First record of Fossa Cryptoprocta ferox in Mariarano Forest, Madagascar

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Small Carnivore Conservation 52 & 53: 45–55
ORIGINAL ARTICLE
First record of Fossa Cryptoprocta ferox in Mariarano forest,
Madagascar
Gareth Kerry Hamilton MANN1, Peter LONG2, Felix RAKOTONDRAPARANY3, Sam THE
SEING4 & Robert GANDOLA5
1 Department of Zoology and
Entomology, Rhodes University,
PO Box 94, Grahamstown 6140,
South Africa.
2 Biodiversity Institute,
Department of Zoology,
Tinbergen Building, South Parks
Road, Oxford, OX1 3PS, UK.
3 Département de Biologie
Animale, Faculté des Sciences,
Université Antananarivo. PO
Box 906, Antananarivo 101,
Madagascar.
4 DBCAM, Lot II A 93L,
Anjanahary, Antananarivo 101,
Madagascar.
5 School of Ocean and Earth
Science, National Oceanography
Centre, University of
Southampton, Waterfront
Campus, Southampton, SO14
3ZH, UK.
Correspondence:
Gareth K. Hamilton Mann
gmann9@gmail.com
Associate editor:
Frank Hawkins
http://www.smallcarnivoreconservation.org
ISSN 1019-5041
Introduction
Madagascar is the worldʼs fourth-largest island and a global biodiversity hotspot with an
abundance of endemic mammalian fauna (Myers et al. 2000, Mittermeier et al. 2005).
However, many of these species are threatened by habitat loss and fragmentation (Irwin et
al. 2010), as Madagascarʼs indigenous forest cover has been reduced by an estimated
43.85% from the 1950ʼs to the year 2000 (Harper et al. 2007). Mammalian carnivores tend
to be especially vulnerable to habitat loss and fragmentation due to their relatively large
Abstract.
We surveyed the carnivore community in two patches of forest around
Mariarano village in north-western Madagascar using camera traps. Cameras
were set along trails in the forest and were active for a total of 517 trap
nights. We recorded the presence of two indigenous carnivore species, Fossa
Cryptoprocta ferox and Western Falanouc Eupleres major, and three
introduced carnivore species; Small Indian Civet Viverricula indica, domestic
dogs Canis familiaris and domestic cats Felis catus. This is the first record of
C. ferox in the Mariarano forest area. We discuss the significance of this
finding, as well as a potential extirpation of E. major in the Matsedroy forest
patch.
Keywords: Camera trapping, deciduous forest, Eupleres major, Eupleridae
Premier signalement de Fossa Cryptoprocta ferox dans la forêt
Mariarano, Madagascar
Résumé.
Nous avons sondé la communauté de carnivores dans deux parcelles de forêt
autour du village de Mariarano dans le nord-ouest de Madagascar à lʼaide de
pièges photographiques. Des caméras ont été fixées le long des sentiers dans
la forêt et ont été actives pour un total de 517 nuits de piégeage. Nous avons
enregistré la présence de deux espèces indigènes de carnivores, le Fossa
Cryptoprocta ferox et le Falanouc occidental Eupleres major, et trois espèces
de carnivores introduits; la Petite civet indienne Viverricula indica, la chien
domestique Canis familiaris et le chat domestique Felis catus. Ceci est le
premier enregistrement de C. ferox dans la zone de la forêt Mariarano. Nous
discutons de la signification de cette découverte, ainsi que dʼune disparition
potentielle de E. major dans la parcelle de forêt Matsedroy.
Mots clés: Eupleres major, Eupleridae, forêt de feuillus, piégeage-caméra

Cryptoprocta ferox in Mariarano forest, Madagascar
Small Carnivore Conservation 52 & 53: 45–55 46
spatial requirements (Woodroffe & Ginsberg 1998). In Madagascar, this situation is
particularly acute, as relatively few studies have been done on the endemic carnivore
species. This lack of data has resulted in carnivores being excluded from formal
conservation plans for Madagascar (Kremen et al. 2008). Improved knowledge of the
distribution, habitat preferences and disturbance tolerance of Madagascarʼs indigenous
carnivores is thus of critical importance to their future conservation.
