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Rapid Decrease in Populations of Wild Ring-Tailed Lemurs (Lemur catta) in Madagascar

  • Franklin Scholars

Abstract and Figures

Lemurs are the most threatened group of mammals on earth. Lemur catta (ring-tailed lemur) represents one of the most iconic lemur species and faces numerous an-thropogenic threats in the wild. In this study, we present population estimates from 32 sites across the range of L. catta , collected from primary and secondary data sources, to assess the number of ring-tailed lemurs left in the wild. We estimate that there are approximately 2,220 individual L. catta remaining in the 32 sites considered. We note local extinctions of populations of L. catta in at least 12 of the 32 sites examined, and that significantly more extinctions occurred in areas without some form of protection. This decrease in extant populations could represent a decrease of more than 95% of all ring-tailed lemurs in Madagascar since the year 2000. While these results should be considered preliminary, we stress the rapid decline of the species and note that habitat loss, bush-meat hunting and the illegal pet trade are driving populations to local extinction. Based on the data presented here, urgent and immediate funding and conservation action are crucial to ensure the viability of the remaining wild populations of ring-tailed lemurs.
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Short Communication
Folia Primatol 2016;87:320–330
DOI: 10.1159/000455121
Rapid Decrease in Populations of
Wild Ring-Tailed Lemurs ( Lemur catta ) i n
Marni LaFleur a Tara A. Clarke b Kim Reuter c Toby Schaeffer d
a Department of Anthropology, University of California San Diego, La Jolla, CA , and
b Evolutionary Anthropology Department, Duke University, Durham, NC , USA;
c Africa Field Division, Conservation International, and d Independent Consultant
(pro bono), Nairobi , Kenya
Ring-tailed lemur · Madagascar · Extinction · Bushmeat · Pet trade ·
Population decline · Lemur catta
Lemurs are the most threatened group of mammals on earth. Lemur catta (ring-
tailed lemur) represents one of the most iconic lemur species and faces numerous an-
thropogenic threats in the wild. In this study, we present population estimates from 32
sites across the range of L. catta , collected from primary and secondary data sources, to
assess the number of ring-tailed lemurs left in the wild. We estimate that there are ap-
proximately 2,220 individual L. catta remaining in the 32 sites considered. We note local
extinctions of populations of L. catta in at least 12 of the 32 sites examined, and that sig-
nificantly more extinctions occurred in areas without some form of protection. This de-
crease in extant populations could represent a decrease of more than 95% of all ring-
tailed lemurs in Madagascar since the year 2000. While these results should be considered
preliminary, we stress the rapid decline of the species and note that habitat loss, bush-
meat hunting and the illegal pet trade are driving populations to local extinction. Based
on the data presented here, urgent and immediate funding and conservation action are
crucial to ensure the viability of the remaining wild populations of ring-tailed lemurs.
© 2017 S. Karger AG, Basel
Lemurs are the most threatened group of mammals on earth and are endemic to
Madagascar, a biodiversity hotspot [Schwitzer et al., 2014]. Lemurs face numerous
threats, including habitat loss due to deforestation and climate change [Brown and
Received: August 29, 2016
Accepted after revision: December 14, 2016
Published online: January 13, 2017
Marni LaFleur
Department of Anthropology
University of California San Diego
Campus Box 0532, La Jolla, CA 92093 (USA)
E-Mail marni.lafleur
© 2017 S. Karger AG, Basel
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DOI: 10.1159/000455121
Yoder 2015], hunting [Schwitzer et al., 2014], and the illegal pet trade [Reuter et al.,
2016]. Lemurs have been described as “the single highest primate conservation prior-
ity in the world” [Schwitzer et al., 2013, p. 5].
Ring-tailed lemurs ( Lemur catta ) are a flagship species for conservation, and are,
perhaps, Madagascar’s best-known primate. These lemurs are characterized by their
behavioural flexibility, in that their sociality (e.g., dominance/rank relationships, ag-
gression) and activity budgets vary according to environmental conditions [Sauther
et al., 1999], and ecological plasticity, in that they can inhabit a plethora of habitat
types [Goodman et al., 2006] and rebound from environmental perturbations [Gould
et al., 1999]. The first large-scale population estimates of wild ring-tailed lemur hab-
itats and individuals estimated a 9.5% habitat and 20% population decrease between
1985 and 2000 (population estimated at 751,251 in the year 2000 [Sussman et al.,
2006]). Since then, census attempts of specific sites have continued [e.g., Kelley, 2013;
Dimilahy et al., 2015; Gould and Andrianomena, 2015], yielding evidence of localized
ring-tailed lemur extinctions [Gardner and Davies, 2014] and a suspected population
decline rate of over 50% over a 3-generation period [Andriaholinirina et al., 2014].
P o p u l a t i o n d e c l i n e s o f L. catta are driven by various threats. Ring-tailed lemurs
represent 28% of individuals kept as in-country illegal pets [Reuter and Schaefer, 2016],
and are sometimes killed and consumed by their human owners [Reuter and Schaefer,
in press]. This species also faces direct hunting pressure [e.g., Goodman 2003], signifi-
cant habitat loss [Brinkmann et al., 2014], and population declines resulting from the
effects of climate change [Brown and Yoder, 2015]. As a result, L. catta is now restricted
to isolated fragments [Sussman et al., 2003] with relatively low population densities
[Kelley 2013; Andriaholinirina et al., 2014; Gould and Andrianomena, 2015], and its
survival in the wild at most locations remains perilous [but see Dimilahy et al., 2015].
In this study, we present a population estimate for ring-tailed lemurs across 32
locations within their natural and historical range across south-western Madagascar.
Moreover, we discuss the consequences of our findings and propose next steps to en-
sure the long-term viability of remaining wild populations.
Estimating Population Numbers at Known Sites
We assessed the status of populations of L. catta at 32 locations where they are known to
occur ( Fig.1 ; Table1 ). Similar to a census of chimpanzees ( Pan troglodytes ) by Plumptre and Cox
[2006], we collected primary and secondary data regarding the largest known and/or protected
habitat fragments in which L. catta is found to estimate the total number of individuals left in the
wild. L. catta is found at locations outside of these 32 known sites [Goodman et al., 2006, map
approx. 100 records of wild populations of L. catta ); however, these 32 sites likely comprise the
largest remaining populations of L. catta , as many of the previously mentioned “known” sites are
now deforested or devoid of animals [Sussman et al., 2006; Gardner and Davies, 2014].
To assess the status of populations of L. catta at these 32 sites, we used a mixture of methods:
in-person site visits (M.L. and T.A.C.), personal communication with other researchers, and in-
formation gleaned from the published literature ( Table1 ). We could not visit all 32 field sites due
to time and budget limitations; for in-person site surveys, we prioritized locations where previous
researchers suggested that populations of L. catta were under significant extraction pressure [e.g.,
Kirindy Mitea Sud hunting pressure, Goodman, 2003; Andohahela hunting pressure, Siers, 2007].
For in-person site surveys ( n = 10 sites, Table1 ), M.L. and T.A.C. spent a minimum of 2
days at each site in June-August 2016 and used rapid assessment methods [Gilles and Reuter,
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DOI: 10.1159/000455121
322 LaFleur/Clarke/Reuter/Schaeffer
2014] to confirm or fail to confirm the presence of L. catta . Since we aimed to confirm the pres-
ence of ring-tailed lemurs in the locations visited, we took steps to increase the chances of detect-
ing animals. These included speaking to local officials and villagers in order to understand where
animals had been seen in the recent past, or where ring-tailed lemur troops’ sleep sites or fre-
quently used trees may be located. Once target areas were identified, we then walked on both
forest trails and off-trail locations for between 5 and 12 km/day, while trying to detect the pres-
ence of ring-tailed lemurs. Confirmation of the presence of L. catta included hearing calls or ob-
serving animal(s), or indirect indicators of presence such as seeing footprints, spur markings on
vegetation, bite marks on plants/foods, and/or presence of faeces. Indirect indicators of animal
presence are reliable when efforts are standardized and consistent [Macdonald et al., 1999], and
can be used to detect population changes in wildlife [Gese, 2001]. At all sites where the presence
of L. catta could not be confirmed, M.L. and T.A.C. spoke with park managers, tour guides, and/
or local community members to understand when and how many L. catta had last been seen at
that site, and whether the animals had faced any known threats in the past. At 7 out of 10 sites,
M.L. and T.A.C. were unable to confirm the presence of L. catta and therefore consider these
populations to be locally extinct unless otherwise noted ( Table1 ). At the remaining 3 sites (Isalo
Historical range (a)
Records of Lemur catt a
Protected areas
Historical range
Current range (b)
Locally or nearly locally extinct
Protected areas
Fig. 1. Historical range ( a ) and estimated current range ( b ) for L. catta , including areas where
populations are present, locally extinct, or likely or nearly locally extinct. a Data are from Good-
man et al. [2006] and Andriaholinirina et al. [2014]. b References are available from Table1.
