis. The spines do not appear to
be arranged geometrically. There are no spines present in the mid- and proximal
regions.
Spines are not present in immature animals with undescended or small
testes, but small nodes or ‘‘buds’’ can be seen in the same areas where spines are
formed in adults (Fig. 3). The relative spine development noted was thought to be
related to different levels of maturity, with the greatest development occurring in
individuals with larger bodies and testes. Upon capture the distal end of the penis
and the ‘‘spinal array’’ were noted to be consistently exposed and visible above
the foreskin in mature males (Fig. 2).
Galagoides orinus
The penile morphology of Galagoides orinus was previously undescribed. The
current description is based on one specimen that, as far as we know, is the only
male of this species in the museum collections. The specimen was judged to be
mature, based on body measurements, body mass, and the presence of descended
testes (Table I). G. orinus has a unique and differentiated penis morphology (Fig.
1). The overall shape of the penis is conical. The proximal section is narrow
(dorsoventral width 5 2.1 mm) and widens through the mid-section (dorsoventral
width 5 2.3 mm), and continues to widen to the distal end (dorsoventral
width 5 3.2 mm) of the glans penis. G. orinus has a small penis in absolute
terms, but its relative penis length index of 0.132 (n 5 1) is similar to that of
G. cocos, G. zanzibaricus, and G. granti (Table II). On the dorsal surface of the
proximal section there is a group of approximately 14 large Type 2 spines
arranged in a regular pattern, but no spines are visible on the lateral and ventral
sections. The mid-section is complex. The Type 2 spines continue to a mid-point
and then there is an area with no spines. Small, geometrically positioned Type 1
spines cover the distal region up to the end of the glans penis. From the end of
the glans penis, the baculum protrudes prominently and is positioned toward the
dorsal edge.
DISCUSSION
The results of this study support the conclusion that penile morphology
can be used to help identify galago species and aid in their classification. All of
the species examined in this study had spined penises, a result that is
contradictory to some of the findings of Anderson [2000]. This may indicate that
the G. udzungwensis (now classified as G. zanzibaricus), G. granti, and
G. rondoensis specimens illustrated in Anderson [2000] were immature animals.
Indeed, Honess [1996] acknowledged that this may be the case for the
G. rondoensis (n 5 1) and G. udzungwensis (n 5 1) individuals trapped in that
study. G. cocos, G. granti, and G. zanzibaricus, which have undergone recent changes
in taxonomic status, are superficially similar in body morphology and may be
closely related species [Butynski et al., in press] (Perkin, unpublished data). This
aspect is also indicated by similarities in their penile morphologies. The penile
Am. J. Primatol. DOI 10.1002/ajp
Penile Morphology in Galagoides / 25
morphologies of G. zanzibaricus examined in this study, sampled from mainland
coastal sites, appear to differ from the description of the penile morphologies of
G. udzungwensis (now renamed G. zanzibaricus) from Matundu Forest Reserve
(ca. 250 km further inland to the southwest), indicating possible intraspecific
variation [Anderson, 2000; Honess, 1996].
G. rondoensis is a highly distinctive species, both morphologically and
behaviorally [Honess, 1996] (Perkin unpublished data). Equally, the penile
morphology is highly distinctive in terms of relative length, shape, and spinal
distribution. G. rondoensis is the only galago species in this study that always
displayed the distal region of the penis when captured (Figs. 2 and 3). Whether
this is display is detectable and registered by females prior to mating is an
intriguing question. Judging from the one specimen examined, G. orinus has a
unique penile morphology that is readily distinguishable from other galago
species. This includes G. demidoff, of which G. orinus was once considered to
be a subspecies [Grubb et al., 2003]. These taxa also differ from each other vocally
and morphologically [Grubb et al., 2003; Honess & Bearder, 1996; Lawrence &
Washburn, 1936; Perkin et al., 2002].
When describing galago penile morphology, investigators should consider the
ontogeny, because immature males may exhibit no or undeveloped spines.
The profile of the penis must be indicated for an accurate comparison of penile
morphology. Further research and greater sample sizes are required before
we can understand the levels of intra- and interspecies variation among galago
penile morphologies. The roles of testes development and social organization in
relation to penis ontogeny are also poorly understood.
ACKNOWLEDGMENTS
My thanks go to Professor S.K. Bearder, Dr. N. Burgess, and N. Doggart
for comments. I thank H. Gideon, WWF Tanzania, IUCN, Wildlife Conservation
Society of Tanzania, Colobus Trust, National Museums of Kenya, and the Field
Museum of Natural History for their assistance. The Tanzania Commission
for Science and Technology kindly granted research permission for the conduct
of this research program.
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implications, genus, morphology, taxonomy, africa, penile, family, comparative, east, galagidae, galagoides, galago
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