Our study presents results of a camera trapping survey conducted in remnant patches
of western dry deciduous forest around Mariarano village located approximately 50 km
north-west of Mahajanga in western Madagascar and builds on an existing dataset (see
Evans et al. 2013). These forest patches are not formally protected, and are threatened by
illegal timber extraction, charcoal production and clearing for agriculture (Washington et
al. 2009, Long et al. 2012). Nevertheless, the forests around Mariarano contain a wide
variety of lemur, reptile and bird species, including threatened species such as Coquerel's
sifaka Propithecus coquereli, Leaf-tailed Geckos Uroplatus sp. and Madagascar Fish Eagle
Haliaeetus vociferoides.
Since 2009, a long-term monitoring project of the areaʼs biodiversity has been
running collaboratively by Operation Wallacea, an international NGO, Development and
Biodiversity Conservation Action for Madagascar, a community-based Malagasy
conservation NGO, local community forest management groups, and the University of
Antananarivo. Biodiversity surveys are done annually from June to August, during the local
dry season. While most species are readily observed by the multidisciplinary teams who
undertake the monitoring, indigenous carnivores are cryptic and seldom directly observed.
Camera traps have been found to be the most effective means of gathering information on
carnivore species at Mariarano (Evans et al. 2013). Our study aimed to gather data on
carnivores in the Mariarano forest as part of the ongoing monitoring programme.
Materials and methods
Study areas
We sampled two discrete patches of forest in the vicinity of Mariarano village (see Figure
1). Mariarano forest (also known as Ankatsabe forest) borders Mariarano village on three
sides (North, East and South), while the area to the west of Mariarano village has been
cleared for cultivation. Sampling routes around Mariarano therefore sampled the western
and central portions of Mariarano forest. In 2014, new sampling routes were demarcated to
the west of Antafiemeva, a village on the eastern fringe of Mariarano forest. These new
routes allowed for systematic sampling of the eastern side of Mariarano forest.
We also sampled in the area around Matsedroy Station, located in Matsedroy forest
(also known as Analabe forest), which is located approximately 5 km to the west of
Mariarano village. Matsedroy forest is separated from Ankatsabe by a broad strip of

Mann et al.
47 Small Carnivore Conservation 52 & 53: 45–55
cultivated land on either side of the Mariarano River. Matsedroy forest was noticeably more
degraded than Mariarano forest, with signs of recent deforestation evident (Ibouroi et al.
2013).
Figure 1. Map of the study area showing the position of the study area in Madagascar (inset), and
locations of camera stations used in 2014. Forest patches are indicated by dark grey, intermediate-
grey areas are wetland and riverine areas, while light grey represents open savannah areas with little
tree cover.
Methods
We set 22 unbaited Bushnell TrophyCam HD camera traps along the existing network of
seven sampling routes in the Mariarano forest, and four sampling routes in Matsedroy
forest (Figure 1). Survey routes range from 1.7 to 3.6 km in length, and typically two
cameras were placed on each survey route a minimum of 1 km apart. Camera sites were
chosen on the basis of evidence of terrestrial mammal activity (i.e., tracks and scats), as
well as the advice of local guides. Two of the shorter routes (length ~1.6 km) only
contained single camera sites, while additional cameras were positioned on two other,
relatively long, sampling routes. A further two cameras were placed in opportunistic
locations away from sampling routes. All cameras were set with their sensors
approximately 30 cm high, and were programmed to record bursts of three photographs,
with a 10-second gap in between capture events. We collected data from late June to late
July 2014. The GPS coordinates of each camera site were recorded using a Garmin

Cryptoprocta ferox in Mariarano forest, Madagascar
Small Carnivore Conservation 52 & 53: 45–55 48
GPSMAP 62s. Sampling effort was measured in trap nights; i.e., a 24-hour period from
midday to midday.