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Table 1. Locations of known populations of L. catta included in the scope of this paper
Location name Coordinates Protected status Population category Number
Size of forest
fragment or
areaa, km2
Year of
Source of
Days of
Citation (most recent
25°04.70 S,
46°47.24 E
Protected area Present 50 0.25 2010 Literature Razafindramanana,
Ambinany (Berenty region) 24°59.53 S,
46°18.07 E
Private reserve Present 65 2015 Literature Dimilahy et al., 2015
Andohahela 24°08.72 S,
43°49.74 E
National park Present*82 760 2000 Literature Moniac and Heitman,
Andranobe (Ambolavoa
21°57.78 S,
46°56.73 E
Present 36 0.381 2013 Literature Gould and
Andrianomena, 2015
Andringitra 22°06.76 S,
46°55.38 E
National park Present 106 31,134 1995 Literature Rakotoarisoa, 1999
Anja Miray (Ambalovoa
21°51.12 S,
46°50.40 E
Present 210 0.36 2013 Literature Gould and
Andrianomena, 2015
Anjapolo (Berenty region) 24°57.66 S,
46°15.06 E
Private reserve Present 151 2015 Literature Dimilahy et al., 2015
Bealoka (Berenty region) 25°00.59 S,
46°18.51 E
Private reserve Present*163 1 2009 Literature Dimilahy et al., 2015;
Sussman et al., 2006
Berenty (Berenty region) 24°59.88 S,
46°19.87 E
Private reserve Present 562 2 2010 Literature Dimilahy et al., 2015
Beza Mahafaly Parcel 1 23°42.00 S,
44°32.33 E
Special reserve Present 124 0.8 2014 Pers. commun. Sauther, pers. commun.
Beza Mahafaly Parcel 2 23°42.00 S,
44°34.33 E
Special reserve Present 205 5.2 2014 Pers. commun. Sauther, pers. commun.
Cap Sainte-Marieb 25°35.75 S,
45°08.35 E
Special reserve Locally or nearly
locally extinct*
50 0.18 2008 Literature Kelley, 2013
Chameleon (Tsaranoro
Unknown Community-level
Present 28 0.08 2013 Literature Gould and
Andrianomena, 2015
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324 LaFleur/Clarke/Reuter/Schaeffer
Location name Coordinates Protected status Population category Number
Size of forest
fragment or
areaa, km2
Year of
Source of
Days of
Citation (most recent
Ifaty 23°08.71 S,
43°36.95 E
Unprotected Extinct*0 – 2015;
Literature; pers.
–, 2 LaFleur et al., 2015
Isalo (Maki canyon)c22°29.26 S,
45°22.73 E
National park Present 118 0.5 1995;
Literature; pers.
–, 1 Hawkins, 1999
Isalo (Namaza circuit)c23°31.82 S,
43°46.01 E
National park Present 1995;
Literature; pers.
–, 1 Hawkins, 1999
Kalambatritra 23°20.50 S,
46°28.66 E
Special reserve Locally or nearly
locally extinct*
0 282.55 2005 Literature Irwin et al., 2005d
Kirindy Mitea 20°44.38 S,
44°10.32 E
National park Extinct*0 722 2016 Pers. commun.;
pers. observati-
–, 1 Lewis B., pers.
Goodman, 2003
Kirindy Mitea de sud 20°44.23 S,
44°00.07 E
Unprotected Extinct*0 2016 Pers. observati-
–, 2 Goodman, 2003
Mangily (Reniala Private
23°07.59 S,
43°36.80 E
Unprotected Extinct*0 0.6 2015 Literature; pers.
–, 2 LaFleur et al., 2015;
Gardner and Davies,
Marody (Tsaranoro Valley) 22°05.57 S,
46°47.65 E
Present 15 0.028 2013 Literature Gould and
Andrianomena, 2015
Petriky 25°04.31 S,
46°53.10 E
Unprotected Locally or nearly
locally extinct*
4 11.8 2012 Literature Malone et al., 2013
Pic d’Ivohibe 22°30.00 S,
46°59.00 E
Special reserve Locally or nearly
locally extinct*
0 34.5 2005 Literature Irwin et al., 2004d
PK 32-Ranobe 22°59.82 S,
43°40.20 E
Protected area Extinct*0 200.46 2010 Literature Gardner and Davies,
Sakarivo Miray
(Ambolavoa region)
21°47.01 S,
46°52.07 E
Present 30 0.142 2013 Literature Gould and
Andrianomena, 2015
(Ambalavoa region)
21°47.18 S,
24°49.83 E
Present 21 0.203 2013 Literature Gould and
Andrianomena, 2015
Table 1 (continued)
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Location name Coordinates Protected status Population category Number
Size of forest
fragment or
areaa, km2
Year of
Source of
Days of
Citation (most recent
Tsaranoro (Tsaranoro
22°05.18 S,
46°46.32 E
Present 78 0.46 2013 Literature Gould and
Andrianomena, 2015
(northern boundary)
24°05.40 S,
43°49.80 E
National park Present 67 2016 Pers. observati-
10 (including
research for a
larger, ongoing
project on
lemur ecology)
(southern boundary)
24°37.98 S,
43°55.98 E
National park Present 40 2015 Pers. commun. Louis E.E., pers.
Tsinjoriake (Moringa
circuit)b23°31.31 S,
43°45.28 E
Protected area Locally or nearly
locally extinct*
5 50 2016 Pers. commun.;
pers. observati-
–, 3 Youssouf J., pers.
Tsinjoriake (southern
mangrove area)
23°32.18 S,
43°44.68 E
Protected area Locally or nearly
locally extinct*
0 2016 Pers. observati-
–, 3
Zombitse-Vohibasiae22°50.50 S,
44°40.91 E
National park Locally or nearly
locally extinct*
10 0.36 2016 Pers. commun.;
pers. observati-
–, 2 Madagascar National
Parks, pers. commun.;
Siers, 2007
Total 2,220
An asterisk indicates known exploitation pressures (e.g., hunting) at the location against L. catta. “Personal observation” indicates that we collected data during a site visit in 2016.
a These area estimates should be interpreted with caution as they were often not reported alongside density estimates but have been extracted from various, publicly available documents.
b Kelley [2013] reports a range of 35 50 individual ring-tailed lemurs at Cap Sainte-Marie. Similarly, although we failed to detect the presence of animals, a student of Dr. Youssouff reported seeing a
range of 4–5 animals in Tsinjoriake (Moringa circuit) in August 2016. For the purposes of this paper, we used the larger value for the number of individual lemurs at both sites.
c We confirmed the presence of ring-tailed lemurs at Isalo but were unable to sample all known groups. We thus used data from Hawkins [1999] to estimate the total number of animals. Hawkins [1999]
does not differentiate between the 2 populations of ring-tailed lemurs present, and thus we cite 1 value here for all of Isalo.
d Irwin et al. [2004] conducted extensive censuses at Pic d’Ivohibe and Kalambatritra. Their censuses do not report the presence of ring-tailed lemurs, and here we use these data to infer the absence of
the species.
e We failed to detect animals, but the Madagascar National Parks ecological monitoring team reported the presence of 10 animals in March 2016 (3 months previously).
