Limestone cliff - face and cave use by wild ring-tailed lemurs (Lemur catta) in southwestern Madagascar

Abstract

Ring - tailed lemurs live in a range of habitats in southwestern Madagascar. To date, much of the knowledge of ring - tailed lemur ecology, biology and behavior come from riverine gallery forests sites. Recent years have seen an expansion of comprehensive research on this resilient species, including areas of limestone spiny forest along Madagascar’s southwestern coast. This work is documenting newly discovered behaviors by this species. The regular use of cliff - faces and embedded crevices and caves by ring - tailed lemurs in southwestern Madagascar are reported here. Cave use by several anthropoid primates has been explained as a thermoregulatory behavior. It is suggested that cliff - face and cave use by these ring-tailed lemurs serves several purposes, including resource acquisition, thermoregulation, and as an anti - predator avoidance strategy in the absence of suitable large sleeping trees. Observations indicate that the limestone boundaries of the Mahafaly Plateau and their associated xerophytic scrub forests warrant further conservation attention, given the presence of behavioral variation and increasing threats to this endangered primate species.

Full text

MADAGA SCAR CONSERVATION & DEVE LOPMENT VOLUME 8
|
ISSUE 2 — NOVEMBER 2013 PAGE 7 3
Limestone cliff - face and cave use by wild ring-tailed
lemurs (Lemur catta) in southwestern Madagascar
ARTICLE http://dx.doi.org/10.4314/mcd.v8i2.5
Michelle L. Sauther, Frank P. CuozzoI, Ibrahim A.
Youssouf JackyII, Krista D. FishIII, Marni LaFleurIV, Lova
A. L. RavelohasindrazanaII and Jean F. RavoavyII
Correspondence:
Michelle L. Sauther
Department of Anthropology, University of Colorado-Boulder,
Boulder, Colorado, 80309, U.S.A.
E - mail: sauther@colorado.edu
I Department of Anthropology, University of North Dakota, Grand Forks, North Dakota, 58202, U.S.A.
II Department of Animal Biology, Faculty of Sciences, University of Toliara (601), Madagascar.
III Department of Anthropology, Colorado College, Colorado Springs, Colorado, U.S.A.
IV Institute for Population Genetics University of Veterinary Medicine, Vienna Josef Baumann Gasse 1 1210 Vienna, Austria.
ABSTRACT
Ring - tailed lemurs live in a range of habitats in southwestern
Madagascar. To date, much of the knowledge of ring - tailed
lemur ecology, biology and behavior come from riverine gallery
forests sites. Recent years have seen an expansion of compre-
hensive research on this resilient species, including areas of
limestone spiny forest along Madagascar’s southwestern coast.
This work is documenting newly discovered behaviors by this
species. The regular use of cliff - faces and embedded crevices
and caves by ring - tailed lemurs in southwestern Madagascar
are reported here. Cave use by several anthropoid primates
has been explained as a thermoregulatory behavior. It is sug-
gested that cliff - face and cave use by these ring-tailed lemurs
serves several purposes, including resource acquisition, ther-
moregulation, and as an anti - predator avoidance strategy in the
absence of suitable large sleeping trees. Observations indicate
that the limestone boundaries of the Mahafaly Plateau and their
associated xerophytic scrub forests warrant further conserva-
tion attention, given the presence of behavioral variation and
increasing threats to this endangered primate species.
RÉSUMÉ
Lemur catta occupe divers habitats dans le Sud - ouest de
Madagascar. L’écologie, la biologie et le comportement de
Lemur catta sont actuellement mieux connus des populations
vivant dans les forêts riveraines et les zones environnantes. Pour
mieux comprendre cette espèce de lémurien, les recherches
ont été étendues à d’autres habitats dont les forêts épineuses
du plateau calcaire qui est situé le long du littoral Sud - ouest
de Madagascar. Dans cette étude nous rapportons les com-
portements récemment découverts de Lemur catta qui utilise
les falaises et les grottes dans le Sud - ouest de Madagascar.
L’utilisation des grottes par la plupart des primates hominoïdes
est liée à un avantage thermorégulateur offert par ce milieu.
Dans notre cas, l’exploitation de falaises et de grottes par
Lemur catta semble être associée à un mécanisme permettant
d’échapper aux prédateurs et à l’absence de grands arbres
qui devaient servir de dortoirs. De sorte que les falaises et les
forêts épineuses du plateau calcaire Mahafaly ont besoin d’une
conservation particulière car nos résultats de suivis montrent
que les changements de comportement du lémurien emblé-
matique de cette région trouve vraisemblablement son origine
dans la dégradation de l’environnement de cette espèce.
INTRODUCTION
Ring - tailed lemurs (Lemur catta) are a resilient primate species,
inhabiting a wide range of habitats across southern and south-
western Madagascar that includes areas of heavy human dis-
turbance (e.g., Sauther et al. 1999, Gould 2006, Jolly et al. 2006).
Spanning a range from the southernmost point in Madagascar
(Cap Sainte Marie), across the spiny forests and dry deciduous
riverine forests of the Atsimo - Andrefana Region, through the
high-altitude Andringitra Massif and surrounding highlands,
ring-tailed lemurs live within a range of challenging environ-
ments in terms of temperature variation, droughts, cyclones
and differing levels of anthropogenic change (Goodman et al.
2006, LaFleur 2012). Lemur catta is also the most terrestrial of all
living lemurs, and exploits a wide range of resources, depending
on the habitat (e.g., Sauther et al. 1999, Goodman et al. 2006,
Gould 2007). To date, most of what is known about this species’
biology, ecology, and behavior comes from long - term studies
(> 25 years) at two riverine gallery sites, the Bezà Mahafaly
Special Reserve and Berenty Private Reserve (Gould 2007). Only
recently has information on this species been available from
more xeric spiny forest areas, including along the limestone pla-
teau of Madagascar’s southwestern region (Sauther and Cuozzo
2008, Kelley 2011, Cuozzo and Sauther 2012, LaFleur 2012). In
this paper, the use of caves and cliff fissures by ring-tailed
lemurs at two sites, Tsimanampesotse National Park and the
Tsinjoriake New Protected Area, both located in southwestern
Madagascar, is described in detail for the first time, and possible
ecological functions of this behavior discussed.