All animals recorded by the camera traps were identified to species level, and these
data were entered into a spreadsheet together with the date, time and camera station at
which the animal was recorded. Domestic animals were not included in our analyses, apart
from domestic cats Felis catus and dogs Canis familiaris that did not appear to be
accompanied by people. Dogs accompanied by people were assumed to be under human
control, and therefore less likely to actively hunt wildlife, both due to social taboos, or
faddy (Jones et al. 2008) and the dogs being fed rather than needing to hunt wildlife to
survive. We assumed that repeated captures of the same species within one hour at a
camera station were non-independent recaptures of the same individual, and therefore
excluded these from subsequent analyses. Species accumulation curves were plotted for the
area as a whole and for the individual forest patches using EstimateS version 9.2 (Colwell
2006). If the species accumulation curve did not reach an asymptote we used the
Abundance Coverage Estimator (ACE) to estimate the total number of species likely to be
present in the area (Chazdon et al. 1998).
Results
We obtained data from 20 camera traps that were active for a total of 517 trap nights, with
each station active for a mean period of 25.85 (± SD) trap nights (± 6.39 days). Five camera
traps were stolen during the course of the study, and no data were obtained from four of
these. We recorded 78 independent captures of wildlife, of which 41 were of feral domestic
cats and dogs. Cats were recorded 21 times at nine sites across the study area, while dogs
were recorded 20 times at eight sites. Bushpigs Potamochoerus larvatus were the most
frequently photographed wildlife species, recorded 25 times across seven sites. Five
carnivore species were recorded in total; all five were present in Mariarano forest, but only
three were recorded in Matsedroy forest. Carnivore capture records are summarised in
Table 1. Species accumulation curves reached asymptote for Matsedroy forest (ACE =
3.00), but not for Mariarano forest (ACE = 5.41) or the area as a whole (ACE = 6.11),
suggesting that overall camera trapping effort was insufficient to record all carnivore
species in the area. Evans et al. (2013) reported local familiarity with Ring-tailed
Mongoose Galidia elegans, and it is possible that this species is present in the area but was
not recorded. Detailed records for wild carnivores are provided below:
Fossa Cryptoprocta ferox
Fossa are classified as Vulnerable on the IUCN Red List of Threatened Species (Hawkins
& Dollar 2008). A single photograph of a C. ferox was recorded south of Mariarano village
at 05h30 on 11 July 2014 on a portion of ox cart track that intersected the sampling route
within an area of secondary forest (see Figure 2). This is the first confirmed record of C.

Mann et al.
49 Small Carnivore Conservation 52 & 53: 45–55
ferox in the Mariarano forest, although previous studies have suggested that they are likely
to be present in the area (Long et al. 2012, Evans et al. 2013). This record does not
constitute a range extension for C. ferox, which are thought to be widely distributed
throughout low-altitude areas of Madagascar (Hawkins & Dollar 2008). Nevertheless, the
nearest published record of C. ferox is 80 km away at Ankarafantsika National Park (Dollar
et al. 2007, Garbutt 2007).
Table 1. Summary of carnivore camera trap data from the Mariarano forest, Madagascar, collected
during the dry season in 2014.
Camera
South
East
Effort1
Species2
MA101
15°28′36′′
46°42′06′′
21
C. familiaris (5)
MA102
15°28′06′′
46°42′18′′
0
Camera stolen
MA201
15°29′36′′
46°42′18′′
33
F. catus (1)
MA301
15°30′06′′
46°42′42′′
31
E. major (2), C. familiaris (5), F. catus (2)
MA302
15°30′3′′
46°43′12′′
31
E. major (2), V. indica (2), C. familiaris (1)
MA303
15°30′18′′
46°42′30′′
31
C. ferox (1), V. indica (3), F. catus (1), C. familiaris (5)
MA401
15°28′24′′
46°41′30′′
0
Camera stolen
MA402
15°29′00′′
46°42′00′′
0
Camera stolen
MA501
15°28′06′′
46°41′36′′
0
Camera stolen
MA502
15°27′24′′
46°41′30′′
30
AN101
15°28′12′′
46°44′36′′
31
F. catus (2), C. familiaris (1)
AN102
15°29′24′′
46°44′18′′
31
E. g. major (2), V. indica (1), F. catus (4)
AN201
15°28′12′′
46°43′30′′
30
AN202
15°28′48′′
46°43′24′′
30
MD101
15°29′24′′
46°38′30′′
23
MD102
15°29′36′′
46°38′12′′
23
F. catus (3), C. familiaris (1)
MD201
15°28′48′′
46°38′36′′
24
F. catus (3)
MD202
15°28′48′′
46°38′36′′
24
F. catus (3)
MD203
15°28′24′′
46°38′06′′
24
MD301
15°28′12′′
46°38′24′′
24
MD401
15°28′06′′
46°39′12′′
24
MD402
15°27′48′′
46°39′00′′
24
V. indica (1), F. catus (4)
MDOPP1
15°28′48′′
46°39′00′′
24
V. indica (2), F. catus (1), C. familiaris (1)
MDOPP2
15°28′54′′
46°39′00′′
4
V. indica (2)
1Effort refers to the number of trap nights for which the camera was active. The number of independent captures
recorded for each species per camera station is shown in brackets after the species name.
