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Mammalia 74 (2010): 177–185 2010 by Walter de Gruyter • Berlin • New York. DOI 10.1515/MAMM.2010.018
2010/201004
Article in press - uncorrected proof
Species limits and distribution of the Malagasy carnivoran
genus Eupleres (Family Eupleridae)
Steven M. Goodman
1,2,
* and Kristofer M. Helgen
3
1
Field Museum of Natural History, 1400 South Lake Shore
Drive, Chicago, IL 60605, USA
2
Vahatra, BP 3972, Antananarivo 101, Madagascar,
e-mail: sgoodman@fieldmuseum.org;
sgoodman@vahatra.mg
3
National Museum of Natural History, Smithsonian
Institution, P.O. Box 37012, MRC 108, Washington,
DC 20013-7012, USA
*Corresponding author
Abstract
A review was conducted of members of the endemic Mala-
gasy carnivoran genus Eupleres (family Eupleridae) based
on published and unpublished records and museum speci-
mens. Classically, one species has been recognized in this
poorly known genus – E. goudotii, divided into two geo-
graphical forms with non-overlapping distributions: E. g.
goudotii distributed in the mesic forests of the east and E. g.
major found in the dry areas of the northwest. Drawing on
external and craniodental comparisons, we demonstrate that
these two forms are highly distinctive morphologically and
can be readily distinguished from each another. Furthermore,
there is some evidence that they both can occur on the slopes
of Montagne d’Ambre in the far north of the island. On this
basis, we recognize these taxa as distinct species, E. goudotii
and E. major.
Keywords: distribution; Eupleres; Madagascar;
morphology; taxonomy.
Introduction
Until recently it was assumed that the native Carnivora of
Madagascar represented several deeply divergent lineages
within the order, and hence that they arrived on the island
via multiple colonization events (Gregory and Hellman 1939,
Simpson 1945). However, recent molecular comparisons
have shown that all native Malagasy Carnivora genera
(Cryptoprocta, Fossa, Eupleres, Galidia, Galidictis, Sala-
noia, and Mungotictis) form a monophyletic group repre-
senting a single colonization event (Yoder et al. 2003) and
all are now placed in an endemic family, Eupleridae (Wozen-
craft 2005). This family represents one of the more extra-
ordinary adaptive radiations among the world’s Carnivora
(Goodman 2009). Although higher level systematics of the
Eupleridae are becoming reasonably well established (Yoder
et al. 2003), the distribution and species limits of several
genera remain poorly studied. Herein we address these
aspects for the enigmatic genus Eupleres (the falanouc),
which possesses a highly reduced dentition and other
remarkable anatomical features (Gray 1870, Carlsson 1902,
Pocock 1915, Scapino 1981, Popowics 2003, Gaubert et al.
2005) and for which little information is available regarding
natural history and geographical distribution (Albignac 1972,
1973a, 1974).
In his extensive review of the Carnivora of Madagascar,
Albignac (1973a) considered Eupleres to be monospecific
and represented by two subspecies: E. g. goudotii Doye`re,
1835, from the eastern wet forests and E. g. major Lavauden,
1929, from the transitional dry deciduous-humid forests of
the northwest. Albignac (1973a) presented several external
morphological characters to separate these two subspecies.
In subsequent taxonomic reviews of the Carnivora of
Madagascar, this monospecific arrangement for Eupleres has
been followed (Wozencraft 1993, 2005), although there has
been some indication that these geographical forms could
warrant specific recognition (Lavauden 1929, Albignac
1973b). The purpose of this paper is to review morphological
characters and measurements distinguishing these two forms,
their geographical distribution, and to assess their specific
status.
Methods and materials
Patterns of morphological variation
To assess patterns of morphological variation in the different
recognized forms of Eupleres we examined and measured
specimens in the following institutions: American Museum
of Natural History (AMNH); Muse´um national d’Histoire
naturelle (MNHN); Museum of Comparative Zoology, Har-
vard University (MCZ); Natural History Museum, London
wBMNH; formerly The British Museum (Natural History)x;
Yale Peabody Museum (YPM), Yale University; Museum fu¨r
Naturkunde, Humboldt Universita¨t, Berlin (ZMB). Some of
these specimens are incomplete, missing teeth or damaged
in other ways. All craniodental measurements included here-
in were taken by the second author, at an accuracy of 0.1 mm
for cranial variables and 0.01 mm for dental variables.
The following measurements were taken for all adult (and
nearly mature subadult) specimens – skulls: greatest length
of skull (GSL), condylobasal length (CBL), zygomatic
breadth (ZB), breadth of braincase at zygomatic roots (BB),
interorbital width (IOW), length of anterior palatal foramina
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178 S.M. Goodman and K.M. Helgen: Species limits of Eupleres (Eupleridae)
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Table 1 External measurements (in mm) of Eupleres taken from
Albignac (1973a) and from linear measurements carefully estimated
directly from study skins at MCZ by the second author.