Cave use is rare among primates, but has been previously
reported among continental African and Asian anthropoid
primates, including chimpanzees, baboons, and langurs. A vari-
ety of hypotheses have been put forth regarding this behavior,

Madagascar Conservation & Development is the journal
of Indian Ocean e-Ink. It is produced under the respon-
sibility of this institution. The views expressed in contri-
butions to MCD are solely those of the authors and not
those of the journal editors or the publisher.
All the Issues and articles are freely available at
http://www.journalmcd.com
Contact Journal MCD
info@journalmcd.net for general inquiries regarding MCD
funding@journalmcd.net to support the journal
Madagascar Conservation & Development
Institute and Museum of Anthropology
University of Zurich
Winterthurerstrasse 190
CH-8057 Zurich, Switzerland
Missouri Botanical Garden (MBG)
Madagascar Research and Conservation Program
BP 3391
Antananarivo, 101, Madagascar
Indian Ocean e-Ink
Promoting African Publishing and Education
www.ioeink.com

MADAGA SCAR CONSERVATION & DEVE LOPMENT VOLUME 8
|
ISSUE 2 — NOVEMBER 2013 PA G E 74
many positing thermoregulation as a key reason (Pruetz 2001,
2007, Huang et al. 2003, McGrew et al. 2003, Barrett et al. 2004,
Workman 2010). For example, cave use among chimpanzees in
southeastern Senegal looks to be an “opportunistic” behavior
in response to high temperatures (Pruetz 2007: 318). Explana-
tions for cave use by primates also include access to water,
obtaining nutrients through geophagy and predator avoidance
(McGrew et al. 2003).
Among Madagascar’s strepsirhine primates, only anecdotal
observations of cave use have been previously reported. Ring-
tailed lemurs at Andrahomana cave, in southern Madagascar
have been seen resting and feeding during the daytime in this
cave (Vasey et al. 2013). At Isalo National Park, on the southern
edge of Madagascar’s central highlands, ring - tailed lemurs
were observed entering caves to lick the soil as well as using
caves during the day and at night (Dinets, pers. com). Finally,
a single report from northern Madagascar describes crowned
lemurs (Eulemur coronatus), using a cave to drink water
(Wilson et al. 1989).
Early reports by Perrier de la Bâthie (1927) described ring-
tailed lemurs using the vertical cliff-faces of the Andringitra
Massif, central Madagascar. A more recent study of these high-
altitude (ranging above 2,000 meters) L. catta on the Andringitra
Massif, near the northern boundary of this species’ range in
south - central Madagascar, has documented the use of fissures
and overhangs in the granitic outcrops as sleeping sites (Good-
man et al. 2006). Goodman et al. (2006) hypothesize that i) caves
serve as shelter in response to cold temperatures at this alti-
tude, and/or ii) caves and rock overhangs may be a way of avoid-
ing predation by the endemic fossa (Cryptoprocta ferox). Gould
(pers. comm.) has recently observed a small L. catta troop (n = 4)
that uses a small cave in the Ambalavao region of the highlands,
in the same region as the Andringitra Massif. This population
lives in habitat with no forest cover due to slash - and - burn
agriculture, and survives on anthropogenic resources such as
crops and fruit trees. This example of ring - tailed lemur cave
use is likely a result of human action rather than ecological
adaptations such as thermoregulation or predator avoidance,
and is similar to langur cave use described by Li and Rogers
(2005), where these monkeys use caves and limestone karst as
a refuge in response to deforestation and expanding cultivation.
In fact, all of the scattered, fragmented L. catta populations in
this highland region may be using the granitic slopes, massifs
and caves as a refuge, as the surrounding areas are void of
continuous forest, subject to frequent intentional burning (e.g.,
Goodman and Langrand 1996), and are marked by vast expanses
of cultivated land (Cameron and Gould 2013). Data on ring - tailed
lemur cliff - face and cave use in areas of continuous forest,
but without the large sleeping trees present in riverine gallery
forest areas such as Bezà Mahafaly Special Reserve and Berenty
Private Reserve, would add to current anecdotal reports from
this and other species and provide valuable data to interpret
and explain this behavior beyond it being a response to anthro-
pogenic effects.
METHODS
As part of the authors’ long - term collaborative research on the
ecology, biology and behavior of ring -tailed lemurs in south-
western Madagascar, research has been carried out along
the western edge of the Mahafaly Plateau, Atsimo - Andrefana
Region. Data presented here are based on a cumulative 11
months of direct observations, supplemented with data from
camera traps. The authors have spent varying amounts of time
at Tsimanampesotse National Park (E43°46’–43°50’, S24°03’–
24°12’) ranging from several weeks to seven months: May/
June 2006 (MLS, FPC, IAJY); May/June 2007 (MLS, FPC, IAJY);
June 2008 (MLS, FPC, IAJY, MML); September 2010 through
April 2011 (MML); August 2013 (ML). During the seven - month
behavioral study, camera traps monitored daily lemur activity
as well as predator behavior. Tsimanampesotse is a 42,000 ha
National Park representing the western most escarpment of
the limestone Mahafaly Plateau and is constructed of Cenozoic
limestone (DuPuy and Moat 1998). A highly seasonal habitat,
most rainfall occurs between December and February with
annual rainfall rarely exceeding 500mm (Donque 1975, APAAT
Protected Areas Report: http://bioval.jrc.ec.europa.eu/APAAT/
pa/2307/), although during this same time period during this
research rainfall was only 400mm (LaFleur 2012). Temperature
also varies dramatically with daytime highs of well over 40°C,
although mean daily temperatures range between 22.5°C
and 35.8°C (LaFleur 2012). The area is also affected by high
winds, frequent droughts and cyclones (Andriatsimietry et al.
2009, LaFleur 2012). At Tsimanampesotse census data on five
groups are available from 2006, 2007 and 2010–11: Vintany (n
= 12–20 adults), ILove (n = 9–13 adults), Akao (n = 9–15 adults),
Capture Be (n = 14 adults) and Miandry (n = 12 adults). For
this report the focus is on three lemur troops that have been
regularly observed, Vintany, ILove and Akao, including during
the above - mentioned continuous seven-month study. Standard
focal sampling methods were used during this study (LaFleur
2012). The Vintany and ILove troops inhabit an area above
saline Lac Tsimanampesotse, in the limestone spiny forest on
the western edge of the Mahafaly Plateau. Akao spends parts
of its time along the eastern edge of Lac Tsimanampesotse,
as well as ascending the western edge of the escarpment.