2The numbers in parentheses correspond to the number of records.
Western Falanouc Eupleres major
Eupleres major is classified as Endangered in the current IUCN Red List (Dollar 2000).
Individuals of this species were recorded six times at three sites. Two records, four days
apart, were obtained from a site near Antafiemeva village, at 01h44 and 19h10,
respectively. The remaining four records were obtained from two sites located south of
Mariarano village, both in secondary forest along a track that ran parallel to the main road
running south-east from Mariarano. Despite the relatively close proximity of these two
sites, records were obtained on four different days, suggesting that these were independent
capture events. Aside from one record at 19h10 (Figure 3), all E. major pictures were
recorded within a period of less than one hour, from 01h44 to 02h39, suggesting a possible
peak of activity during this time. Other studies have shown E. goudotii to be predominantly
nocturnal (Gerber et al. 2012a), and our results suggest that this trend holds true for the E.

Cryptoprocta ferox in Mariarano forest, Madagascar
Small Carnivore Conservation 52 & 53: 45–55 50
major population of Mariarano forest. Published records of E. major are scarce, but the
species range is thought to extend from Antsiranana at the northern tip of Madagascar to
close to Soalala on the west coast, at elevations ranging from 10 to 1,500 m (Goodman &
Helgen 2010). Eupleres major has previously been recorded in the Mariarano Forest area
(Evans et al. 2013), as well as approximately 70 km south-east of Mariarano at Marovoay
and near Port Bergé, approximately 100 km to the east (Goodman & Helgen 2010).
Figure 2. Camera trap image of a Fossa Cryotoprocta ferox (left) recorded in the Mariarano forest,
Madagascar, in July 2014. Although only the rear half of the animal was captured, the long tail is
sufficient to identify the subject as C. ferox.
Small Indian Civet Viverricula indica
Viverricula indica is classified as Least Concern in the latest IUCN Red List (Choudhury et
al. 2015), but it is not native to Madagascar (Garbutt 2007). This species was the most
frequently photographed and widely distributed wild carnivore within the Mariarano Forest
complex; it was recorded on 12 occasions at six sites spread across the study area. One
individual was recorded by a camera set on a track in recovered secondary forest near
Antafiemeva village at the easternmost extent of the study area. Six captures were recorded
at two sites in secondary forest south of Mariarano village near the centre of the study area.
One of these sites was on a major road frequently used by local people, both on foot and by
ox cart, while the other was in a more isolated location near the forest edge to the south.
Small Indian Civets were also recorded on the western side of the study are at three sites
near Matsedroy research station. All the cameras that recorded V. indica in this area were
situated in open areas or severely degraded secondary forest.

Mann et al.
