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Lemur Distribution and Resident Attitudes Towards Forest Loss and Degradation in Ankarafantsika National Park, Madagascar


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Lemurs across Madagascar are threatened with extinction. This threat is largely associated with lemur habitat being converted for agriculture resulting in increased fragmentation of their forest. It is important, therefore, to record species presence across their geographic range in order to observe any local extinctions and at-risk populations, to provide accurate information on population size, and identify species that are most at risk of extinction in the wild. Ankarafantsika National Park is one of the last deciduous dry forests in Madagascar under state protection. Eight lemur species are known to occur in and around the park: Avahi occidentalis, Eulemur fulvus, Eulemur mongoz, Lepilemur edwardsi, Microcebus murinus, Microcebus ravelobensis, Cheiro-galeus medius, and Propithecus coquereli. Pressures on the forest of Ankarafantsika, such as logging and fire, have reduced forest cover and increased fragmentation. We conducted a rapid assessment of the occurrence of lemurs and their relative abundance around the periphery of the park, including sites that had not been surveyed previously. We also aimed to determine how the local communities perceived forest loss and the presence of these lemurs. We completed 10 diurnal and 19 nocturnal lemur surveys at nine sites and interviewed 11 residents in four villages close to the park boundary. We found seven of the eight species present within Ankarafantsika, and site-and species-level differences in encounter rates. Species richness differed across sites and may be negatively related to levels of anthropogenic disturbance at these sites. We found differences in the respondents' knowledge of the forest according to their sex, and residents reported declining numbers of lemurs, and increased fires and charcoal production that has negatively impacted the forest during their lifetimes. We highlight the need for long-term monitoring of lemur populations across the park and immediate conservation action to protect forests and lemurs.
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Primate Conservation 2020 (34): xx-xx
Lemur Distribution and Resident Attitudes Towards Forest Loss
and Degradation in Ankarafantsika National Park, Madagascar
Travis S. Steffens1,2, Hasina Mamy Rabe Maheritaka3, Jesse Hildebrand4 and Megan Aylward5
1Department of Sociology and Anthropology, University of Guelph, Guelph, Ontario, Canada
2Planet Madagascar, Guelph, Ontario, Canada
3Department of Zoology and Animal Biodiversity, University of Antananarivo, Ambohitsaina, Madagascar
4Exploring By The Seat of Your Pants, Elora, Ontario, Canada
5National Centre for Biological Sciences, TATA Institute of Fundamental Research, Bengaluru, Karnataka, India
Abstract: Lemurs across Madagascar are threatened with extinction. This threat is largely associated with lemur habitat being
converted for agriculture resulting in increased fragmentation of their forest. It is important, therefore, to record species presence
across their geographic range in order to observe any local extinctions and at-risk populations, to provide accurate information on
population size, and identify species that are most at risk of extinction in the wild. Ankarafantsika National Park is one of the last
deciduous dry forests in Madagascar under state protection. Eight lemur species are known to occur in and around the park: Avahi
occidentalis, Eulemur fulvus, Eulemur mongoz, Lepilemur edwardsi, Microcebus murinus, Microcebus ravelobensis, Cheiro-
galeus medius, and Propithecus coquereli. Pressures on the forest of Ankarafantsika, such as logging and re, have reduced forest
cover and increased fragmentation. We conducted a rapid assessment of the occurrence of lemurs and their relative abundance
around the periphery of the park, including sites that had not been surveyed previously. We also aimed to determine how the local
communities perceived forest loss and the presence of these lemurs. We completed 10 diurnal and 19 nocturnal lemur surveys at
nine sites and interviewed 11 residents in four villages close to the park boundary. We found seven of the eight species present
within Ankarafantsika, and site- and species-level differences in encounter rates. Species richness differed across sites and may
be negatively related to levels of anthropogenic disturbance at these sites. We found differences in the respondents’ knowledge of
the forest according to their sex, and residents reported declining numbers of lemurs, and increased res and charcoal production
that has negatively impacted the forest during their lifetimes. We highlight the need for long-term monitoring of lemur popula-
tions across the park and immediate conservation action to protect forests and lemurs.
Key words: Biogeography, lemur communities, conservation, deforestation, species richness, rapid assessment
The lemurs of Madagascar and the Comoros are the
most endangered animal group in the world (Schwitzer et
al. 2013); the most recent IUCN Red List assessment (July
2020) lists 105 (94.6%) of the 111 species and subspecies
as threatened (IUCN 2020). These primates are primarily
arboreal, and many species have very restricted distributions
(Goodman and Benstead 2005; Schwitzer et al. 2013). Even
within the species’ ranges, their populations are patchily dis-
tributed (Wright et al. 2008). Approximately 44% of forests
in Madagascar have been converted to non-forest since the
1950s (Vieilledent et al. 2018) and, given this forest loss and
fragmentation across Madagascar, it is imperative that we
monitor the occurrence and abundance of the different spe-
cies of lemur over time and across different spatial scales to
gain a better understanding of the threats and their impacts.