Table 1 (continued)
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326 LaFleur/Clarke/Reuter/Schaeffer
Maki canyon, Namaza circuit, Tsimanampesotse northern boundary), we estimated how many
L. catta were likely present by counting animals in known groups. More specifically, at Isalo
(Maki canyon), we walked a circuit until we encountered ring-tailed lemur groups and then fol-
lowed the animals for a minimum of 1 h, in order to ascertain how many animals were travelling
with the group. At Isalo (Namaza circuit), we remained in a tourist picnic spot, and when ring-
tailed lemur groups came through we monitored/followed individuals to determine the group
size and identity. At Tsimanampesotse, we monitored known groups and detailed any intergroup
encounters with neighbouring/unknown groups. For sites not visited in person ( n = 22), we col-
lected data from the literature and from colleagues who had visited these sites ( Table1 ).
Given the variations in sample size, methodological approach, sampling effort, and date of
assessment, our compiled data largely violate assumptions of parametric and non-parametric
statistics. Thus, the majority of our analyses are descriptive in nature. We present the mean,
range, and median numbers of ring-tailed lemurs present at each of the 32 study sites, and com-
pare the likelihood of local extinction between protected and unprotected areas (Fisher exact test,
p = 0.05). We recognize that the data presented here are limited and must be treated with caution.
Both over- and underestimations are possible in the data presented here. Overestimations are
possible in the data that were collected several years ago [Gould and Andrianomena, 2015], as
animals may no longer be present. Underestimates are also possible in areas where sampling ef-
fort has not been sufficient, such as large protected areas without extensive monitoring (i.e., Tsi-
manampesotse National Park, Isalo National Park). Despite these limitations, the data presented
here are still informative in understanding decreasing population trends and the urgent conser-
vation threats faced by ring-tailed lemurs.
Parameters of the Data Set
We sourced data for 84% of the sites ( n = 27 out of 32) from information made
public in 2010 or thereafter ( Table1 ). Ring-tailed lemurs’ historical range covers
roughly 100,000 km
2 throughout southern Madagascar, although this generalized es-
timate includes both habitable (e.g., forested) and uninhabitable (e.g., rivers, cities)
areas [Sussman et al., 2006]. The sites assessed here covered the habitable area of ap-
proximately 33,208 km
2 ( Table1 ).
Estimated Population Size across 32 Sites
We estimate the presence/absence and total number of animals remaining at the
32 sites considered in this study. L. catta were deemed locally extinct or nearly lo-
cally extinct (i.e., we were unable to detect the animals’ presence; fewer than 5 animals
remaining at last known sighting) in 37.5% of the sites included in this study ( n = 12
of 32, Table1 ). A higher percentage of areas with some form of protection or man-
agement status (69% of n = 29 sites) had extant ring-tailed lemur populations, when
compared to unprotected areas (0% of n = 3 sites, Fisher exact test, n = 32, p = 0.0444,
sites with extinct and likely extinct populations grouped together).
The sites had an average of 72 ± 110 (mean ± SD; range: 0–562; median 36) in-
dividuals of L. catta per site ( Table1 ). In 2 instances the numbers of animals were
reported as a range including 35–50 individual ring-tailed lemurs at Cap Sainte-Ma-
rie [Kelley, 2013] and 4–5 individual ring-tailed lemurs at Tsinjoriake [Youssouff,
pers. commun.]. For these 2 instances we included the high estimates of the number
of animals present ( Table1 ). Only 8 of 32 sites had populations of more than 100 in-
dividuals ( Table1 ). Sites with some protection ( n = 28 sites) had an average of 79 ±
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113 individuals, while sites without protection ( n = 3 sites) had an average of 3 ± 1
individuals left at the site. From the sum of all sites examined, we estimate that there
are approximately 2,220 ring-tailed lemurs remaining in the sites included in this
study ( Table1 ).
We estimate that there are 2,220 L. catta remaining at the 32 sites included in this
study. This estimate may not account for all small forest fragments where ring-tailed
lemurs could persist or all possible groups of animals in unmonitored areas of large
National Parks (i.e., Tsimanampesoste, Isalo), and thus could underestimate the total
number of animals remaining. Alternatively, the estimate could include animals that
have been extirpated very recently, and thus be inflated. Given that the previous esti-
mates for individual wild ring-tailed lemurs were 2 orders of magnitude larger (i.e.,
751,251 [Sussman et al., 2006]), we are confident that there has been a rapid decrease
in the numbers of animals living in the wild in Madagascar. Though we acknowledge
that both ours and the population estimates of Sussman et al. [2006] were calculated
with caveats, this sharp decline is concerning. Both complete deforestation of land
parcels and decline of their lemur inhabitants [e.g., Kelley et al., 2007, Antserana-
nomby Forest] and localized extirpations of ring-tailed lemurs without forest clearing
(e.g., Ranobe area [Gardner and Davies, 2014]) have been previously documented.
Current and detailed data regarding the numbers of ring-tailed lemurs persisting in
all of their habitats are urgently needed to understand fully the dramatic decline in
wild ring-tailed lemurs and prevent complete extinction in the wild.
The rapid decline in populations of L. catta has implications for conservation of
the species. First, we found only 1 site with a population of >500 L. catta ( Table1 ). In
Uganda, sites with <500 chimpanzees ( Pan troglodytes ) are of conservation concern
as smaller populations are not able to sustain genetic viability in the long term
[Plumptre and Cox 2006]. In the case of L. catta , Clarke et al. [2015] found that ring-
tailed lemur populations with fewer than 500 individuals in the central highlands
maintain moderate levels of genetic diversity. These populations, however, are likely
exhibiting a time lag response whereby genetic diversity will decrease substantially in
the future due to continued population fragmentation and associated impacts [Clarke
et al., 2015]. Both the low numbers of individuals at isolated sites (and the individu-
als’ potential inability to find mates), and the consequences of low genetic diversity
are problematic for ring-tailed lemurs’ long-term survival in the wild. Second, we
found that unprotected areas are more likely to experience population extinctions of
ring-tailed lemurs. The general effectiveness of protected areas in maintaining species
presence has been documented globally [see Naughton-Treves et al., 2005], and with-
in the order Primates [see Pusey et al., 2007], although effective management of pro-
tected areas is also necessary, as has been noted for lemur species specifically [Schwit-
zer et al., 2014]. Madagascar has been plagued with political turmoil, yet even during
times of crisis, conservation threats are more significant outside protected areas
[Schwitzer et al., 2014]. The combination of higher levels of genetic diversity within
larger populations and better survivorship within protected areas suggests that con-
servation priorities for ring-tailed lemurs should include areas with more animals and
established protection ( Table1 ).
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328 LaFleur/Clarke/Reuter/Schaeffer
A t t h e g l o b a l s c a l e , p o p u l a t i o n e x t i n c t i o n s a r e a m o r e s e n s i t i v e i n d i c a t o r o f t h e l o s s
of biodiversity than species extinctions [Ceballos and Ehrlich, 2002]. Likewise, because
L. catta i s k n o w n f o r i t s e c o l o g i c a l f l e x i b i l i t y [ S a u t h e r e t a l . , 1 9 9 9 ] , i t c o u l d b e a p r o x y
for biodiversity loss as its population decline underscores the difficulty that other, less
adaptable species have in overcoming a multitude of threats. As a well-known and fre-
quently encountered species (i.e., globally in zoos, and at select popular tourist sites in
Madagascar), the overarching perception is that L. catta i s n o t g l o b a l l y t h r e a t e n e d , n o r
in imminent threat of extinction. A primate species’ prevalence in the media has been
associated with the belief that wild populations are stable and not threatened by extinc-
tion [Leighty et al., 2015; Nekaris et al., 2016]. This false impression that ring-tailed
lemurs are not threatened with extinction could be devastating for the species (and
larger communities of biodiversity in Madagascar) as it may impede the ability to se-
cure funding for rapid conservation action. That being said, L. catta i s c u r r e n t l y u n d e r
consideration for inclusion in the 2016–2018 “World’s 25 Most Endangered Primates”
listing, which is published bi-annually by the IUCN/SSC Primate Specialist Group
[IUCN, 2016]. Inclusion of ring-tailed lemurs in this list will highlight the threats these
animals are facing and inform the larger conservation community about the fact that
even adaptable, flexible species such as ring-tailed lemurs can experience rapid decline.