Tsimanampesotse contains dry spiny forest with open - canopied
dwarf flora of the Euphorbiaceae, Didiereaceae, Bombaceae,
and Fabaceae families as well as areas of semi - deciduous trees
(LaFleur 2012, Jacky Youssouf, unpub. data). To characterize the
habitat at Tsimanampesotse in 2007 three areas encompassing
the lemur groups have been evaluated using seven transects
covering a total of 300x10 meters for each of the three areas.
Most recently (2012), the authors expanded their work to
include an area north of Tsimanampesotse and the Onilahy River
mouth/Saint Augustin Bay, in the Tsinjoriake New Protected
Area, between 15km and 25km south of Toliara (E43°45’36”,
S23°26’53”). This newly protected area encompasses approxi-
mately 25,000 ha of limestone cliffs and southwestern dry spiny
forest thicket as well as coastal mangroves. A total of 12 20x50
m vegetation transects were carried out to characterize the
lemur’s habitats. During a brief previous survey in 2007 by the
University of Brighton, Lemur catta were reported to sleep in
trees near a cave area and within nearby crevices (Scott et al.
no date). In April 2012 members of the research team carried out
an intensive month long study of ring - tailed lemurs at Tsinjori-
ake that included daily census, habitat analyses and behavioral
ecology (Ravelohasindrazana 2013, Ravoavy 2013). LALR and
JFR also conducted interviews and administered questionnaires
to 30 community members regarding their use of resources in
the area. Eight ring - tailed lemur groups were observed using

MADAGA SCAR CONSERVATION & DEVE LOPMENT VOLUME 8
|
ISSUE 2 — NOVEMBER 2013 PAGE 75
focal sampling within two locations during April 2012. These
were: Ambanilia troops; Mailaka (n = 5), Ekipa (n = 11), Fetry (n
= 3), Tsy misaraky (n = 2), and Antsifotse troops; Troop 1 (n = 2),
Troop 2 (n = 3), Troop 3 (n = 4) and Troop 4 (n =5). In July/August
2013, a second preliminary ecological survey was conducted,
including the use of camera traps, of Lemur catta and their
habitats at two locations in the Tsinjoriake Protected Area, one
in the Binabe area including the Grotte (or cave) of Binabe and
the Grotte Ambanilia area where the Sarodrano Peninsula meets
the mainland, north of the mouth of Onilahy River.
RESULTS
During each of the seven time periods spent observing Lemur
catta at Tsimanampesotse and/or Tsinjoriake, at least some
troops within these larger populations regularly, and for the
ILove troop at Tsimanampesotse exclusively, used cliff - face
crevices and/or small caves as sleeping sites at night (Figure
1). This was true during the continuous seven-month study of
this troop from September 2010 through April 2011. Of inter-
est, not all troops at these localities exhibit this behavior. At
Tsimanampesotse the Vintany troop regularly sleeps in a large
cluster of Ficus sp. trees at the top of a vertical limestone
depression, where these lemurs sometimes descend to drink
water. The Akao troop at Tsimanampesotse was never seen
using cliff - face areas as sleeping sites; rather, they regularly
slept in a stand of introduced Australian pines (Casuarina equi-
setifolia) along the eastern lake margin. Vegetation transects
revealed that there were very few large (> 10 cm DBH) Ficus sp.
or Tamarindus indica trees available for sleeping (20 of 225 trees
measured), and the majority of these are found near limestone
sinks that may also contain steep cliff - faces and/or caves.
At Tsinjoriake, of the eight troops observed, four commonly
used one of three caves for sleeping at night (Grotte Binabe:
Troops 2 and 4; Grotte Binakely: Troop 3; Grotte Ambanilia: Ekipa
Troop). During the 2013 visit to the area no direct observations
of the lemurs using Grotte Ambanilia were made, but fresh
L. catta fecal material near the entrance was found, indicat-
ing they were still using this cave as resting or sleeping sites.
In addition to their use as nightly sleeping sites, larger caves
within limestone sinks/depressions commonly contained pools
of water that were used by lemurs as drinking sites at both
Tsimanampesotse (Figure 2, cf. Video 1) and Tsinjoriake (at
Grotte Sarodrano and Grotte Binabe). The larger caves were
also used as ‘day caves’ during the hot season apparently to
cool off at Tsimanampesotse. At Tsimanampesotse members of
ILove group also lick the limestone walls around the caves and
fissures (cf. Video 2). Large trees in the area, those greater than
10 cm in DBH (Ficus marmorata, Poupartia sylvatica, Tamarindus
indica and Noronhia sp.), are limited to coastal mangrove forest
and a small gallery forest as well as within the immediate vicinity
of large limestone caves that contain pools of water. The major-
ity of the vegetation is dwarfed, with heights less than 2.5m
(cf. Zafisamimanana 2012, Ravelohasindrazana 2013, Ravoavy
2013). Thus, four of the eleven troops observed at both loca-
tions used cliff - face crevices and/or caves as sleeping sites and
most troops were also observed using large caves as drinking
sources and/or potentially as refugia from excessive heat along
the western edges of the limestone Mahafaly Plateau. These
data greatly expand the knowledge of ring - tailed lemur cave
use in Madagascar, which to date has been primarily anecdotal.
DISCUSSION
Cave use by non - human primates has been linked to ther-
moregulation, predator avoidance, and/or resource access, as
well as refugia in response to human actions such as defor-
estation (e.g., Pruetz 2001, 2007, Huang et al. 2003, McGrew
et al. 2003, Goodman 2006, Workman 2010). Similar to the
langurs of Southeast Asia discussed earlier (Huang et al. 2003,
Workman 2010), ring - tailed lemur cave use in the Andringitra
and Tsaranoro region much further north of Tsimanampesotse
and Tsinjoriake is likely linked to deforestation, given only rem-
nant forests exist in these areas (Goodman and Langrand 1996,
Cameron and Gould 2013). As the areas in which observations
of cliff-face and cave use display intact forests, at least until
recently, it is unlikely that cave use in this region is a direct
response to deforestation.
FIGURE 1. The ILove troop lemurs on cliff-face, about to enter small sleeping
caves at Tsimanampesotse National Park. Arrow denotes one of the sleep-
ing caves.
FIGURE 2. The ILove troop drinking water from the Grotte Mitoho at
Tsimanampesotse. Photo by Violaine Pellicheroo, with permission from the
author.

MADAGA SCAR CONSERVATION & DEVE LOPMENT VOLUME 8
|
ISSUE 2 — NOVEMBER 2013 PA G E 7 6
Large caves with pools of fresh water do provide important
sources of drinking water during the day at both study sites,
especially since these are available year round (LaFleur 2012).