The effects of habitat loss and degradation on lemurs
are well studied (Schwitzer et al. 2013; Schüßler et al. 2018;
Kling et al. 2020). Habitat loss is the greatest threat (Schwit-
zer et al. 2013) but the impact of habitat degradation is more
complex. Some lemurs have shown positive responses to hab-
itat degradation. Avahi laniger and Microcebus cf. simmonsi,
for example, have been found at higher densities in regen-
erating secondary forest compared to mature forest (Miller
et al. 2018). Bamboo lemurs (Hapalemur meridionalis) use
Steffens et al
habitats dominated by invasive plants, which allow them
to tolerate habitat degradation and fragmentation (Eppley
et al. 2015). Schüßler et al. (2018), however, reported that
lemur species diversity was negatively impacted by habitat
Ankarafantsika National Park in northwest Madagascar
is primarily deciduous dry forest. It hosts eight species of
lemurs (Alonso et al. 2002; Razafy Fara 2003; Mittermeier
et al. 2010; Gautier et al. 2018). Research and associated
conservation efforts in the park have been primarily centred
around Ampijoroa, an area located close to the Madagascar
National Parks (MNP) headquarters along the national road
RN4 (e.g., Thorén et al. 2010; Joly 2011; Ramsay et al. 2017).
Information on the degree of disturbance to the forest, and the
effect of this disturbance on lemur species in other areas of
the park is, therefore, limited. Ankarafantsika has been high-
lighted as a priority area for lemur conservation by the IUCN
Species Survival Commission (Schwitzer et al. 2013). The
IUCN SSC action plan for lemur conservation (Schwitzer et
al. 2013) recommends increased rapid assessment surveys of
remote locations within the park (Radespiel and Razandra-
manana 2013). Despite its status as a national park, activities
such as logging, deforestation, and hunting threaten the lemur
populations there (García and Goodman 2003; Guschanski et
al. 2007; Radespiel and Raveloson, 2001), and while res are
typical components of the seasonal cycle of deciduous dry
forests, increased instances of uncontrolled res may have
detrimental effects on the ora and fauna and their ability to
recover from severe damage (Gautier et al. 2018).
Studies of lemurs in Ankarafantsika but outside of Ampi-
joroa have revealed that some species are widespread, mouse
lemurs, for example (Guschanski et al. 2007; Radespiel et
al. 2008; Rakotondravony and Radespiel 2009; Steffens and
Lehman 2018, 2019), whereas others may have been locally
extirpated, the woolly lemur Avahi occidentalis, for example
(Steffens and Lehman 2019). Species richness and occur-
rence in some areas of Ankarafantsika have been shown to
be positively related to the habitat available, especially for
species larger than Microcebus (Steffens and Lehman 2018,
2019). Propithecus coquereli numbers drop near forest edge
and roads (Kun-Rodrigues et al. 2014). In 1997, Schmid and
Rasoloarison (2002) carried out a rapid assessment at three
sites in the interior of the park that revealed the presence of
eight lemurs—ve nocturnal, two cathemeral, and one diur-
nal. Besides providing valuable information on lemur diver-
sity in the three locations, it revealed lower levels of distur-
bance in the east of the park, furthest from the national road
(Schmid and Rasoloarison 2002).
Community-based conservation initiatives aim for a
bottom-up approach to managing landscapes in contrast to
the top-down approach of so-called “fortress conservation”
(Adams and Hulme 2001). When Ankarafantsika was rst
established as a protected area in 1927, it was classed as a
“Réserve Naturelle Intégrale” (Virah-Sawmy et al. 2014). As
a result of the declared “Durban Vision”, announced at the
Vth World Parks Congress in Durban in 2003, it was elevated
to a national park, and there was, besides, consideration given
to including residents of the local communities into manage-
ment decisions, management activities, and as benefactors of
management (Aymoz 2013). For example, there is a buffer
zone on the edge of the park that is managed by local com-
munities, and 50% of the fee charged to visit the park is dedi-
cated to conservation/development projects aimed to benet
local people (Aymoz et al. 2013). The implementation of
community-based conservation projects has, however, suf-
fered some set-backs. A commercial-based community forest
management program near Ankarafantsika, for example,
was unsuccessful in protecting the forest (Rasolofoson et al.
2015). In the Ambohitantely Special Reserve in the central
highlands, Klein et al. (2007) found that the implementation
of community-based conservation strategies actually fol-
lowed a top-down approach, as they neglected to understand
the needs and interests of local communities; a situation that
resulted in raised tensions towards conservation activities.