Several urgent conservation actions should be considered with regard to protect-
ing L. catta from extinction in the wild. First, it has been suggested that primate pop-
ulations threatened with extinction should be regularly monitored using routine site
visits [Plumptre and Cox, 2006]. For L. catta this monitoring is especially necessary
at sites that have large remaining populations and in areas that are data deficient; a
complete site-based census of L. catta sites is recommended. Second, long-term ge-
netic viability is greater in populations with more individuals [Plumptre and Cox,
2006]. Since there are likely only 8 ring-tailed lemur populations remaining that have
more than 100 individual animals ( Table1 ), these populations should be targeted for
protection in order to maintain genetic diversity of the species. Additionally, reintro-
duction of captive ring-tailed lemurs, particularly those confiscated from the illegal
pet trade [LaFleur et al., 2015] could be used as a mechanism to increase wild genetic
diversity and to repopulate forested areas where populations are now extinct. Rein-
troductions are not without risks however, and comprehensive assessments of both
prospective animals and release sites would need to be carried out, as per IUCN re-
introduction guidelines [IUCN/SSC, 2013]. Third, live capture for the within-coun-
try pet trade poses a significant threat to many lemur species. More than 28,000 le-
murs were kept in illegal captivity between 2010 and mid-2013, and of these ring-
tailed lemurs were the most prevalent, comprising 28% of reported animals [Reuter
et al., 2016]. Information regarding drivers and locations of capture, trade routes, and
absolute numbers of ring-tailed lemurs in the illegal pet trade is imperative to future
conservation initiatives [Reuter and Schaefer, 2016].
Madagascar is one of the world’s poorest countries, and poverty is a significant
predictor of extinction for primate species [Estrada, 2013]. The primary threats to
ring-tailed lemurs, including deforestation and capture for the bushmeat and pet
trades, can each be linked to poverty in that subsistence living depends heavily on
natural resource exploitation [Brimont et al., 2015] and that forest products, includ-
ing lemurs, can be sold to generate income [Gardner and Davis, 2014]. It is critical to
link conservation efforts with social and economic development projects, in order to
ensure that both Madagascar’s human population and wildlife thrive.
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Folia Primatol 2016;87:320–330
DOI: 10.1159/000455121
The authors thank the Government of Madagascar, Ministry of the Environment and Ecol-
ogy of Forests, Madagascar National Parks, the University of Toliara, and MICET, for their per-
missions and facilitations of our research. We also wish to thank Lydia Greene and Holly Schnei-
der-Brown for their dedication to forwarding lemur conservation and awareness. Additionally,
we thank Dr. Anna Nekaris and 3 anonymous reviewers for their feedback, which significantly
improved this paper. Funding was provided by the Margot Marsh Biodiversity Foundation, Mar-
got Marsh Primate Action Fund, Duke University’s Arts and Sciences Council: Committee on
Faculty Research Grant (awarded to T.A.C.), and Lemur Love Inc. private donors.
Disclosure Statement
We understand Folia Primatologica ’s declaration of interests and declare that we have no
competing interests.
Andriaholinirina N, Baden A, Blanco M, Chikhi L, Cooke A, Davies N, Dolch R, Donati G, Ganzhorn J,
Golden C, Groeneveld LF, Hapke A, Irwin M, Johnson S, Kappeler P, King T, Lewis R, Louis EE,
Markolf M, Mass V, Mittermeier RA, Nichols R, Patel E, Rabarivola CJ, Raharivololona B, Rajaobe-
lina S, Rakotoarisoa G, et al. (2014). Lemur catta . The IUCN Red List of Threatened Species 2014:
Brinkmann K, Noromiarilanto F, Ratovonamana RY, Buerkert A (2014). Deforestation processes in
south-western Madagascar over the past 40 years: what can we learn from settlement characteristics?
Agriculture, Ecosystems & Environment 195: 231–243.
Brimont L, Ezzine-de-Blas D, Karsenty A, Toulon A (2015). Achieving conservation and equity amidst
extreme poverty and climate risk: the Makira REDD+ project in Madagascar. Forests 6: 748–768.
Brown JL, Yoder AD (2015). Shifting ranges and conservation challenges for lemurs in the face of climate
change. Ecology and Evolution 5: 1131–1142.
Ceballos G, Ehrlich PR (2002). Mammal population losses and the extinction crisis. Science 296: 904–907.
Clarke TA, Gray O, Gould L, Burrell AS (2015). Genetic diversity of the ring-tailed lemur ( Lemur catta )
in south-central Madagascar. Folia Primatologica 86: 76–84.
Dimilahy L, Foulon C, Mhia P, Rahanitriniaina O, Andrianarimalala V, Fenomanana R, Randrianady J,
Ratovolala M, Sehenomalala C, Walker-Bolton A, Tahirisoa R, Rafenoarisoa F, Soppelsa J, Randri-
anirina T, Trapon D, Ravahatramanan-Jarasoa FN (2015). Livret de recensement 2015 Lemur catta,
Réserve Privée de Berenty 1936–2015. Unpublished report.
Estrada A (2013). Socioeconomic context of primate conservation: population, poverty, global economic
demands and sustainable land use. American Journal of Primatology 75: 30–45.
Gardner CJ, Davies ZG (2014). Rural bushmeat consumption within multiple-use protected areas: quali-
tative evidence from southwest Madagascar. Human Ecology 42: 21–34.
Gese EM (2001). Monitoring of terrestrial carnivore populations. USDA National Wildlife Research Cen-
ter Publication No 576.
Gilles H, Reuter KE (2014). The presence of diurnal lemurs and human-lemur interactions within the buf-
fer zone of the Ankarana National Park. Lemur News 18: 27–30.
Goodman S (2003). Hunting of wild animals by Sakalava of the Menabe region: a field report from Kirin-
dy-Mite. Lemur News 8: 4–6.
Goodman SM, Rakotoarisoa SV, Wilmé L (2006). The distribution and biogeography of the ringtailed
lemur ( Lemur catta ) in Madagascar. In Ringtailed Lemur Biology (Jolly A, Sussman RW, Koyama N,
Rasamimanana H, eds.), pp 3–15. New York, Springer.
Gould L, Andrianomena P (2015). Ring-tailed lemurs ( Lemur catta ), forest fragments, and community-
level conservation in south-central Madagascar. Primate Conservation 29: 67–73.
Gould L, Sussman RW, Sauther ML (1999). Natural disasters and primate populations: the effects of a
2-year drought on a naturally occurring population of ring-tailed lemurs (Lemur catta) in south-
western Madagascar. International Journal of Primatology 20: 69–84.
Hawkins A (1999). The primates of Isalo National Park, Madagascar. Lemur News 4: 11–14.
Downloaded by:
Temple University - 1/13/2017 2:51:32 PM
Folia Primatol 2016;87:320–330
DOI: 10.1159/000455121
330 LaFleur/Clarke/Reuter/Schaeffer
Irwin MT, Samonds KE, Raharison JL, Wright PC (2004). Lemur latrines: observations of latrine behavior
in wild primates and possible ecological significance. Journal of Mammalogy 85: 420–427.
Irwin MT, Johnson SE, Wright PC (2005). The state of lemur conservation in southeastern Madagascar:
population and habitat assessments for diurnal and cathemeral lemurs using surveys, satellite imag-
ery and GIS. Oryx 39: 204–218.
IUCN (2016). The IUCN Red List of Threatened Species. Version 2016-1 . (ac-
cessed June 30, 2016).
IUCN/SSC (2013). Guidelines for Reintroductions and Other Conservation Translocations . Version 1.0 .
Gland, IUCN Species Survival Commission.
Kelley EA (2013). The ranging behavior of Lemur catta in the region of Cap Sainte-Marie, Madagascar.