51 Small Carnivore Conservation 52 & 53: 45–55
Figure 3. Camera trap image of a Western Falanouc Eupleres major recorded in the Mariarano
forest, Madagascar, in July 2014
Discussion
This study provides further detail following the initial assessment of the wild carnivores of
the Mahamavo forest (Evans et al. 2013), recording C. ferox in the area for the first time, as
well as the Western Falanouc at a further three locations. The discovery of C. ferox in the
area is interesting, given the isolation of the remnant forest at Mariarano from other patches
of western dry deciduous forest (Moat & Smith 2007, Long et al. 2012). Fossa population
density estimates range between 0.18 and 0.26 individuals per 100 km2 in Kirindy Forest,
another patch of Western dry deciduous forest (Hawkins & Racey 2005). Extrapolation of
these estimates to the Mariarano forest would suggest a population of between 12 and 17 C.
ferox individuals. However, these figures are likely to be overestimates, in that the
Mariarano forest area also contains sizeable tracts of agricultural land, as well as a number
of villages and smaller settlements, all of which are likely to adversely influence C. ferox
abundance (Gerber et al. 2012b). Regardless, it can be safely assumed that the Mariarano
forest C. ferox population falls well below the often-accepted threshold of 500 individuals
required for a population to be viable in the long term (Thomas 1990). This suggests that
the Mariarano forest C. ferox population is either a remnant population on the verge of
extirpation, or that it forms part of a larger metapopulation that may include the nearest
known population at Ankarafantsika National Park. However, Fossas̕ are thought to be
relatively intolerant of disturbed habitats, preferring to remain close to forests (Gerber et al.
2012b, Kotschwar Logan et al. 2014). C. ferox presence has been suspected in Matsedroy
forest since 2010, when local guides from Mariarano village claimed to have detected one

Cryptoprocta ferox in Mariarano forest, Madagascar
Small Carnivore Conservation 52 & 53: 45–55 52
while leading a research group, and guides have consistently claimed that C. ferox is
present in the Mariarano area. However, when findings of the 2014 survey were presented
to the GIZ Boeny and Tanteraka (i.e., local council), members stated that they were
unaware of C. ferox presence in the area. This lends further credence to the notion that C.
ferox densities are low in the area.
There was frequent overlap between carnivores at camera sites. The camera site
where we recorded C. ferox also obtained records of V. indica, C. familiaris and F. catus.
Similarly, introduced carnivores were also present at all three sites at which E. major were
recorded. It is possible that both dogs and cats have a strong negative influence on non-
domestic carnivores, through direct mortality, competition for food and space, and the
spread of disease and parasites (Hawkins & Racey 2008, Barcala 2009, Gerber et al.
2012a). An apparent decline in C. ferox abundance at Ankarafantsika National Park was
attributed to the growing population of stray dogs within the park (Barcala 2009). Although
C. ferox are known to predate lemurs, ground-dwelling species are an important part of
their diet, and all three introduced carnivore species are thus likely to compete with C. ferox
for food (Hawkins & Racey 2008, Barcala 2009). Future monitoring of the local
distribution of endemic and introduced carnivores is thus essential for conservation
planning and management.
Evans et al. (2013) recorded six captures of E. major in 227 trap nights at a mean
capture rate of 0.02 captures per night. Our study also recorded six captures, but with a far
greater sampling effort of 517 trap nights (mean capture rate 0.01 records per night). This
may indicate a decline in the abundance of E. major, especially as all the records obtained
by Evans et al. (2013) were obtained in Matsedroy forest. We did not record any E. major
in Matsedroy forest, despite our species accumulation curves suggesting that all species
present in the area had been recorded. No records of E. major were obtained in 2013 either,
albeit with a much lower sampling effort (i.e., 80 trap nights). Eupleres goudotii is thought
to be sensitive to habitat fragmentation (Gerber et al. 2012b), and it is possible that the
continued degradation of Matsedroy forest has reached a threshold at which they are no
longer able to persist in that forest patch. However, the low capture rate of E. major (six
captures in 517 trap nights) suggests that this result be treated with caution. Future
monitoring in Matsedroy forest, together with analytical tools such as occupancy
modelling, will be used to develop more robust measures of the local distribution of this
species.
While forest fragmentation has been identified as a major driver of extirpations of
Madagascar's endemic carnivores, there is evidence to suggest that intact carnivore
communities can persist in anthropogenically-modified forest areas (Gerber et al. 2012b).