They found that, although poverty was considered the greatest
threat to forest conservation, most damage to the forest was
caused by res set by relatively wealthy people, who own and
graze cattle near the protected area. Thus, it is important to
understand local dynamics, perceptions, and attitudes related
to conservation and management of protected areas in Mada-
gascar (Kaplin 2005). In Ankarafantsika, local residents
around the periphery of the park have the least positive per-
spectives on conservation actions, compared to those in the
interior (near Ampijoroa) or where the Madagascar National
Parks agents are employed (Aymoz et al. 2013). These areas
could, therefore, benet from integration of these communi-
ties into conservation initiatives, taking into account commu-
nity perspectives on effective ways to protect the habitat and
the lemur species present (Fritz-Vietta et al. 2011).
We walked the perimeter of Ankarafantsika National
Park, to (i) assess the distribution of lemur species across the
park and (ii) discuss changes to the forest that residents had
observed over their lifetimes, as well as their current attitudes
towards conservation. We used a rapid assessment survey
technique to obtain information on lemur species occurrence
and relative abundance around the national park, and inter-
viewed people who have lived within or near to the park over
the last few decades to see their perspective on changes to
the forest and lemur species. Dialogue with older residents
allowed us to gain qualitative information on rates of changes
and how these are viewed. These perspectives can help us to
identify priority areas for species monitoring and the com-
munities that are interested in future conservation initiatives.
Lemurs of Ankarafantsika National Park
Figure 1. Map of rapid assessment path, camp, and sighting locations in Ankarafantsika National Park (ANP), Madagascar. C# = locations for each camp near survey
Study area
Ankarafantsika National Park (135,800 ha) is located at
16°6'55.29”S, 47°05'49.10"E on the Route National 4 (RN4),
approximately 115 km southeast of Mahajunga in the Boeny
region, northwest Madagascar. The park has eight known
lemur species (Table 1) and comprises a mosaic of dry decidu-
ous forest and secondary grassland created by anthropogenic
activities (Gautier et al. 2018; Radespiel and Razandrama-
nana 2013). There are approximately 60 communities within
and near the park boundaries, with an estimated total popula-
tion of 100,000 (Madagascar National Parks 2017). We con-
ducted this survey over a period of 14 days from 28 June – 11
July 2019. We walked the perimeter of the park (Fig. 1) to
gain access to areas distant from Ampijoroa and the national
highway (RN4).
Lemur surveys
We selected nine villages around the periphery of the park
as bases, near to which we could set up temporary camps (Fig.
1), to facilitate access to trails, local assistants, and water. We
conducted lemur surveys in forest near to each of the nine
camps around the park using the line-transect method on
existing trails. These trails were all in the continuous forest
inside the park boundary with the exception of Camp 5 where
we surveyed a large fragment on the edge of the park bound-
ary (Fig. 1). We conducted a total of n = 29 (10 diurnal; 19
Steffens et al
nocturnal) surveys over 14 days and nights surveying a total
distance of 22.15 km. Each survey team consisted of 2–4
We collected the following data at the beginning and end
of each transect; Survey number, date, surveyors’ names, start
time, GPS coordinates at the start of the trail, heading of trail,
temperature, cloud cover, moon phase if applicable, end time,
GPS at end of trail, distance walked, total number of sight-
ings, number of species. We walked each transect at a speed
of 0.75 and 1 km/hr for nocturnal and diurnal surveys, respec-
tively. When an animal was seen (visual sighting) or heard
(auditory sighting), we recorded the following: date, time,
side of transect, species, distance along transect (m), observer
GPS, trail heading, lemur heading, distance to the center of
the group, perpendicular distance along transect, perpendicu-
lar GPS, perpendicular distance to centre of the group, group
spread, height of the animal, number of individuals, behav-
iour, and method of detection (visual or auditory). A visual
or auditory record was considered one sighting regardless of
the number of individuals observed (i.e. groups were treated
as a single sighting).
We calculated the relative abundance of each species (by
camp and by species overall) as the encounter rate (number of
sightings of each species per km walked). We dene species
richness as the total number of species observed near each
camp. Unfortunately, we could not discriminate between the
two, mouse lemur species, found in the park, M. raveloben-
sis and M. murinus. Although there are differences in occur-
rence and abundance for each species at small scales, both
species were considered to occur throughout the park (Jolly et
al. 2011; Andriaholinirina et al. 2020). We, therefore, consid-
ered occurrence of Microcebus spp. to represent both.
Qualitative assessments of forest disturbance and resource
In addition to walking survey transects, at three commu-
nities (Andranomidtra, Beronono, and Sainte Marie), we con-
ducted reconnaissance walks across larger areas (2–10 km).
On transects and these reconnaissance walks we observed
the integrity of the forest, signs of resource use, and wildlife
in the area. We did not systematically quantify the amount
of disturbance we observed but qualitatively noted the main
anthropogenic activities near each camp such as: existence
and type of trails, existence and the relative amount of the
following: charcoal production, re disturbance, extraction of
tubers (Dioscorea maciba), traps for hunting, and cut trees.