American Journal of Physical Anthropology 150: 122–132.
Kelley EA, Sussman RW, Muldoon KM (2007). The status of lemur species and Antserananomby: an up-
date. Primate Conservation 11: 71–77.
LaFleur M, Clarke T, Giraud L, Youssouf J, Gould L, Adiba M (2015). Reniala Lemur Rescue Center for
ring-tailed lemurs in Madagascar. Lemur News 19: 11–13.
Leighty KA, Valuska AJ, Grand AP, Bettinger TL, Mellen JD, Ross SR, Boyle P, Ogden JJ (2015). Impact
of visual context on public perceptions of non-human primate performers. PLoS One 10: e0118487.
Macdonald DW, Mace GM, Barreto GR (1999). The effects of predators on fragmented prey populations:
a case study for the conservation of endangered prey. Journal of Zoology 247: 487–506.
Malone M, Ramanamanjato JB, Randriatfika F, Donati G (2013). Habitat structure and grey mouse lemur
( Microcebus murinus ) abundance in the transitional littoral forest of Petriky, south-east Madagas-
car. Lemur News 17: 22–26.
Moniac N, Heitmann A (2007). Lemur catta and hunting around Andohahela. Lemur News 12: 11.
Naughton-Treves L, Holland MB, Brandon K (2005). The role of protected areas in conserving biodiver-
sity and sustaining local livelihoods. Annual Review of Environment and Resources 30: 219–252.
Nekaris KAI, Musing L, Vazquez AG, Donati G (2016). Is tickling torture? Assessing welfare towards slow
lorises ( Nycticebus spp.) within web 2.0 videos. Folia Primatologica 86: 534–551.
Plumptre AJ, Cox D (2006). Counting primates for conservation: primate surveys in Uganda. Primates 47:
Pusey A, Pintea L, Wilson ML, Kamenya S, Goodall J (2007). The contribution of long-term research at
Gombe National Park to chimpanzee conservation. Conservation Biology 21: 623–634.
Rakotoarisoa SV (1999). Contribution à l’étude de l’adaptation de Lemur catta Linnaeus, 1758, aux zones
sommitales de la Réserve Naturelle d’Andringitra. Mémoire DEA, Université d’Antananarivo.
Razafindramanana J (2011). Behavioural Ecology of Sympatric Lemur Species Lemur catta and Eulemur sp.
in Forest Fragments, South-Eastern Madagascar. D o c t o r a l d i s s e r t a t i o n , O x f o r d B r o o k e s U n i v e r s i t y .
Reuter KE, Schaefer MS (in press). Motivations for the ownership of captive lemurs in Madagascar. An-
throzoos .
Reuter KE, Schaefer MS (2016). Illegal captive lemurs in Madagascar: comparing the use of online and
in-person data collection methods. American Journal of Primatology DOI: 10.1002/ajp.22541.
Reuter KE, Schaefer MS (2016). Captive conditions of pet lemurs in Madagascar. Folia Primatologica 8 7 :
Reuter KE, Gilles H, Wills AR, Sewall BJ (2016). Live capture and ownership of pet lemurs in Madagascar:
extent and conservation implications. Oryx 40: 344–354.
Sauther ML, Sussman RW, Gould L (1999). The socioecology of the ring-tailed lemur: thirty-five years of
research. Evolutionary Anthropology 8: 102–132.
Schwitzer C, Mittermeier RA, Davies N, Johnson S, Ratsimbazafy J, Razafindramanana J, Louis EE Jr, Ra-
jaobelina S (eds.) (2013). Lemurs of Madagascar: A Strategy for their Conservation 2013–2016. Bris-
tol, IUCN SSC Primate Specialist Group, Bristol Conservation and Science Foundation, and Con-
servation International.
Schwitzer C, Mittermeier RA, Johnson SE, Ratsimbazafy J, Razafindramanana J, Louis EE, Chikhi L,
Colquhoun IC, Crick J, Dolch R, Donati G, Irwin M, LaFleur M, Nash S, Patel E, Peacock H, Ran-
drianambinina B, Rasolofoharivelo T, Wright P (2014). Averting lemur extinctions amid Madagas-
car’s political crisis. Science 343: 842–843.
Siers RS (2007). Research experience in Vohibasia Forest, southwestern Madagaskar. Lemur News 12:
Sussman RW, Green GM, Porton I, Andrianasolondraibe OL, Ratsirarson J (2003). A survey of the habi-
tat of Lemur catta in southwestern and southern Madagascar. Primate Conservation 19: 1631.
Sussman RW, Sweeney S, Green GM, Porton I, Andrianasolondraibe OL, Ratsirarson J (2006). A pre-
liminary estimate of Lemur catta population density using satellite imagery. In Ringtailed Lemur
Biology (Jolly A, Sussman RW, Koyama N, Rasamimanana H, eds.), pp 16–31. New York, Springer.
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... This study aims to assess the learning experiences available at zoos and their retrospectively quality in terms of knowledge retention: using a well-known species as a case-study: the ringtailed lemur (Lemur catta). Lemurs are the most threatened mammal group (LaFleur, et al., 2017, Estrada, et al., 2017, and are "the single highest conservation priority in the world" (Schwitzer, et al. 2013). ...
... Acting as a flagship species for Madagascar, the ringtailed lemur is the most common lemur kept in captivity (Andriaholinirina, et al., 2014;LaFleur, et al., 2017), with a current captive population totalling 2,500 individuals (ISIS, 2006, in Andriaholinrina, et al., 2014. ...
... Despite the sustainable captive population, wild L. catta populations saw a 20% decrease between 1985 to 2000 (Sussman, et al., 2006) and further local extinctions suspected a decline rate of over 50%, estimating the current population between 2000 to 2400 individuals (Andriaholinirina, et al., 2014;LaFleur, et al., 2017LaFleur, et al., , 2018. The result reflected previous population trends; seeing the species upgraded from Near Threatened to Endangered on the IUCN Red List (Andriaholinirina, et al., 2014;Gould & Sauther, 2016), and being identified as one of the world's top 25 endangered primates (Schwitzer, et al., 2017). ...
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Environmental education programs focus on improving the public’s knowledge and support of conservation motives, improving polices, and making decisions which have an overall positive effect on the natural world . Zoological establishes zoos are in a critical position for providing educational opportunities to the public, as both direct and indirect approaches. Using a simple survey to collate data, this study assess the learning experiences available at zoos and their retrospectively quality in terms of knowledge retention using a well known captive species; Lemur catta. This study found that visitors who have been exposed to more focused education experiences will have greater general knowledge regarding lemur species and L. catta in particular, as there were significant positive correlations between learning categories. With this in mind, establishments should aim to continue providing regular talks and review the content and species targeted to maximise effectiveness.
... Ring-tailed lemurs are under severe conservation pressure; they are classified as "endangered" by the International Union for Conservation of Nature (IUCN) [8], resulting primarily from deforestation, hunting, and capture for the pet trade. A recent population census has revealed a dramatic population decline, with as few as 2,200 individuals remaining in the wild [9]. Of further concern, the species is distributed across a highly fragmented range with only 8 populations of ≥100 individuals remaining [9]. ...
... A recent population census has revealed a dramatic population decline, with as few as 2,200 individuals remaining in the wild [9]. Of further concern, the species is distributed across a highly fragmented range with only 8 populations of ≥100 individuals remaining [9]. Despite this near-term population decline, a recent microsatellite analysis indicates that the genetic diversity of L. catta populations could be exceptionally high, with evidence of genetic isolation by distance throughout their geographic range [6]. ...