In addition, at Tsinjoriake there are also limestone seeps along
the edges of the coastal Mangrove forests which make fresh
water available during low tide, and which are used by some
lemur troops (e.g., Ekipa troop). Goodman et al. (2006) put forth
the interesting question of whether ring - tailed lemurs can
exist in areas without fresh water and suggest that, especially
in spiny forests, they may meet their water needs from their
diet and/or from dew that collects on plants overnight. At a
highly anthropogencially altered habitat in the Cap Sainte - Marie
region of southern Madagascar, Kelley (2011) reports only one
instance of ring - tailed lemurs drinking water and suggests the
lemurs there also receive their water requirements from their
foods, particularly the introduced Opuntia sp. Nevertheless,
the fact that lemurs living within the spiny forest habitats of
Tsimanampesotse and Tsinjoriake do seek out and use fresh
water resources support the hypothesis that access to water
may be significant, and that pools of fresh water within such
caves or fresh water emanating from limestone seeps are an
important aspect of their ecology in these intact spiny forests.
It is also possible that, like at Isalo National Park, the lemurs
may practice geophagia by ingesting soil within cave environs.
To date this has not been observed within the actual caves but
has been seen in other areas (LaFleur 2012). However, as noted
above, in August 2013, MML observed ring - tailed lemurs licking
the walls of limestone cliffs where they sleep at Tsimanamp-
esotse. As water precipitates out of the limestone and dries it
may provide access to minerals for these lemurs. This licking
behavior, along with geophagia, has been widely noted among
ring-tailed lemurs living in gallery forest sites (e.g., Jolly 1966,
Loudon et al. 2006).
Cave use in this region may also be a way to stay cool
during the hot wet season as well as to stay warm during the
cool dry season (at Tsimanampesotse temperatures range from
10.4° C to 41.9° C but can be even hotter or cooler in more open
areas (LaFleur 2012)). While some ring - tailed lemurs likely use
day caves to escape the extreme heat, as observed for Fongoli
chimpanzees (Pruetz 2001, 2007), it is important to distinguish
between day and night use of caves, as different pressures
(e.g., predation, ambient temperatures) likely vary throughout
the day. At Tsimanampesotse, the ILove troop used the small
cliff - face crevices and caves for sleeping without exception
during all the observations, and at all times of the year, from the
scorching Austral summer, where temperatures regularly range
above 40°C to the cool late Austral fall and early winter when
temperatures can fall as low as the single digits °C (LaFleur
2012). In addition, for the troops studied at Tsimanampesotse
only the ILove troop used caves and fissures for night sleeping,
while neither the Vintany nor the Akao troops did. As the ILove
and Vintany troops live only several hundred meters from each
other on top of the western escarpment of the Mahafaly Plateau,
they experience similar temperature conditions, and thus would
be expected to respond to these stressors in a similar way,
given the availability of numerous limestone depressions and
cliff-faces in Vintany’s range. Similarly, the Akao troop used
a large stand of Casuarina equisetifolia at the eastern edge
of the Lac Tsimanampesotse, along a marshy area each night
throughout the observations in May/June 2006. During that
period, temperatures at night dropped near the single digits °C,
with high humidity. Thus, these lemurs sleep under quite cold
conditions, yet have never been observed using caves along
the escarpment as night sleeping sites, despite their daily use
of this escarpment area for foraging. There are also a variety
of behavioral responses to temperature challenges. At Tsima-
nampesotse during periods of excessively high temperatures
(at or greater than 40°C) the lemurs spent considerable time
resting in the shade and licking their hands and feet to cool
off (LaFleur 2012). During especially cold mornings the ILove
troop left its caves at daylight and formed ‘lemur balls’ on the
limestone surface above their sleeping caves until the sun was
high enough to allow individual sunning. Similarly, at Tsinjoriake
the lemurs visited the mangroves during the hottest period of
the day, where they could rest and feed in the shade of these
larger trees, but did not use these trees to sleep overnight. Thus,
thermoregulation alone is unlikely to be the primary reason for
sleeping at night in caves and fissures.
As noted by Anderson (1998), primates spend about half of
their lives within sleeping sites, and where primates choose to
sleep is thus an important aspect of their behavioral ecology.
Numerous studies have indicated that sleeping sites appear to
be selected to provide some safety from predation (cf. reviews
in Anderson 1998, Bitetti et al. 2000, Cui et al. 2006, Cheyne et
al. 2012). Especially critical is a sleeping site that allows early
predator detection or provides difficult access for an approach-
ing predator. For example, both talapoin and tufted capuchin
monkeys choose sleeping areas where approaching predators
create noise and vibrations (Gautier-Hion 1970, Zhang 1995).
Sympatric Kloss’s gibbons (Hylobates klossi) and Mentawi
langurs (Presbytis potenziana) on Siberut Island, Indonesia, both
use tall emergent sleeping trees, but the gibbons preferentially
select trees with few large lianas. Human hunters, who use the
large lianas to climb into the trees, are thus able to kill far more
langurs than gibbons (Tenaza and Tilson 1985, Cheyne et al.
2012). Habitat structure is especially essential for understanding
variation in anti - predation behavior (Enstam 2007). For example,
patas monkeys on continental Africa live in open savannas with
few large sleeping trees and thus individuals have a dispersed
sleeping pattern with one individual sleeping in a tree. Within
the same habitat, African vervets will sleep in tall trees within
riverine forest habitats and thus all individuals may sleep in
one tree (Enstam 2007). Hamadryas baboons (Papio hamadryas)
living in the highlands of Ethiopia, are especially relevant here:
their habitat has few sleeping trees, but they solve this dilemma
by sleeping on sheer cliff - faces (Kummer 1968).
Goodman et al. (2006) were the first to suggest that the
use of limestone crevices and/or overhangs on vertical cliff-
faces as night sleeping sites by ring - tailed lemurs could be an
anti - predator strategy, specifically in response to endemic fossa
(Cryptoprocta ferox). Thus, another possible explanation for cave
use among the studied lemurs is that using crevices and caves
as night sleeping sites at Tsimanampesotse and Tsinjoriake may
provide some safety from night - active predators within a habitat
with a relative paucity of large trees for suitable sleeping sites.