Based on the above qualitative assessment, we ranked each
camp as having a high or low level of disturbance, and if high
indicated what was the driver (for example, re, charcoal pro-
duction, or both). To achieve a high ranking the forest around
the camp had to have extensive forest loss or disturbance,
while a low ranking had little forest loss or disturbance. We
then compared the qualitative level of anthropogenic distur-
bance to lemur species richness observed at each camp.
Interviews with local residents
We conducted interviews in four communities (see
Supplementary material for Interview questions and replies,
Table S1: Andranomidtra, Beronono, Sainte Marie, and
Ampombolira) using an open-ended questionnaire. In each
community we asked the community leader to facilitate inter-
views with volunteers among elder members of each commu-
nity. Prior to being interviewed, we read an ethics statement
to each participant about the use of the information being col-
lected and asked whether they consented to being interviewed.
Interviews typically lasted 20–30 minutes and were recorded
using an Olympus Vn-8100PC digital voice recorder.
We completed our walk around the park by primarily fol-
lowing a rebreak/vehicle track around the perimeter. We
covered 220 km. We completed a total of 29 lemur surveys
and three opportunistic walks (Table 2). We interviewed
11 members of the four communities (Andranomidtra, Ber-
onono, Sainte Marie, and Ampombolira) including seven men
and four women (Table S1).
Lemur surveys
We recorded the presence of the following lemurs in the
Ankarafantsika National Park: Microcebus spp. (probably
two species), Lepilemur edwardsi, Avahi occidentalis, Eul-
emur fulvus, Eulemur mongoz, and Propithecus coquereli
(Fig. 1). We did not observe Cheirogaleus medius, which
hibernates during the dry season (Dausmann et al. 2005).
Species richness was highest at camps 4 (Beronono; n = 7)
and 7 (Ambarindahy; n = 6; Table 3). Relative abundance
was highest for Microcebus and lowest for Eulemur mongoz.
Our one sighting of E. mongoz was opportunistic, during a
reconnaissance walk and not on a survey (Table 4). The rela-
tive abundance of all species by camp ranged from 0.0–3.28
individuals/km for diurnal walks and 6.27–22.00 individuals/
km for nocturnal walks.
Community member responses about their daily tasks
differed by sex. The women reported that their daily tasks
typically included cooking meals and monitoring rice elds;
they didn’t know about wildlife because they rarely went into
the forest. The men reported that their daily tasks included
monitoring rice elds and occasionally going into the forest.
Six of the 11 men interviewed said that they had perceived a
decline in the wildlife over time (Table S1).
There were some regional differences in the nature of
deforestation reported by respondents around Ankarafan-
tsika. At Camp 1 (Andranomidtra), three of four respondents
reported that deforestation was caused by immigrants from
the “south,” who cut the trees to produce charcoal (Table
S1). Respondents at Camp 4 (Beronono) reported that the
Lemurs of Ankarafantsika National Park
park limits were moved from nearby the village to farther
away, and the community members used the forest between
the village and the new park limit for their daily needs (Table
S1). At Camp 5 (Saint Marie) a respondent suggested that
re was the main factor reducing the size and quality of the
forest around the community (Table S1). Both respondents
at Camp 8 (Ampombolira) also suggested re generated from
outside the community was the main cause of forest loss near
the community.
The majority of respondents found value in the forest
but said that the forests near their communities had dimin-
ished in recent years (Table S1). Most respondents claimed
that forest is either farther from their community (8/11) or
degraded (3/11). They reported a reduction in forest from
increased res, local residents clearing land, and immigrants
clearing forest for charcoal. Two respondents noted that the
remaining forest is now warmer than when there was more
forest. Eight of the respondents commented that there is less
wildlife including lemurs following the reduction in forest
near their community. Most respondents felt that the forest
is useful. Respondents reported that the forest provided
trees for human use (10/11), for example for building and
rewood; water (7/11) and clean air (3/11); medicinal plants
(2/11); homes for wildlife (2/11), and shade (1/11). Nine of
the eleven respondents felt that the loss of forest has affected
their community negatively. One felt that there was a posi-
tive benet from forest loss (rewood was easier to collect),
and another claimed there was no impact. Three respondents
reported that their community did not participate in conserva-
tion activities, while seven reported that their communities
had helped create and maintain rebreaks, and one told us
that their community had planted trees. Only one respondent
(C5 Sainte Marie) knew of external organizations helping to
conserve forest. Nine respondents commented that Mada-
gascar National Parks engaged in conservation activities, and
six reported that local communities did the same (Table S1).
Nine respondents wanted forest restoration activities near
their communities, one respondent did not want change, and
another did not respond to this question.