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Background: The ring-tailed lemur (Lemur catta) is a charismatic strepsirrhine primate endemic to Madagascar. These lemurs are of particular interest, given their status as a flagship species and widespread publicity in the popular media. Unfortunately, a recent population decline has resulted in the census population decreasing to <2,500 individuals in the wild, and the species's classification as an endangered species by the IUCN. As is the case for most strepsirrhine primates, only a limited amount of genomic research has been conducted on L. catta, in part owing to the lack of genomic resources. Results: We generated a new high-quality reference genome assembly for L. catta (mLemCat1) that conforms to the standards of the Vertebrate Genomes Project. This new long-read assembly is composed of Pacific Biosciences continuous long reads (CLR data), Optical Mapping Bionano reads, Arima HiC data, and 10X linked reads. The contiguity and completeness of the assembly are extremely high, with scaffold and contig N50 values of 90.982 and 10.570 Mb, respectively. Additionally, when compared to other high-quality primate assemblies, L. catta has the lowest reported number of Alu elements, which results predominantly from a lack of AluS and AluY elements. Conclusions: mLemCat1 is an excellent genomic resource not only for the ring-tailed lemur community, but also for other members of the Lemuridae family, and is the first very long read assembly for a strepsirrhine.
... conservation genetics, Madagascar, microsatellites, strepsirrhines have lost over 45% of their habitat (Brinkmann, Noromiarilanto, Ratovonamana, & Buerkert, 2014;LaFleur, Clarke, Ratzimbazafy, & Reuter, 2017a); and by 2080, it is estimated that 63% of their remaining range will shift due to climate change alone (Brown & Yoder, 2015). Furthermore, there has been a recent uptick in exploitation for the illegal pet trade, causing severe population declines, and in some cases local extinctions, throughout their remaining geographic range (Gardner & Davies, 2014;Gould & Sauther, 2016;LaFleur, Clarke, Reuter, Schaefer, & ter-Horst, 2019;LaFleur, Clarke, Reuter, & Schaeffer, 2017b;LaFleur & Gould, 2009;Reuter et al., 2019;Reuter & Schaefer, 2016). At present, there are estimated to be fewer than 2,400 individuals within sampled locations (Gould & Sauther, 2016;LaFleur et al., 2017b), though population estimates are still lacking throughout much of their range (e.g., Murphy, Ferguson, & Gardner, 2017). ...
... Furthermore, there has been a recent uptick in exploitation for the illegal pet trade, causing severe population declines, and in some cases local extinctions, throughout their remaining geographic range (Gardner & Davies, 2014;Gould & Sauther, 2016;LaFleur, Clarke, Reuter, Schaefer, & ter-Horst, 2019;LaFleur, Clarke, Reuter, & Schaeffer, 2017b;LaFleur & Gould, 2009;Reuter et al., 2019;Reuter & Schaefer, 2016). At present, there are estimated to be fewer than 2,400 individuals within sampled locations (Gould & Sauther, 2016;LaFleur et al., 2017b), though population estimates are still lacking throughout much of their range (e.g., Murphy, Ferguson, & Gardner, 2017). ...
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Madagascar's ring‐tailed lemurs (Lemur catta ) are experiencing rapid population declines due to ongoing habitat loss and fragmentation, as well as increasing exploitation for bushmeat and the illegal pet trade. Despite being the focus of extensive and ongoing behavioral studies, there is comparatively little known about the genetic population structuring of the species. Here, we present the most comprehensive population genetic analysis of ring‐tailed lemurs to date from across their likely remaining geographic range. We assessed levels of genetic diversity and population genetic structure using multilocus genotypes for 106 adult individuals from nine geographically representative localities. Population structure and F ST analyses revealed moderate genetic differentiation with localities being geographically partitioned into northern, southern, western and also potentially central clusters. Overall genetic diversity, in terms of allelic richness and observed heterozygosity, was high in the species (AR = 4.74, H O = 0.811). In fact, it is the highest among all published lemur estimates to date. While these results are encouraging, ring‐tailed lemurs are currently affected by ongoing habitat fragmentation and occur at lower densities in poorer quality habitats. The effects of continued isolation and fragmentation, coupled with climate‐driven environmental instability, will therefore likely impede the long‐term viability of the species.
... Among other issues were the status of Lemur catta and Propithecus verreauxi and of Lepilemur leucopus in the south of Madagascar. While L. catta and P. verreauxi have a large area of distribution, the actual presence of Lemur catta at many sites within this area of distribution is uncertain and thus the area of occupancy might be substantially smaller that suggested by maps showing the area of distribution (Gould and Sauther, 2016;LaFleur et al., 2016;Murphy et al., 2017). ...
... Increased rates of deforestation in Madagascar, primarily driven by the need to meet economic and subsistence needs at the household level, have resulted in high degrees of habitat fragmentation (Schwitzer et al., 2013a). Fragmented, patchy areas lead to increased encounters with lemurs, increases in hunting opportunities (e.g., to meet subsistence needs, supply wildlife trade), and in turn potential disease transmission (C2; C7; C9; D1) (Barrett and Ratsimbazafy, 2009;Gilles and Reuter, 2014;LaFleur et al., 2016LaFleur et al., , 2018LaFleur et al., , 2019. The decline in the populations of red-collared brown lemurs (Eulemur collaris) in Saint Luce (southeast Madagascar) is attributed to hunting, although a population persists in the littoral forest fragments of the area (Roberts et al., 2019/20). ...
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Globally, non-human primates face mounting threats due to unsustainable harvest by humans. There is a need to better understand the diverse drivers of primate harvest and the complex social-ecological interactions influencing harvest in shared human-primate systems. Here, we take an interdisciplinary, systems approach to assess how complex interactions among primate biological and ecological characteristics and human social factors affect primate harvest. We apply our approach through a review and synthesis of the literature on lemur harvest in Madagascar, a country with one of the highest primate species richness in the world coupled with high rates of threatened primate species and populations in decline. We identify social and ecological factors affecting primate harvest, including the characteristics of lemurs that may make them vulnerable to harvest by humans; factors describing human motivations for (or deterrents to) harvest; and political and governance factors related to power and accessibility. We then discuss social-ecological interactions that emerge from: (1) the prevalence of informal institutions (e.g., cultural taboos), (2) adoption of human predatory strategies, (3) synergies with habitat use and habitat loss, and (4) interactions among regional- and local-scale factors (multi-level interactions). Our results illustrate that social-ecological interactions influencing lemur harvest in Madagascar are complex and context-specific, while influenced by a combination of interactions between species-specific characteristics and human social factors. These context-specific interactions may be also influenced by local-level cultural practices, land use change, and effects from regional-level social complexities such as political upheaval and food insecurity. We conclude by discussing the importance of identifying and explicitly accounting for nuances in underlying social-ecological systems and putting forth ideas for future research on primate harvest in shared human-primate systems, including research on social-ecological feedbacks and the application of Routine Activities Theory.
... Where disagreements among researchers and those in the conservation community have been unresolved, they have in some cases undermined the list's potential desired impact. For example, ring-tailed lemurs ( Lemur catta ) were included in the 2016-2018 edition following worrying signs of rapid population declines due to drought ( Vyawahere, 2020 ), habitat degradation, and the pet trade ( Gould & Sather, 2016 ;LaFleur, Clarke, Reuter, & Schae er, 2017 ). Ring-tailed lemurs, a fl agship species for the Madagascar tourism industry, have a large range in the country's south. ...
... It seems, however, that the trade in galagos as pets in general is only small scale and often opportunistic, as has been found for the nocturnal pottos and angwantibos (Svensson and Friant 2014). Capture and trade of wild ring-tailed lemurs (Lemur catta) is similarly opportunistic but, once believed to be wide ranging and common, they were estimated to have suffered a population decline of nearly 95% since 2000, mainly driven by bushmeat and the pet trade (LaFleur et al. 2016). ...