We did not encounter any evidence of hunting of endemic carnivores, and it is likely that
these are protected by faddy (i.e., local taboo) as has been recorded elsewhere in
Madagascar (Jones et al. 2008). However, faddy does not confer universal protection; both

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53 Small Carnivore Conservation 52 & 53: 45–55
the perceived threat to humans and livestock and bushmeat hunting have led to the killing
of endemic carnivores elsewhere in Madagascar (Jones et al. 2008, Barcala 2009, Golden
2009). Consequently, the ongoing involvement of local communities in biodiversity
monitoring and conservation efforts in the Mariarano forest is thus key to preserving and
maintaining the forest’s carnivore biodiversity.
Acknowledgements
The authors would like to thank the Ministry of Forests and Environment for providing permits to
work in the Mariarano forest and DREF Boeny. The surveys were implemented and funded by
Operation Wallacea, Development and Biodiversity Conservation Action for Madagascar and the
Département de Biologie Animale (DBA), University of Antananarivo. We are grateful to GIZ
Boeny and VOI Tanteraka, and VOI Tanandava for accommodating our field camp at Mariarano
village and for providing field guides and easy access to the forest at Mariarano and Matsedroy
respectively. We would also like to thank the field guides at Mariarano, Matsedroy and
Antafiemeva, who assisted our field work, as well as the Operation Wallacea students, volunteers,
and support staff for their assistance. The authors would like to thank Dr. Frank Hawkins and one
anonymous reviewer for comments that led to the improvement of this manuscript.
References
Barcala O. 2009. Invasive stray and feral dogs limit fosa (Cryptoprocta ferox) populations
in Ankarafantsika National Park, Madagascar. Duke University (M.Sc. thesis),
Durham, North Carolina, U.S.A.
Chazdon R. L., Colwell R. K., Denslow J. S. & Guariga M. R. 1998. Statistical methods for
estimating species richness of woody regeneration in primary and secondary forests
of NE Costa Rica. Pp. 285–309 in Dallmeier F. & Comiskey J. A. (eds) Forest
biodiversity research, monitoring and modelling: conceptual background and Old
World case studies. Parthenon Publishing, Paris, France.
Choudhury A., Duckworth J. W., Timmins R., Chutipong W., Willcox D. H. A., Rahman
H., Ghimerey Y. & Mudappa D. 2015. Viverricula indica. The IUCN Red List of
Threatened Species 2015. <www.iucnredlist.org>. Downloaded on 5 December
2015.
Colwell R. K. 2006. EstimateS: Statistical estimation of species richness and shared
species from samples. www.purl.oclc.org/estimates
Dollar L. (Mustelid, Viverrid & Procyonid Specialist Group) 2000. Eupleres major. The
IUCN Red List of Threatened Species 2000. <www.iucnredlist.org>. Downloaded
on 5 December 2015.
Dollar L., Ganzhorn J. U. & Goodman S. M. 2007. Primates and other prey in the
seasonally variable diet of Cryptoprocta ferox in the dry deciduous forest of
Western Madagascar. Pp. 63–76 in Gursky, S. L. & Nekaris, K. A. I. (eds) Primate
anti-predator strategies. Springer, New York, U.S.A.

Cryptoprocta ferox in Mariarano forest, Madagascar
Small Carnivore Conservation 52 & 53: 45–55 54
Evans B., Rakotondraparany F., Cole L., Long P. & Gandola R. 2013. The carnivores of
Mariarano forest, Madagascar: first insights. Small Carnivore Conservation 49: 15–
19.
Garbutt N. 2007. Mammals of Madagascar – A Complete Guide. Yale University Press,
New Haven, U.S.A.
Gerber B. D., Karpanty S. M. & Randrianantenaina J. 2012a. Activity patterns of
carnivores in the rain forests of Madagascar: implications for species coexistence.
Journal of Mammalogy 93: 667–676.
Gerber B. D., Karpanty S. M. & Randrianantenaina J. 2012b. The impact of forest logging
and fragmentation on carnivore species composition, density and occupancy in
Madagascar’s rainforests. Oryx 46: 414–422.
Golden C. 2009. Bushmeat hunting and use in the Makira Forest, north-eastern
Madagascar: a conservation and livelihoods issue. Oryx 46: 386–392.
Goodman S. M. & Helgen K. M. 2010. Species limits and distribution of the Malagasy
carnivoran genus Eupleres (Family Eupleridae). Mammalia 74: 177–185.