Forest disturbance and resource use
Although we did not collect systematic data on forest dis-
turbance or resource use over the course of the expedition, we
made note of anthropogenic activities. We observed that C1
(Andranomidtra) was characterized by extensive forest loss
and we saw evidence of charcoal production. There were
large and clear existing trails and evidence of the forest being
used by zebu cattle (Bos taurus indicus) near to these transects.
Steffens et al
At C2 (Bevazaha) we observed low levels of forest distur-
bance, and only little evidence of charcoal production. At
C3 (Ambahimalandy) we encountered one trap set for bush-
pigs (Potamochoerus larvatus). C4 (Beronono) is a large vil-
lage along the banks of the Mahajamba River. The forest
itself was relatively intact with little evidence of logging or
charcoal production. However, we did observe evidence of
collection of maciba (Dioscorea maciba; small tubers similar
to manioc) and one area that had been affected by re. C5
(Sainte Marie) was situated in a fragment on the edge of the
park. The fragment contained a small hamlet and the park
base station. Compared to the other camps, C5 appeared to
have more evidence of maciba extraction, re, and trails. C6
(Mahatazana) was characterized by heavy re damage which
penetrated into the forest. C7 (Ambarindahy) was situated in
a river valley and had moderate levels of disturbance includ-
ing trails, cut trees, evidence of maciba extraction, and re.
C8 (Ampombolira) was characterized as having numerous
trails, evidence of re, and some cut trees. At C9 (Bealana)
we camped at the Madagascar National Parks’ base station
and the forest was approximately 1.5 km away. Here the for-
ests have been cleared and there was substantial charcoal pro-
duction. In summary, C1 and C9 shared similar charcoal pro-
duction issues, C2, C3, and C8 had moderate levels of impact
from various sources. C4 and C7 were the least disturbed and
C5 and C6 were the most impacted by re.
Lemur species richness and forest disturbance
We found seven species near C4 and six near C7—both
communities have relatively low levels of anthropogenic dis-
turbance. In the areas with relatively high levels of charcoal
production such as C1 and C9, areas that see more frequent
and large res such as C5 and C6, or had a combination of
anthropogenic disturbance (C2, C3, and C8) fewer species
were present (2–4 species).
Lemurs of Ankarafantsika National Park
Through our rapid assessment we assessed the occur-
rence and relative abundance of lemurs along the periphery
of Ankarafantsika National Park. We assessed areas near to
the forest edge which are accessible to the communities and
are exposed to daily anthropogenic activities. Although many
studies have assessed the occurrence of lemurs in the park
(Radespiel and Raveloson 2001; Schmid and Rasoloarison
2002; Steffens and Lehman 2019), to date, no studies have
looked at species richness across the entire perimeter of the
park. During this study we observed seven of the eight spe-
cies that have previously been recorded in Ankarafantsika (we
did not see C. medius). There were, however, differences in
the occurrence rate and relative abundance of lemurs between
survey locations. Observed differences in species richness
and relative abundance among survey sites may be because
of variation in disturbance regimes and their intensity. Lemur
species richness was not uniform around the perimeter of the
park. We qualitatively assessed species richness in relation to
disturbance and found that it was lower in areas with higher
anthropogenic disturbance. Although our sample sizes are
small, our results suggest that human disturbance such as
charcoal production, and large frequent res impacts lemur
species richness.
There are differences in species richness and relative
abundance between our study and other studies. C4 (Ber-
onono) was the only survey site where we observed all seven
species at one location. This differs from Schmid and Raso-
loarison (2002) who recorded seven species (including C.
medius) at Ankarokaroka and six species each at Antsikoky
and Tsimaloto (sites located in the forest interior). While Tsi-
maloto is in the same area as our survey site C2 at Bevazah, it
is approximately 3 km farther into the interior than our survey
transects where we only observed four species (Microcebus
spp., E. fulvus and L. edwardsi). Near C2 (Bevazaha) and C5
(Sainte Marie) we recorded four species of lemur in each site,
whereas Radespiel and Raveloson (2001) recorded four near
Bevazaha and six near Sainte Marie. Two of our survey sites
(C4 and C9) were previously surveyed by Kun-Rodrigues et
al. (2014), during their survey of P. coquereli at four sites
across the park. Both studies found P. coquereli at each
site. We, however, observed only one group of P. coquereli
opportunistically at C4 (Beronono) and one group during a
nocturnal survey at C9 (Bealana). With greater survey effort
Kun-Rodrigues et al. (2014) reported nding 42 groups
at Beronono and 23 groups at Bealana. However, it is dif-
cult to compare the abundance of lemurs between studies
and study sites due to multiple factors that are not consistent
among studies, such as survey effort and methods. We did
not have the same level of survey effort in each site as other
studies had as our main aim was to survey the perimeter of the
entire park rather than carry out intensive local surveys.
Reports from local residents highlight that increased
forest loss and disturbance is occurring within and near the
park. The interviewees also made connections between a lack
of forest and a reduction in wildlife near their communities.