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Primates are traded yearly in the tens of thousands for reasons such as biomedical research, as trophies and pets, for consumption and to be used in traditional medicine. In many cases, this trade is illegal, unsustainable and considered a major impediment to primate conservation. Diurnal primates make up the vast majority of this trade, but recent studies have found that the trade in nocturnal primates is more common than previously thought, and among them are the galagos. There are currently 19 galagos recognized but there is still a dearth of research on these species and subspecies. The purpose of our study was to provide a more comprehensive picture of the trade in galagos within and across their African range countries, to help determine whether it is illegal or its sustainability needs to be assessed, and to provide baseline data and management recommendations to better regulate this trade, including strengthening policy, enforcement and conservation interventions. We gathered information on trade and use of galagos using an online questionnaire (May-August 2020), and on country-specific legislation relating to wildlife trade, hunting and legal protection of galagos, and looked at each range country's Corruption Perception Index score to gain an understanding of the obstacles in the way of effective law enforcement. We received 140 responses to our online questionnaire, from 31 of the 39 galago range countries. Respondents from 16 of these countries reported on first-hand observations of galagos being traded or used. Out of these, 36% reported seeing galagos sold or used for consumption, 33% as pets and 25% had observed them sold or used for traditional practices (including medical and magical purposes and for witchcraft). Most reports came from West Africa followed by Central Africa, East Africa and Southern Africa. We found that the number of reports on galagos being traded was higher in countries with higher numbers of galago species. Countries with more restrictive legislation experienced a higher number of reports of trade. Galagos observed in the pet trade was more common in East Africa, whilst reports of them in the bushmeat trade were more common in Central and West Africa. Galagos observed in the trade for traditional practices was by far most common from West Africa. We found that all galago range countries have some level of legal protection for some or all of their native galago species. It is evident that use and trade of galagos occurs throughout their range, albeit localized to certain areas. We urge galago range countries to adequately protect all species and to ensure legal trade is effectively regulated. Range countries that prohibit the use and trade in galagos must ensure legislation is adequately enforced. Further research into the drivers behind the use and trade of galagos should be initiated in countries with high levels of use and trade to further inform conservation and policy actions and to catalyze enforcement actions against poaching and illegal trade.
... The dry forests of Madagascar's southern regions have been reported as the most seriously fragmented forest type in the country, and such fragmentation has occurred due to factors such as extensive slash and burn practices for cattle grazing, charcoal production, and various forms of agriculture (Bodin, Tengo, Norman, Lundberg, & Elmqvist, 2006;Harper, Steininger, Compton, Tucker, & Hawkins, 2007;Sauther et al., 2013;Scott et al., 2006). Such habitat loss, as well as illegal bushmeat hunting for local consumption (Borgerson, McKean, Sutherland, & Godfrey, 2016;Golden, 2009) and capture for the illegal pet trade (LaFleur, Clarke, Reuter, Schaefer, & terHorst, 2019;Reuter, Gilles, Wills, & Sewall, 2016) have resulted local extinctions or drastic reductions of ring-tailed lemur populations in many areas of southern Madagascar where they are or were previously reported (Gould & Sauther, 2016;Kelley, Sussman, & Muldoon, 2007;LaFleur, Clarke, Reuter, & Schaeffer, 2016;Sauther et al., 2013;Sussman et al., 2006). ...
Habitat fragmentation is an increasingly serious issue affecting primates in most regions where they are found today. Populations of Lemur catta (ring-tailed lemur) in Madagascar's south-central region are increasingly restricted to small, isolated forest fragments, surrounded by grasslands or small-scale agriculture. Our aim was to evaluate the potential for population viability of L. catta in nine forest fragments of varying sizes (2-46 ha, population range: 6-210 animals) in south-central Madagascar, using a set of comparative, quantitative ecological measures. We used Poisson regression models with a log link function to examine the effects of fragment size, within-fragment food availability, and abundance of matrix resources (food and water sources) on L. catta population sizes and juvenile recruitment. We found a strong association between overall population size and (a) fragment size and (b) abundance of key food resources Melia azedarach and Ficus spp. (per 100 m along transect lines). Juvenile recruitment was also associated with fragment size and abundance of the two above-mentioned food resources. When the largest population, an outlier, was removed from the analysis, again, the model containing fragment size and abundance of M. azedarach and Ficus spp. was the best fitting, but the model that best predicted juvenile recruitment contained only fragment size. While our results are useful for predicting population presence and possible persistence in these fragments, both the potential for male dispersal and the extent of human disturbance within most fragments play crucial roles regarding the likelihood of long-term L. catta survival. While seven of the nine fragments were reasonably protected from human disturbance, only three offered the strong potential for male dispersal, thus the long-term viability of many of these populations is highly uncertain.
... Current wild ring-tailed lemur populations are facing several anthropogenic threats, with multiple recent studies establishing a severe drop in numbers across 32-34 sites, with at least 12 showing local extinctions of populations LaFleur et al., 2016). An earlier study in 2012 found suggestions of a recent genetic bottleneck, further supporting the concern that ring-tailed lemur populations could become critically endangered (Parga et al., 2012). ...
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Lemurs are naturally endemic to the island of Madagascar. Captive lemur diets generally consist of commercial dry pellets supplemented with vegetables and forages. The dry pellets given are their main source of fibre and are key to ensuring all nutritional requirements are met in a man-made setting. These are often the blandest and least desirable part of the diet, and therefore usually fed first, separately to more palatable foods, and ad lib. Incorrect feeding in captivity for lemurs is a prominent issue seen in many institutions. Enrichment feeds have proven to have a beneficial effect on encouraging natural behaviours, such as grooming and foraging. However, there is minimal research on whether enrichment can be used to affect feed intake. Three species of lemur, the red ruffed lemur (Varecia rubra), the black and white ruffed lemur (Varecia variegate), and the ring-tailed lemur (Lemur catta) were given three different feeding enrichments. Two aims were being investigated: 1) whether presenting dry pellets through enrichment devices changes the overall feed intake in three species of lemur, and 2) whether different enrichment items have different effects on pellet consumption. Results presented no significant change in pellet consumption when fed through enrichment devices for ring-tailed lemurs (p>0.05), which was also the case for both ruffed lemur species (p>0.1). They also distinguished no significant difference between feeding enrichments in pellet intake in any of the three species of lemur. This study helped identify the possibility of using enrichment for pellet consumption in zoos, but showed that the devices used didn’t have a significant enough effect to suggest future implementation of enrichments for the sake of nutrition. Further research is needed to fully understand whether the application of enrichment can be beneficial for encouraging adequate diets in lemurs, and more factors need to be considered, such as species difference in locomotion and enclosure use.
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A key aspect of a primate’s ecology is its food source, the very nature of which is spatially and seasonally dependent and may be affected by anthropic pressures. One of the most endangered, yet best-studied, strepsirrhine primates is the ring-tailed lemur (Lemur catta), a species that has experienced significant human-induced habitat loss over many decades. To help understand feeding variability across time and space, I present a literature review of plant species (and parts) fed on by ring-tailed lemurs at nine sites in Madagascar: Ambatotsirongorongo, Andringitra Massif, Anja Reserve, Antserananomby, Berenty Reserve, Bezà Mahafaly Special Reserve, Cap Sainte-Marie, Tsaranoro Valley Forest, and Tsimanampetsotsa National Park. I gathered literature using keyword searches on Google Scholar ( and verified scientific names using the “Catalogue of the Plants of Madagascar” ( From 24 studies, I identify 221 genera and 241 species of consumed plants, with 92 genera and 70 species consumed at two or more sites. Based on the available distribution data, 63% of species are endemic and 22% native. Sixty-seven plants are known only by Malagasy common names and excluded from analyses. When authors identify the plant tissue consumed, 52% of species in the diet are represented by a single tissue type, typically leaves (mature and immature) or fruit (ripe or unripe). This review highlights the importance of studying multiple populations when creating dietary summaries of species and should prove valuable to those exploring ecological trends and habitat use by ring-tailed lemurs.
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Videos, memes and images of pet slow lorises have become increasingly popular on the Internet. Although some video sites allow viewers to tag material as 'animal cruelty', no site has yet acknowledged the presence of cruelty in slow loris videos. We examined 100 online videos to assess whether they violated the 'five freedoms' of animal welfare and whether presence or absence of these conditions contributed to the number of thumbs up and views received by the videos. We found that all 100 videos showed at least 1 condition known as negative for lorises, indicating absence of the necessary freedom; 4% showed only 1 condition, but in nearly one third (31.3%) all 5 chosen criteria were present, including human contact (57%), daylight (87%), signs of stress/ill health (53%), unnatural environment (91%) and isolation from conspecifics (77%). The public were more likely to like videos where a slow loris was kept in the light or displayed signs of stress. Recent work on primates has shown that imagery of primates in a human context can cause viewers to perceive them as less threatened. Prevalence of a positive public opinion of such videos is a real threat towards awareness of the conservation crisis faced by slow lorises.