Studies of ring - tailed lemurs in gallery forests document that
this species either chooses sleeping trees that are tall and have
broad dense canopies that are large enough to accommodate
the entire troop, or stands of tall trees that allow all individuals to
sleep near one another (Sauther et al. 1999). In over 25 years of

MADAGA SCAR CONSERVATION & DEVE LOPMENT VOLUME 8
|
ISSUE 2 — NOVEMBER 2013 PAG E 7 7
study at the Bezà Mahafaly Special Reserve, ring - tailed lemurs
have never been observed to use any night sleeping sites other
than large trees, even when troop size exceeds 20 individuals
(MLS, pers. observ.). Given the variety of known lemur predators
at Bezà Mahafaly Special Reserve, including Cryptoprocta ferox,
wild cats, feral dogs and a diversity of avian raptors (Sauther
1989, Gould and Sauther 2007) use of such sleeping trees may
be an anti - predator strategy, especially as ring - tailed lemurs
also exhibit predation sensitive foraging strategies (Gould and
Sauther 2007). That ring - tailed lemurs seek out safe night sleep-
ing sites is also supported at other sites. For example, two troops
of ring - tailed lemurs studied in the Cap Sainte - Marie region of
southern Madagascar used stands of introduced Opuntia s p.
as night sleeping areas, despite there being remnant stands
of larger trees in traditionally protected sacred forests (Kelley
2011). Here, remnant stands of sacred forests are very small in
area and widely dispersed, and ring - tailed lemurs were never
observed to use these as night sleeping areas except in one
instance, where the lemurs slept in a hedge of Opuntia sp. near
one of these sacred forests.
At Tsimanampesotse Ficus sp., Tamarindus indica and
Casuarina equisetifolia are the only trees large enough to
provide potential sleeping trees for a lemur troop. Ficus
sp. and T. indica are rare. Only 20 of the 255 trees on our
transects with a DBH > 10 cm were these species, and these
were typically found near the caves that form in limestone
depressions. C. equisetifolia is only found along the marshy
banks of Lac Tsimanampesotse. Unlike the degraded areas of
Cap Sainte Marie (Kelley 2011), there are no hedges of Opuntia
sp. at Tsimanampesotse. One habitat difference between the
Vintany, Akao and ILove troops at Tsimanampesotse is that
both Vintany and Akao troops have access to trees tall and
large enough to accommodate an entire ring-tailed lemur troop
and are thus suitable as sleeping sites. It is also important
to note that the stand of C. equisetifolia trees used by the
Akao lemurs is surrounded by thick marsh ferns of Acrosti-
chum aureum at the base of the trees, which are difficult and
noisy to move through, thus likely inhibiting terrestrial predator
access to these trees. Also, these lemurs have been observed
to use the plants to avoid avian predators, by quickly descend-
ing from the Casuarina equisetifolia into the ferns. The ILove
troop sleeps in the cliff - face crevices/caves that border a deep
limestone depression despite there being a few large T. indica
and Ficus sp. trees in their range. In August 2013, a new troop
was observed in a different area at Tsimanampesotse using a
large cave as their nightly sleeping site. Around this cave are a
number of large Ficus sp. trees, yet this group does not use them
as night sleeping sites. While a long - term study of ring - tailed
lemur cave use is needed, these observations indicate caves
are viewed as acceptable and, in some areas with larger trees,
even as preferred night sleeping sites. Recent observations
also indicate that ring - tailed lemurs may perceive sleeping
caves as important refugia from nocturnal predators but not
necessarily diurnal ones. In one case, as the ILove troop was
entering their caves and crevices in the evening, one indi-
vidual started alarm calling at a raptor flying over the caves. All
lemurs quickly came out of the cave and scanned the area for
the hawk. This is similar to anti - predator behaviors exhibited
by Bezà Mahafaly gallery forest ring - tailed lemurs, which will
orient towards avian prey and often mob them (Sauther 1989).
Both, the fossa (Cryptoprocta ferox) as well as wild cats
(Felis silvestris) have been documented to occur at Tsiman-
ampesotse and fecal analyses indicate fossa prey on ring - tailed
lemurs here (LaFleur 2012). While fossa are adept climbers, they
are not able to climb high canopy trees very well, due to their
large size (Hawkins 2003). At Tsimanampesotse the camera
trap data indicate that fossa can scale narrow tall trees, thus
rendering these types of trees useless as safe sleeping sites.
Also, the only known successful predation of a lemur during the
2010–2011 study period was on an infant in the Vintany group,
whose body was found beneath this troop’s sleeping tree, a
large broad Ficus sp. The infant clearly showed two large canine
puncture wounds on the neck (LaFleur 2012). Thus at Tsiman-
ampesotse, given that access to sleeping caves and crevices
requires scaling a vertical cliff - face, it likely does provide safety
from night - active fossa and nocturnal wild cat predation and
this may explain why some lemur troops use caves even though
larger trees may be nearby.
Further evidence of cave use for predator avoidance is
seen at Tsinjoriake. Important predators include packs of village
dogs as well as fossa. At this site there are only a few areas
that contain large trees. This includes the mangrove forest and
a small gallery forest as well as areas directly next to Grotte
Binabe and Grotte Binakely. Within both the mangrove forest and
the small gallery forest the majority of trees have a DBH of < 10
cm (mangrove: 92 % ; gallery forest: 64 % , Ravelohasindrazana
2013). There are large trees (> 10 cm DBH) around Grotte Binabe,
but these are few in number and are found only directly next
to the cave. At the Grotte of Binabe and Binakely the lemurs
regularly uses cliff - face crevices and caves for sleeping that
would be difficult for fossa and wild cats to scale. Similarly, the
Ekipa troop uses a group of caves that are located on the side
of a limestone cliff that vertically drops precipitously into the
ocean. The situation for the Ekipa troop is, however, slightly
more complex. Earlier observations (Scott et al. no date) noted
that, in addition to this cave, this group also used a stand of
Tamarindus indica trees by a hotel for sleeping. Work by the
research team in April 2012 also noted this. However, in June
2012, one month of camera trapping in this area showed no
images of lemurs there. Most recently, in July/August 2013, the
local research guide informed the research team that the Ekipa
troop is no longer using these tamarind trees due to a large and
growing pack of domestic dogs in the area, which apparently
blocks the terrestrial access route to this fragmentary stand of
trees. Thus, the Ekipa troop appears to now exclusively use the
cave located on the limestone cliffs for sleeping, presumably to
avoid these introduced predators. Given the long coexistence
of L. catta and endemic predators in southwestern Madagascar,
it is suggested that cliff - face and cave use is a natural part of
the L. catta behavioral repertoire and is then available when
needed, such as in the highland region in response to human
actions (i.e., deforestation).