Reasons why forest was removed near their communities dif-
fered across the park, however. At C1, for example, residents
reported that forest loss along the southern end of the park is
the result of “southern” Tandroy immigrants removing trees
for charcoal production. The Tandroy are a nomadic ethnic
group originally from the Androy region of southern Mada-
gascar, who rely less on water-based agriculture and more
on other crops such as maize and raising zebu cattle (Waeber
et al. 2015). They collectively pioneer new land to extract
charcoal then divide the transformed land into smaller parcels
to be used by the Tandroy immigrants to grow maize to buy
zebu (Muttenzer 2012; Waeber et al. 2015). Waeber et al.
(2015) suggest that the land use practices of Tandroy explains
the decrease in forest cover in Androy and now the southern
side of the park. We noticed evidence of charcoal production,
including trees felled and stacked ready to burn, bags of char-
coal, and previously used charcoal repits, just outside the park
at C1 and within the park boundaries at C9, including areas
where we saw lemurs. At C8 and C9, residents report that
re is the main cause of forest loss around their communities.
Fire is a natural process typical to dry forest environments but
is also used by local cattle owners to promote new grasses for
their cattle to graze (Bloesch et al. 1999; Gautier et al. 2018).
However, there has been an increase in res, coupled with
the increased use of burning land for cattle, that has resulted
in less forest and more secondary grassland within and along
the periphery of the park (Gautier et al. 2018). Steffens and
Lehman (2018, 2019) reported that repeated use of re and
cattle grazing has created a fragmented landscape that has
negatively impacted lemur species richness and occurrence.
Finally, residents in C4 reported that deforestation near their
community was conducted by community members to plant
rice elds to grow food for the community. Converting forest
for agriculture is common practice in Madagascar (Harper
et al. 2007). The practice of cutting down forest has been
integral to many communities to grow food (Scales 2014).
Although this has complex effects on the environment (Scales
2014), some of the unintended consequences of removing
forest result in decreased biodiversity (Schwitzer et al. 2013),
increased erosion, and localized changes in rain and water
tables (Zwartendijk et al. 2017).
Our rapid assessment of lemurs and interviews of local
residents around Ankarafantsika suggest that the forest and
lemurs in and around park continue to be under threat. The
main threats include charcoal production and large res. We
found few lemur species per site and had lower encounter
rates for all species within the park relative to previous rapid
assessments, with the exception of Microcebus, which are
relatively abundant and ubiquitous. We cannot speak for C.
medius, which was hibernating during our study. Anthro-
pogenic disturbance varies across the park. We suggest that
urgent action is needed to alleviate the impact of charcoal pro-
duction in the south and res in the north.
Providing solutions for cattle owners may include local
and federal strategies that allow for controlled res to be
Steffens et al
set (Klein et al. 2007) that allow for this income-generating
activity while minimizing the impacts on the ground. Simi-
lar to Ambohitantely Special Reserve, re is not tolerated
in some parts of the park, and cattle owners there have no
incentive to stop runaway wildres from burning the forest.
Strategies that allow cattle owners to use res in a way that
is also benecial to future forest growth (i.e. reduce fuel load
to prevent even larger res) could be used in areas where re
poses a greater risk to forest. Patrols and physical re-breaks
have been found to provide some level of protection to forest
within the park (T. S. Steffens unpubl. data). The few sight-
ings of larger species, including the Critically Endangered P.
coquereli and E. mongoz, warrants continued investigation to
determine the parkwide distribution and population structure
for these species. Species-specic action plans should be
drawn up for these species in order to reverse what appears to
be a declining trend in their geographic range and population
Our study is limited by two main factors. The rst, is
the wide area covered and the limited time of our study that
resulted in low survey effort per site. As such, it is difcult to
compare our encounter rates to studies using different meth-
ods or with greater survey effort. The second was that we
were not able to systematically assess the habitat structure
and human disturbance at each site. Our qualitative data
limited our ability to determine how human disturbance is
impacting lemur species richness, occurrence, and abundance.
It is the rst, however, of a potentially longer-term study
aimed to survey lemurs around the whole of Ankarafantsika
National Park, and is a rst step in meeting the proposal set
out by Radespiel and Razandramanana (2013) to (i) conduct
continuous and long-term assessments of lemurs near 12 base
camps situated around the park, and (ii) carry out additional
rapid assessments to understand the changes to distribution
and population structure of lemurs within it. We have pro-
vided baseline data at these sites and we have engaged with
the communities and CLP at sites around the park, which can
be used for future surveys and expanded monitoring efforts.
To be successful, conservation efforts must engage commu-
nity members directly in order to address direct and indirect
causes of this conservation crisis and include development
activities that help offset the high levels of poverty, consid-
ering the historical context and the cultural perspectives of
community members engaged in activities that puts pressure
on forests and lemurs.