Conference Paper
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Overexploitation represents a severe threat to biodiversity, with live capture affecting millions of animals yearly. An improved understanding of primate live capture is needed, especially for Madagascar’s endangered lemurs. Our objectives were to provide the first quantitative estimates of (1) the prevalence, spatial extent, correlates, and timing of lemur ownership; and (2) procurement methods, within-country movements, and numbers and durations of ownership. Using semi-structured interviews of 1093 households and 61 transporters across 17 study sites, we found that lemur ownership was common, widespread, and affected a variety of taxa. Lemur ownership is ongoing, impacting an estimated 28,253 animals since 2010. Most lemurs were caught by owners, and kept for short (≤ 1 week) or long (≥ 3 years) periods. The live capture of lemurs within Madagascar is not highly organized, but may threaten several already-endangered species.
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Madagascar's lemurs, now deemed the most endangered group of mammals, represent the highest primate conservation priority in the world. Due to anthropogenic disturbances, an estimated 10% of Malagasy forest cover remains. The endangered Lemur catta is endemic to the southern regions of Madagascar and now occupies primarily fragmented forest habitats. We examined the influence of habitat fragmentation and isolation on the genetic diversity of L. catta across 3 different forest fragments in south-central Madagascar. Our analysis revealed moderate levels of genetic diversity. Genetic differentiation among the sites ranged from 0.05 to 0.11. These data suggest that the L. catta populations within south-central Madagascar have not yet lost significant genetic variation. However, due to ongoing anthropogenic threats faced by ring-tailed lemurs, continued conservation and research initiatives are imperative for long-term viability of the species. © 2015 S. Karger AG, Basel.
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Achieving forest conservation together with poverty alleviation and equity is an unending challenge in the tropics. The Makira REDD+ pilot project located in northeastern Madagascar is a well-suited case to explore this challenge in conditions of extreme poverty and climatic vulnerability. We assessed the potential effect of project siting on the livelihoods of the local population and which households would be the most strongly impacted by conservation measures. Farmers living in hilly areas must resort to slash-and-burn agriculture (tavy) since a combination of topographic and climatic constraints, such as cyclones, makes permanent rice cultivation very difficult. These are the people who suffer most from conservation-related restriction measures. For practical reasons the project, unfortunately, did not target these farmers. The main focus was on communities with a lower cyclonic risk that are able to practice permanent rice agriculture in the lowlands. To reduce deforestation without violating the principles of equity, REDD+ projects in Madagascar need to better target populations facing high climatic risks and invest in efforts to improve the farmers' agricultural systems.
The live capture of primates is occurring throughout the tropics and can be a threat to their conservation. Primates are owned as pets for a variety of reasons. Studies of the motivations for primate ownership have been conducted in several countries where they are endemic, but no study has examined this issue in Madagascar. Madagascar is home to the highest number of threatened primate taxa in any one country, and an estimated 28,000 lemurs were kept in illegal captivity from 2010 to mid-2013. We aimed to expand knowledge about the motivations of lemur ownership in Madagascar. Data were collected via a web-based survey (n = 229 respondents) and from the websites and social media pages of 25 hotels. We found that many lemurs (45%) were seen on the premises of a business or in a private home (27%). Many lemurs were perceived to be kept as personal pets (37%) or for money-making or tourism purposes (20%). When lemurs were used for money-making, owners could receive indirect (72% of the time) and direct benefits (28%). Hotels showing photographs of captive lemurs on their websites and social media sites charged USD 25.69 more per night for a standard room than hotels that did not show such photographs. We found little evidence that captive lemurs are kept as a social status symbol, for captive breeding, or as a fallback food. These findings provide evidence that the motivations for the ownership of, usually illegal, captive lemurs is typically linked with money-making or with the desire to have a lemur as a pet. These data can help target new outreach programs.
Live extraction of wildlife is a threat to biodiversity and can compromise animal welfare standards. Studies of the captive environments and welfare of pet primates are known, but none has focused on Madagascar. We aimed to expand knowledge about the captive conditions of pet lemurs in Madagascar. We hypothesized that captive le- murs would often be kept in restrictive settings, including small cages, would be fed foods inconsistent with their natural diets and, as a result, would be in bad physical or psychological health. Data were collected via a web-based survey (n = 253 reports) and from the websites and social media pages of 25 hotels. Most lemurs seen by respon- dents were either kept on a rope/leash/chain or in a cage (67%), though some lemurs were habituated and were not restrained (28%). Most of the time (72%) cages were con- sidered small, and lemurs were rarely kept in captivity together with other lemurs (81% of lemurs were caged alone). Pet lemurs were often fed foods inconsistent with their natural diets, and most (53%) were described as being in bad health. These findings point to a need to undertake outreach to pet lemur owners in Madagascar about the captivity requirements of primates.
Although it is illegal to capture, sell, and trade lemurs, the live capture of lemurs in Madagascar is ongoing and may have impacted over 28,000 lemurs between 2010 and 2013. Only one study has examined this trade and did so using in-person interviews in northern Madagascar. The current study sought to expand this existing dataset and examine the comparability of online surveys to more traditional on-location data collection methods. In this study, we collected data through a web-based survey resulting in 302 sightings of 685 captive lemurs. We also collected data from 171 hotel and 43 restaurant websites and social media profiles. Survey submissions included sightings of 30 species from 10 genera, nearly twice as many species as identified via the in-person interviews. Lemur catta, Varecia variegata, and Eulemur fulvus were the most common species sighted in captivity. Captive lemurs were reported in 19 of Madagascar's 22 administrative regions and most were seen in urban areas near their habitat ranges. This represents a wider geographic distribution of captive lemurs than previously found through in-person interviews. The online survey results were broadly similar to those of the in-person surveys though greater in species and geographic diversity demonstrating advantages to the use of online surveys. The online research methods were low in cost (USD $100) compared to on-location data collection (USD $12,000). Identified disadvantages included sample bias; most of the respondents to the online survey were researchers and many captive sightings were near study sites. The results illustrate the benefits of incorporating a social science approach using online surveys as a complement to traditional fieldwork. Am. J. Primatol. © 2016 Wiley Periodicals, Inc.
The south-central highland region of Madagascar is largely deforested, except for rare, small forest fragments scattered across the landscape. Some of these fragments are now being managed by local community conservation associations, after village residents have seen, first hand, how decades of deforestation and, more recently, climate change in the region have affected their agricultural crops. Furthermore, ecotourism is becoming more common in this area, with the two largest fragments, Anja Community Reserve and the Tsaranoro Valley forest, frequently receiving both Malagasy and foreign visitors. The combination of revenue from such tourism and the prospects of greater food security (by conserving and expanding existing fragments, leading to better soil conditions for nearby crops and rice fields) has resulted in increased fragment preservation over the past few years. We surveyed seven of these fragments in August and September, 2013, with two goals: (1) to evaluate the viability of ring-tailed lemur populations in each fragment in relation to food and water resource availability; and (2) to discuss, with village conservation association leaders, the goals of each association regarding fragment conservation and benefits to local residents. Anja Reserve, operating since 1999, is highly successful, and the association's ecotourism and community development projects have won them national and international recognition. In 2012, village associations Sakaviro Miray, FI.MI.VA Samisorany, and Antokinihoavy-Andranobe received NGO funding to help build a tourism presence in the region, and infrastructure for such activities had begun when we visited these sites in 2013. The Tsaranoro Valley, where three of the fragments are located, is an adventure tourism destination; however, much of the tourism revenue goes to the owners of the trekking camps, although a portion of the funds are donated to the local Association Tantely each year, and these camps also provide jobs for local villagers. Recent efforts are underway to expand these three fragments via reforestation, and to create a corridor between two of them in order to assist with the dispersal of L. catta males, and thus assist with potential population gene flow.