This novel behavior by ring - tailed lemurs expands the
knowledge of primate cave use, previously reported in detail
only for anthropoid primates, and when viewed through the lens
of human evolution provides a broader context to interpret cave
use by human ancestors (e.g., McGrew et al. 2003, Pruetz 2007).
These data also aid the interpretation of cave use in primate
paleobiology. For example, a recent study found fecal pellets in
Anjohikely Cave in northern Madagascar, attributed to extinct

MADAGA SCAR CONSERVATION & DEVE LOPMENT VOLUME 8
|
ISSUE 2 — NOVEMBER 2013 PAGE 7 8
Archaeolemur (Vasey et al. 2013), suggesting that this large,
semi - terrestrial lemur may have used the cave for shelter and
nutrients. A clearer understanding of cave use in extant lemurs
could thus expand the understanding of the paleoecology of
this extinct species. The ongoing collaborative work on these
ring - tailed lemur populations will allow the testing of various
hypotheses regarding primate cave use.
Tsimanampesotse is one of the only protected areas of
Cenozoic limestone habitats on the Mahafaly Plateau (DuPuy
and Moat 1998). These authors also noted the importance of
other areas of the Mahafaly Plateau, specifically along the
escarpment edge towards Saint Augustin, which today encom-
passes the new Tsinjoriake Protected Area. However, we are
just beginning to understand the ecology of ring - tailed lemurs
in spiny forest habitats (e.g., Kelley 2011, LaFleur 2012) as most
studies of ring-tailed lemurs have been in gallery forest habitats
(Sauther 1999, Gould 2007). Given the importance of limestone
caves for safe shelter, nutrition and drinking sites among ring-
tailed lemurs it is essential that these habitats remain intact.
However, the limestone cliffs and southwestern dry spiny forest
thicket as well as coastal mangroves of southwestern Mada-
gascar are currently under intense human pressure, including
the development of limestone mining operations such as Gulf
Industrial Limited’s Soalara Limestone Project near the mouth
of the Onilahy River. Ring - tailed lemurs, recently classified
as an endangered species by the International Union for the
Conservation of Nature (IUCN) Species Survival Commission
(SSC), are threatened by deforestation and habitat degradation
throughout southwestern Madagascar. Tsimanampesotse can
now be added as a habitat undergoing extensive anthropo-
genic change. As recently as August, 2013 trees were being
actively harvested as cattle forage, construction materials, and
for carving pirogues. Based on the discovery of forest traps
and communications with local people, ring - tailed lemurs
are also now actively hunted in the area. Until now, poaching
of ring - tailed lemurs (and other mammals) in southwestern
Madagascar has remained relatively rare, due to widespread
traditional taboos. However, rapid social change, along with
the well - established existing black - market trade in radiated
tortoises (Astrochelys radiata) and the demand for alternative
or luxury bushmeat in Madagascar’s cities have resulted in a
niche for harvesting ring - tailed lemurs. Similarly, Tsinjoriake
is especially affected by its close location to the large city of
Toliara. For example, questionnaires indicate that 14 of the
39 tree species used by ring - tailed lemurs are also used to
produce charcoal in the area including Tamarindus indica
and Ficus sp. (Ravelohasindrazana, 2013, Ravoavy, 2013). The
questionnaires also indicate that adult ring - tailed lemurs are
hunted by humans (using dogs) as a food resource. Infant ring-
tailed lemurs become pets and are sold in Toliara for between
3,000 and 4,000 Malagasy Ariary (about $US1.36–1.81) while
smoked ring - tailed lemurs sell for 4,000–5,000 Malagasy Ariary
($US1.77–2.21). One of the biggest threats to this protected area
is the collection of firewood to fuel the production of bricks at
Ankoronga. It is estimated that the production of bricks in the
village consumes 1,200–1,500 cart - loads of dead wood per
month (Andriamahafaly 2010). The Tsinjoriake New Protected
Area has recently been established as a community - based
ecological reserve (initially developed under the auspices
and guidance of the Programme Germano - Malgache pour
l’Environnement Coopération Allemande / GIZ, a European NGO
that provided funds for local development of this reserve. GIZ/
GTZ also laid the groundwork for local community development).
The continued support of the Tsinjoriake reserve and community
development project now rests on local Malagasy researchers,
scientists and community developers, who are responsible for
obtaining funds to fully implement the project. Towards this
end, JY and the University of Toliara are working with GIZ/GTZ
and the local community TAMIA to facilitate community devel-
opment and training to enable sustainable development and
energy use at Tsinjoriake.
These areas have rarely been included in long - term lemur
(and other faunal) research and conservation. Although not
as speciose (in terms of lemurs) as rainforest habitats, with
only Lemur catta and Microcebus griseorufus observed by the
authors, these coastal areas are rich in endemic plant, bird and
reptile species and do hold important information for under-
standing lemur evolution and the presence of unique lemur
traits (e.g., LaFleur 2012). As such, they deserve immediate
conservation attention.
ACKNOWLEDGEMENTS
We thank the many local and international students and field
assistants without whom our work would not be possible.
We thank MNP (formerly ANGAP) for permission to work at
Tsimanampesotse National Park, Programme Germano-
Malgache pour l’Environnement Coopération Allemande /GIZ,
the University of Toliara and the TAMIA (Tontolo Alandriake
Mitambatse Ianatsono Andatabo) local cooperative for their
support of our work at Tsinjoriake. We also thank the three
MCD reviewers whose comments greatly improved our
paper. Funding for this work has been provided by: Primate
Conservation Inc., the International Primatological Society, the
American Society of Primatologists, the National Geographic
Society, the University of Colorado-Boulder (CRCW; IGP;
Outreach Grant), the University of North Dakota (UND Arts,
Social Sciences and Humanities fund; Faculty Seed Money
Council; SSAC), ND EPSCoR, Colorado College, NSERC PGS
296264, and the National Science Foundation (BCS 0922645,
DDIG 1028708).
REFERENCES
Anderson, J. R. 1998. Sleep, sleeping sites and sleep-related activities:
Awakening to their significance. American Journal of Primatology 46,
1: 63–75.
Andriamahafaly, M. N. 2010. Plan d’Aménagement et de Gestion de la
Nouvelle Aire Protégée Communautaire de Tsinjoriake, Toliara, 135pp.
Unpubl. Report.