We thank Jean Paul Soa, and the Comité Locaux des
Parcs representatives, who assisted us in the research; Mamy
Razatsalama and Georges Marcellin, who provided logisti-
cal support; the Ankarafantsika National Park Director and
Madagascar National Parks staff, who helped facilitate the
project; the Ministère de l'Environnement et du Développe-
ment Durable; and all local community members who hosted
us and supported our project. We are grateful to the following
funding agencies: Primate Conservation Inc. (PCI#152), and
IUCN SOS through Global Wildlife Conservation’s Lemur
Conservation Action Fund.
Ethical standards: All research, including our lemur sur-
veys and questionnaires in this study abided by the national
laws of Madagascar and was authorized and received ethical
clearance by the Ministère de l'Environnement et du Dével-
oppement Durable via permit No. 147/19/MEDD/SG/DGEF/
DGRNE issued to Travis Steffens.
Supplementary material: Interview questions and Table SI
- interview responses. Go to <
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Authors’ addresses
Travis S. Steffens, Department of Sociology and Anthro-
pology, University of Guelph, 50 Stone Rd E., Guelph, ON,
Canada, N1G 2W1; Hasina Mamy Rabe Maheritaka,
Mention of Zoology and Animal Biodiversity, University of
Antananarivo, BP Ambohitsaina, Madagascar; Jesse Hildeb-
rand, Exploring By The Seat of Your Pants, 45 Clegg Road,
Elora, ON, Canada, N0B 1S0; Megan Aylward, National
Centre for Biological Sciences, TATA Institute of Fundamen-
tal Research, GKVK Road, Bellary Road, Bengaluru, Kar-
nataka, 560065. Corresponding author: Travis S. Steffens,
e-mail <>.
Received for publication: 18 March 2020
Revised: 10 July 2020
... For instance, 82.67% of Malagasy professionals but just 19.28% of non-professionals selected logging as a top threat (p = 0.00); 78.67% of professionals but just 33.73% of non-professionals selected hunting (p = 0.00); 52.00% of professionals but just 16.87% of non-professionals selected charcoal production (p = 0.00); and 40.00% of professionals but just 9.64% of non-professionals selected the pet trade (p = 0.00). As logging (Rakotomanana et al. 2013), hunting (Borgerson 2015), charcoal production (Steffens et al. 2020), and the pet trade (LaFleur et al. 2017) are conservation threats for lemurs, education programs about these topics may help gain support among Malagasy non-professionals for solutions that address these threats. ...
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The dry forests in Madagascar represent a remarkable tropical forest ecosystem, occupying almost the entire west slope of the island up to the very northern tip, especially on substrates associated with sedimentary formations. These forests span several woody vegetation types of the island, including (i) the southwestern coastal bushland, (ii) the southwestern dry spiny forest-thicket, and (iii) the western dry forest. These landscapes show a high degree of biodiversity with several centers of endemism hosting a globally unique fauna, with disparities in richness and diversity according to the groups, probably related to paleo-refugia. These landscapes also provide important ecosystem services for various ethnic groups residing along the coast, also hosting the only autochthonous group in Madagascar, the Mikea forest people. In this paper we review the scientific literature to highlight the importance of dry forests socio-ecological landscapes in order to identify knowledge gaps where future research is required to better inform management and policy to better balance conservation and development interests. For this, we recommend the adoption of transdisciplinary approaches that engage with a broad number of stakeholders in order to allow policy adaptations to better cope with current and future changes (e.g., agriculture, energy demands and needs). Les forêts sèches de Madagascar représentent un remarquable écosystème tropical forestier et sont distribuées sur la quasi-totalité du versant occidental jusqu'à l'extrême nord de l'île, et plus particulièrement sur les terrains associés à des formations sédimentaires. Ces forêts compren-nent plusieurs types de végétation sylvicole dont: (i) la formation buissonnante côtière du sud-ouest, (ii) la forêt sèche épineuse du sud-ouest, et (iii) la forêt sèche de l'ouest. Ces paysages abritent une importante biodiversité avec plusieurs centres d'endémisme qui hébergent des faunes uniques à l'échelle globale, avec des particularités en matière de richesse et de diversité en fonction des groupes, probablement liées aux paléo-refuges. Ces paysages fournissent également d'importants services écosystémiques pour les divers groupes ethniques résidant sur la bande côtière occidentale, notamment aux Mikea, qui sont le seul groupe autochtone reconnu pour Madagascar. Dans cet article, nous passons en revue la littérature scientifique pour souligner l'importance des paysages socio-écologiques des forêts sèches, afin d'identifier les lacunes de connaissance à combler pour mieux informer une gestion et une politique capables d'harmoniser les besoins de la conservation et du développe-ment. Dans ce cadre, nous préconisons la nécessité des approches transdisciplinaires portées par un grand nombre de parties prenantes pour permettre les adaptations des politiques face aux changements actuels et futurs, comme, par exemple, pour l'agriculture et les besoins et les demandes en énergie. Los bosques secos de Madagascar representan un ecosistema de bosque tropical notable, que ocupa casi la totalidad de la ladera oeste de la isla hasta la punta norte, especialmente en sustratos asociados con formaciones sedimentarias. Estos bosques abarcan varios tipos de vegetación