Harper G. J., Steininger M. K., Tucker C. J., Juhn D. & Hawkins F. 2007. Fifty years of
deforestation and forest fragmentation in Madagascar. Environmental Conservation
34: 325–333.
Hawkins A. F. A. & Dollar L. 2008. Cryptoprocta ferox. The IUCN Red List of Threatened
Species 2008. <www.iucnredlist.org>. Downloaded on 6 October 2014.
Hawkins C. E. & Racey P. A. 2005. Low population density of a tropical forest carnivore,
Cryptoprocta ferox: implications for protected area management. Oryx 39: 35–43.
Hawkins C. E. & Racey P. A. 2008. Food habits of an endangered carnivore, Cryptoprocta
ferox, in the dry deciduous forests of Western Madagascar. Journal of Mammalogy
89: 64–74.
Ibouroi M. T., Schwitzer C. & Rabarivola J. C. 2013. Population density estimates of two
endangered nocturnal and sympatric lemur species from the Mariarano Forest,
northern Madagascar, using multiple approaches. Lemur News 17: 49–54.
Irwin M. T., Wright P. C., Birkinshaw C., Fisher B. L., Gardner C. J., Glos J., Goodman S.
M., Loiselle P., Rabeson P. & Raharison J.L. 2010. Patterns of species change in
anthropogenically disturbed forests of Madagascar. Biological Conservation 143:
2351–2362.
Jones J. P. G., Andriamarovololona M. M. & Hockley N. 2008. The importance of taboos
and social norms to conservation in Madagascar. Conservation Biology 22: 976-986
Kotschwar Logan M., Gerber B. D., Karpanty S. M., Justin S. & Rabenahy F. N. 2014.
Assessing carnivore distribution from local knowledge across a human-dominated
landscape in central-southeastern Madagascar. Animal Conservation 18: 82–91.
Kremen C., Cameron A, Moilanen A, Phillips S. J., Thomas C. D., Beentje H., Dransfield
J., Fisher B. L., Glaw F., Good T. C., Harper G. J., Hijmans R. J., Lees D. C., Louis
E., Nussbaum R. A, Raxworthy C. J., Razafimpahanana A, Schatz G. E., Vences
M., Vieites D. R., Wright P. C. & Zihra M. L. 2008. Aligning conservation

Mann et al.
55 Small Carnivore Conservation 52 & 53: 45–55
priorities across taxa in Madagascar with high-resolution planning tools. Science
320: 222–226.
Long P., Rabenandrasana M., Rabenoro M., Darlington A., McCann R., Gandola R.,
Morrison R., Graham S., Evans C., Washington H., Evans B., Palfrey R., Raveloson
B., Rakontondravony F., Jocque M. & Andriambelo H. 2012. Biodiversity surveys
of Mariarano and Matsedroy tropical dry forests and associated wetlands, Western
Madagascar; 2010–2012, status report. Unpublished report to the Ministry of
Forests and the Environment, Antananarivo, Madagascar.
Mittermeier R. A., Gil P. R., Hoffman M., Pilgrim J., Brooks T., Mittermeier C. G.,
Lamoreux J. & Da Fonseca G. A. B. 2005. Hotspots revisited. Earth’s biologically
richest and most endangered terrestrial ecoregions. Conservation International,
Sierra Madre, California, U.S.A.
Moat J. & Smith P. P. 2007. Atlas of the vegetation of Madagascar. Kew Publishing, Kew,
U.K.
Myers N., Mittermeier R. A., Mittermeier C. G., Da Fonseca G. A. & Kent J. 2000.
Biodiversity hotspots for conservation priorities. Nature 403: 853–858.
Thomas C. D. 1990. What do real population dynamics tell us about minimum population
sizes? Conservation Biology 4: 324–327.
Washington H., Rabenoro M., Andriambololona N., Griffin O., Higgins F., Osmond T.,
Shah P. & Clementine V. M. 2009. University of Bath Mahamavo Expedition 2009:
Final Report. Bath, U.K.
Woodroffe R. & Ginsberg J. R. 1998. Edge effects and the extinction of populations inside
protected areas. Science 280: 2126–2128.