Andriatsimietry, R., Goodman, S. M., Razafimahatratra, E., Jeglinski, J. W. E.,
Marquard, M., and Ganzhorn, J. U. 2009. Seasonal variation in the diet of
Galidictis grandidieri Wozencraft, 1986 (Carnivora: Eupleridae) in a sub-
arid zone of extreme south-western Madagascar. Journal of Zoology
279, 4: 410–415. (doi:10.1111/j.1469-7998.2009.00633.x)
Barrett, L., Gaynor, D., Rendall, D., Mitchell, D. and Henzi, S. P. 2004 Habitual
cave use and thermoregulation in chacma baboons (Papio hamadryas
ursinus). Journal of Human Evolution 46, 2: 215–222. (doi:10.1016/j.
jhevol.2003.11.005)
Cameron, A. and Gould, L. 2013. Fragment-Adaptive behavioural strate-
gies and intersite variation in the ring-tailed lemur (Lemur catta) in
South-Central Madagascar. In: Primates in Fragments. Developments in
Primatology: Progress and Prospects. L. K. Marsh and C. A. Chapman
(eds.), pp 227–243. Springer, New York, NY. (doi:10.1007/978-1-4614-
8839-2_16)

MADAGA SCAR CONSERVATION & DEVE LOPMENT VOLUME 8
|
ISSUE 2 — NOVEMBER 2013 PAGE 7 9
Cheyne, S. M., Höing, A., Rinear, J. and Sheeran, L. K. 2012. Sleeping site
selection by Agile gibbons: The influence of tree stability, fruit
availability and predation risk. Folia Primatologica 83: 299–311.
(doi:10.1159/000342145)
Cui, L.-W., Quan, R.-C. and Xiao, W. 2006. Sleeping sites of black-and-white
snub nosed monkeys (Rhinopithecus bieti) at Baima Snow Mountain,
China. Journal of Zoology 270, 1:192–198. (doi:10.1111/j.1469-
7998.2006.00137.x)
Cuozzo, F.P. and Sauther, M.L. 2012. The dental ecology of ring-tailed lemurs
(Lemur catta). In: Leaping Ahead: Advances in Prosimian Biology. J. C.
Masters, M. Gamba, M. and F. Génin (eds.), pp 157–163. Springer, New
York, NY.
Di Bitetti, M. S., Luengos, E. M., Baldovino M. C. and Benesovsky V. 2000.
Sleeping site preferences in tufted capuchin monkeys (Cebus apella
nigritus). American Journal of Primatology 50, 4: 257–274. (doi:10.1002/
(SICI)1098-2345(200004)50:4<257::AID-AJP3>3.0.CO;2-J)
Donque, G. 1975. Contribution à l’Étude du Climat de Madagascar. Nouvelle
Imprimerie des Arts Graphiques, Tananarive.
Du Puy, D. J. and Moat, J. F. 1998. Vegetation mapping and classification in
Madagascar (using GIS): implications and recommendations for the
conservation of biodiversity. In: Chorology, Taxonomy and Ecology of
the African and Madagascan Floras. C. R. Huxley, J. M. Lock and D. F.
Cutler D.F. (eds.), pp 97–117. Royal Botanic Gardens, Kew.
Enstam, K. L. 2007. Effects of habitat structure on perceived risk of preda-
tion and anti-predator behavior of vervet (Crecopithecus aethiops)
and patas (Erythrocebus patas) monkeys. In: Predation in Primates.
S. Gursky S. and A. Nekaris (eds.), pp 308–338. Cambridge University
Press, Cambridge.
Gautier-Hion, A. 1970. L’organisation sociale d’une bande de talapoins
(Miopithecus talapoin) dans le nord-est du Gabon. Folia Primatologica
12, 2: 116–141. (doi:10.1159/000155285)
Goodman, S. M. and Langrand, O. 1996. A high mountain population of the
ring-tailed lemur (Lemur catta) on the Andringitra Massif, Madagascar.
Oryx 30, 4: 259–268. (doi:10.1017/S003060530002175X)
Goodman, S. M., Rakotoarisoa, S. V. and Wilmé, L. 2006 The distribution and
biogeography of the Ringtailed lemur (Lemur catta) in Madagascar. In:
Ringtailed Lemur Biology Developments in Primatology: Progress and
Prospects. A. Jolly, N. Koyama, H. Rasamimanana. and R. W. Sussman
(eds.), pp 3–15. Kluwer Academic/Plenum, NY.
Gould, L. 2007. Lemur catta ecology: What we know and what we need to
know. In: Lemurs: Ecology and Evolution. L. Gould and M. L. Sauther
(eds.), pp 257–276. Springer, New York, NY.
Gould, L. and Sauther, M. L. 2007. Anti-predator strategies in a semi-terres-
trial diurnal prosimian, the ring-tailed lemur (Lemur catta), at the Beza
Mahafaly Special Reserve, Madagascar. In: Predation in Primates. S.
Gursky S. and A. Nekaris, A. (eds.), pp 275–288. Cambridge University
Press, Cambridge.
Hawkins, C. E. 2003. Cryptoprocta ferox, Fossa. In: The Natural History of
Madagascar. S. M. Goodman and J. P. Benstead (eds.), pp 1360–1363.
The University of Chicago Press, Chicago.
Huang, C., Wei, F., Li, M., Li, Y. and Sun, R. 2003 Sleeping cave
selection, activity pattern and time budget of white-headed
langurs. International Journal of Primatology 24, 4:813–824.
(doi:10.1023/A:1024628822271)
Jolly, A. 1966. Lemur Behavior. The University of Chicago Press, Chicago.
Jolly, A., Sussman, R. W., Koyama, N. and Rasmimanana, H. (eds.). 2006.
Ringtailed Lemur Biology. Springer, New York, NY.
Kelley, E. A. 2011. Lemur catta in the Region of Cap Sainte-Marie, Madagascar:
Introduced Cacti, Xerophytic Didiereaceae-Euphorbia Bush, and Tombs.
Unpublished Ph.D. Thesis. Washington University, St. Louis, MO.
Kummer, H. 1968. Social organization of hamadryas baboons. The University
of Chicago Press, Chicago.
LaFleur, M. 2012. Ecology of Ring-tailed Lemurs (Lemur catta) at the
Tsimanampetsotsa National Park, Madagascar: Implications for Female
Dominance and the Evolution of Lemur Traits. Unpublished Ph.D.