The rise in research investigating fragmentation and its impact on primates and other taxa reflects the growing presence of fragmented landscapes themselves. Although numerous studies report the negative effects of fragmentation on biodiversity, it is difficult to generalize responses to fragmentation for specific taxonomic groups, such as non-human primates, when studies have not employed a definitive concept of fragmentation or fragments themselves. Madagascar's high degree of fragmentation, wealth of endemic taxa, and extensive history of ecological research provide the opportunity to compare fragmentation studies across similar contexts. We conducted a literature search of peer-reviewed articles on fragmentation in Madagascar to characterize its trends. A total of 70 articles, 46 of which concentrated on lemurs, tested the impacts of fragmentation on Malagasy taxa, while additional sources conducted research in one or more fragments without testing its effects (n = 112 total, 79 on lemurs). Studies on lemurs most frequently tested fragmentation's impacts on genetics and biodiversity metrics (n = 16 and 15 studies, respectively), although health, modeling, behavioral, and cross-disciplinary techniques were also reported. Responses to fragmentation were reported for 49 lemur species, with most studies concentrated in eastern Madagascar (87%). Although there was variation in the metrics reported in studies testing the effects of fragmentation on Malagasy species, the most common measures were fragment area, isolation, or comparison to a control site. Landscape-scale approaches and examination of fragmentation per se were rarely employed. Characterizing trends of fragmentation research in Madagascar emphasizes the challenges of documenting fragmentation's effects while highlighting the benefits of research within fragmented landscapes, particularly when combined with consideration for how the matrix within human-modified landscapes may impact primate populations.
We used species‐area relationships (SARs) to investigate the effects of habitat loss on lemur biogeography. We measured species richness via visual surveys on line transects within 42 fragments of dry deciduous forest at the Ambanjabe field site in Ankarafantsika National Park, Madagascar. We measured human disturbance and habitat characteristics within 38 of the 42 fragments. We measured the distance between each fragment and the nearest settlement, continuous forest, and nearest neighboring fragment. We fit 10 candidate SAR models to the data using nonlinear least squares regression and compared them using Akaike's Information Criterion (AIC). To determine how habitat characteristics, as well as area, influenced species richness, we ran a hierarchical partitioning procedure to select which variables to include in generalized additive models (GAMs) and compared them using AIC. Contrary to expectations, we found that lemurs form convex SARs, without a “small island effect”, and with the power model being the most likely SAR model. Although we found that four variables (area, survey effort, and total human disturbance, and mean tree height) independently contributed greater than 10% of the variation in lemur species richness, only area was included in the most likely model. We suggest that the power model was the most likely SAR model and our inability to detect a “small island effect” are the result of Microcebus spp. being edge tolerant and capable of dispersing through matrix, scale issues in the study design, and low γ‐diversity in the landscape. However, more study is needed to determine what role human disturbance plays in influencing species richness in lemurs. RESEARCH HIGHLIGHTS • 1. Understanding the pattern of the species‐area relationship (SAR) of lemurs is important as suitable forests are shrinking • 2. Lemurs form a convex rather than sigmoidal shaped SAR in a fragmented landscape • 3. Lemurs can occupy fragments <1 ha
A majority of Madagascar's iconic lemurs (Primates, Strepsirrhini) is threatened with extinction due to anthro-pogenic activities like land use change (deforestation) and bushmeat hunting. We used a multivariate approach combining land cover mapping, vegetation/degradation monitoring, the degree of anthropogenic disturbance and the status of forest protection by the local community to model their impact on lemur diversity, population densities and encounter rates within a rural area of lowland rain forest in northeastern Madagascar. High mean annual deforestation rates (2.4%) were calculated since 1990, resulting in a landscape of small and isolated forest fragments. A limited number of eight lemur species belonging to five lemur families were encountered. Diurnal species were absent, while cathemeral lemurs avoided human disturbance. Small and nocturnal species were relatively abundant. Overall lemur diversity was best explained by forest size and a combination of disturbance and hunting. Encounter rates of three nocturnal taxa were influenced by forest size and habitat degradation. Community-level forest protection had no effect on lemur diversity, but coincided with lower levels of habitat degradation. Lemur population sizes were relatively small and only few forests remain that offer suitable habitats for viable populations. We highly recommend external conservation NGOs to support local forest management by improving the existing community-based approach. Actions should include expansion of protected habitats to increase population connectivity (reforestation) and to decrease lemur disturbance by villagers. Without external support, the last remaining forest habitats will be devastated within a few years resulting in the local extinction of most lemur populations.