Thesis. University of Colorado-Boulder. Published online at http://
vetmeduni.academia.edu/MarniLaFleur
Li, Z. and Rogers, M. E. 2005. Are limestone hills a refuge or essential habi-
tat for whiteheaded langurs in Fusui, China? International Journal of
Primatology 26, 2: 437–452. (doi:10.1007/s10764-005-2932-y)
Loudon, J. E., Sauther, M. L. Fish, K. D., Hunter-Ishikawa, M. and Jacky
Youssouf, I. 2006. One reserve, three primates: applying a holistic
approach to understand the interconnections among ring-tailed
lemurs (Lemur catta), Verreaux’s sifaka (Propithecus verreauxi),
and humans (Homo sapiens) at Beza Mahafaly Special Reserve,
Madagascar. Ecological and Environmental Anthropology 2, 2: 54–74.
Published online at http://www.uga.edu/eea
McGrew, W. C., McKee, J. K. and Tutin, C. E. G. 2003. Primates in caves: two
new reports of Papio spp. Journal of Human Evolution 44, 4: 521–526.
(doi:10.1016/S0047-2484(03)00042-3)
Perrier de la Bâthie, H. 1927. Le Tsaratanana, l’Ankaratra el l’Andringitra.
Mémoires de l’Académie Malgache 3: 1–71.
Pruetz, J. D. 2001. Use of caves by savanna chimpanzees (Pan troglodytes
verus) in the Tomboronkoto region of southeastern Senegal. Pan
Africa News 8, 2: 26–28.
Pruetz, J. D. 2007. Evidence of cave use by savanna chimpanzees (Pan
troglodytes verus) at Fongoli, Senegal: implications for thermoregula-
tory behavior. International Journal of Primatology 48: 316-319.
(doi:10.1007/s10329-007-0038-1)
Ravelohasindrazana, L. A. L. 2013. Etude de la population Lemur catta de la
NAP Tsinjoriake site d’Antsifotse. Mémoire de fin d’étude en vue de
l’obtention du diplôme de Licence Professionnelle en Biodiversité et
Environnement (5e promotion). Option: Biologie Animale. Université
de Toliara, Faculté de Science.
Ravoavy, J. F. 2013. Etude de la Population de Lemur catta (Maki), dans la
Nouvelle Aire Protégée Tsinjoriaky, Région Sud-Ouest de Madagascar.
Mémoire en vue de l’obtention du Diplôme de Licence Professionnelle
en Biodiversité et Environnement. Université de Toliara, Faculté de
Science.
Sauther, M. L. 1989. Antipredator behavior in troops of free-ranging Lemur
catta at Beza Mahafaly special reserve, Madagascar. International
Journal of Primatolology 10, 6: 595–606. (doi:10.1007/BF02739366)
Sauther, M. L., Sussman, R. W. and Gould, L. 1999. The socio-
ecology of the ring-tailed lemur: thirty-five years of research.
Evolutionary Anthropology 8, 4:120–132. (doi:10.1002/(SICI)1520-
6505(1999)8:4<120::AID-EVAN3>3.0.CO;2-O)
Sauther, M. L. and Cuozzo, F. P. 2008. Somatic variation in living, wild
ring-tailed lemurs (Lemur catta). Folia Primatologia 79: 55–78.
(doi:10.1159/000108589)
Scott, D., Annael, O., Mbohoahy, T., Raharinirina, N. L., Smith, M.,
Tanteliniaina, S., Thompson, S. and Tsagnangara, C. No Date. University
of Brighton (UOB) Spiny Forest Research Expedition 2007/2008 Final
Report. Unpubl. report to University of Brighton.
Tenaza, R. and Tilson R. 1985. Human predation and Kloss›s gibbon
(Hylobates klossii) sleeping trees in Siberut Island, Indonesia.
American Journal of Primatology 8, 4: 299–308. (doi:10.1002/
ajp.1350080405)
Vasey, N., Burney, D. A. and Godfrey, L. 2013. Coprolites associated with
Archaeolemur remains in North-western Madagascar suggest dietary
diversity and cave use in a subfossil prosimian. In: Leaping Ahead:
Advances in Prosimian Biology. J. Masters, M. Gamba and F. Génin
(eds.), pp 149–156. Springer New York, NY.
Video 1: <http://www.youtube.com/watch?v=8qnBWSMqF_I&feature=youtu.
be>
Video 2: <http://www.youtube.com/watch?v=mf_DSTCDyfI&feature=youtu.
be>
Wilson, J. M., Stewart, P. D., Ramangason, G.-S., Denning, A. M. and
Hutchings, M. S. 1989. Ecology and conservation of the crowned lemur,
Lemur coronatus, at Ankarana N. Madagascar. Folia Primatologica 52,
1–2: 1–26.
Workman, C. 2010. The foraging ecology of the Delacour’s langur
(Trachypithecus delacouri) in Van Long Nature Reserve, Vietnam.
Unpublished Ph.D. Thesis. Duke University. http://dukespace.lib.
duke.edu/dspace/bitstream/handle/10161/2457/D_Workman_
Catherine_a_201005.pdf?sequence=1
Zhang, S.-Y. 1995. Sleeping habits of brown capuchin monkeys (Cebus
apella) in French Guiana. American Journal of Primatology 36, 4:
327–335. (doi:10.1002/ajp.135036040)
Zafisamimanana, L. M. 2012. Cartographie de l’Occupation du Sol dans la
NAP Tsinjoriake. Unpubl. Report.

MADAGA SCAR CONSERVATION & DEVE LOPMENT VOLUME 8
|
ISSUE 2 — NOVEMBER 2013 PAG E 8 0
SUPPLEMENTARY MATERIAL.
AVAILABLE ONLINE ONLY.
Video clip showing Ilove troop lemurs at Tsimanampesotse scal-
ing vertical limestone cliff and entering sleeping caves. Video
by Michelle Sauther.
FIGURE S1. Lemur from the Akao troop in Casuarina equi-
setifolia tree on lake margin (A) and within the marshy ferns
of Acrostichum aureum at the base of these trees (B) at
Tsimanampesotse National Park, 2006.
FIGURE S2. Lemurs from the Grotte de Binabe troop on the cliff-
face at Tsinjoriake Protected Area.
FIGURE S3. Cave above Sarodrano peninsula, where the “Ekipa”
troop sometimes sleeps.
FIGURE S4. Spiny forest habitat at Tsinjoriake. The arrow is point-
ing at a tree that is 2 m. tall.
FIGURE S5. ILove troop ring - tailed lemurs huddling in a “lemur
ball” on limestone rocks after leaving sleeping caves at
Tsimanampesotse National Park.