Form Head and Tail Hind Ear
body length length foot length length
Eupleres goudotii
Albignac 455–495 220–240 80–82 40–44
MCZ (ns4) 470–525 200–250
Eupleres major
Albignac 515–650 240–250 81–92 47–50
MCZ (ns4) 610–740 210–310
(APFL), maximum length of nasal bones (NL), palatal length
(PL), auditory bulla length (AUDL), auditory bulla width
(AUDW), and mandible length (ML); teeth – length of upper
first molar (LM1); width of upper first molar (WM1); length
of upper second molar (LM2); width of upper second molar
(WM2); external alveolar distance between upper first
molars (M1–M1); external alveolar distance between upper
canines (C–C); length of maxillary toothrow (C–M2); width
of lower first molar (wm1); length of second lower molar
(lm2); length of third lower premolar (lp3).
Results
Morphology
External morphology
External measurements tabulated
both by Albignac (1973a) from non-cited sources and from
museum study skins at MCZ (Table 1) demonstrate the con-
sistently larger body size of major relative to goudotii.
Another striking distinction between major and goudotii is
in pelage coloration; major is distinctly darker brown than
goudotii, which is paler and more fawn brown in overall
coloration (both species are paler ventrally than dorsally).
Albignac (1973a) noted sexual dimorphism in both of these
forms, with males being slightly larger than females, but
based on current limited sample sizes we are unable to test
for these differences in a meaningful way.
Another consistent difference between these two forms is
in the position and size of the foot-pads and fur cover on the
distal portions of the fore and hind feet (see Albignac 1973a,
Figures 9–12). On the palmar surface of the forelimb of
nominate goudotii the digital pads are only moderately
developed, and the interdigital and metacarpal pads are large-
ly fused and form a large central surface, whereas in major
the digital pads are markedly larger and inflated, and the
hypothenar and thenar pads are notably separated from the
inflated interdigital pads (terminology follows Brown and
Yalden 1973). Furthermore, in major the palmar surface pos-
terior to the metacarpal pads is largely naked and furred in
goudotii. On the plantar surface of goudotii, the digital pads
are reduced, and the largely fused interdigital pads form a
central rounded unit with a notable separation from the dig-
ital pads, whereas in major the digital pads are conspicuously
inflated structures, the fused interdigital pads form a large
proportion of the foot-pad surface, and the metatarsal pads
(thenar and hypothenar) are more developed. Finally, a con-
siderable portion of the plantar surface posterior to the
metatarsal pads is naked in major and largely covered with
fur in goudotii.
Cranial measurements and morphology There are
several cranial and mandibular measurements that distinguish
goudotii and major (Table 2, Figure 1). In general, major has
a distinctly more robust skull, both in cranial length (e.g.,
greatest length of skull) and breadth (e.g., zygomatic
breadth), as well as a longer and more robust mandible. In
fact, there is a large discrepancy, with no overlap, in condy-
lobasal length between adults of goudotii (F87 mm) and
major (G90 mm), offering a convenient single metric for
discriminating adult skulls. Furthermore, major has a dis-
tinctly longer palate, but shorter anterior palatal foramina
than those of goudotii, an important cranioproportional dis-
tinction. One of the most noticeable diagnostic differences
between these two animals is the much more expansive audi-
tory bullae in major, as reflected both in average values for
length and width dimensions (tabulated in Table 2), but most
notably expressed by inflation of the bullae in the vertical
(dorsoventral) dimension (difficult to measure consistently
and not tabulated here).
Dental measurements Many of the specimens available
for this study are missing teeth, so that there are numerous
dental variables for which few measurements are available,
particularly for major. In general, the upper molar teeth in
major are more robust in length (e.g., length first upper
molar, length second upper molar) and width (e.g., width first
upper molar, width second upper molar) than in goudotii.
The length of the maxillary toothrow in major is notably
longer than in goudotii. Furthermore, the distance across the
rostrum (e.g., distance outer C–C) and palate (distance outer
M1–M1) are greater in major compared with goudotii. The
lower molar measurements in goudotii show little overlap in
length and width to those measurements of major (e.g., sec-
ond to last lower width, second to last lower width, length
of fourth to last tooth in lower jaw) (Table 3). Hence, animals
assigned to nominate goudotii and major have conspicuously
different dentitions, which in major are mainly more robust.
Taxonomic conclusions
Given the notable differences between the two Eupleres taxa
in body size (as shown by external and skull measurements),
pelage coloration, foot-pad morphology and flexor fur pat-
terns, cranial-mandibular proportions and measurements, and
dental measurements, as well as differences in their associ-
ated habitats, and some evidence (see below) that these taxa
can occur in sympatrically (or in near sympatry), we suggest
that these geographical forms should be elevated to the rank
of species. Specimens of Eupleres are rare in museum col-
lections, and to our knowledge no frozen tissue samples are
available of both species to test easily this recommendation
with molecular genetic data. Studies drawing on genetic sam-
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Table 2 Cranial and mandibular measurements (in mm) of Eupleres specimens based on age classes.
Form and GSL CBL ZB BB IOW APFL NL PL AUDL AUDW ML
age class
E. goudotii
Older subadult 88.5, 89.2 84.0, 84.9 35.2, 35.4 31.7, 32.2 14.9, 15.3 6.3, 6.6 21.0, 23.5 41.2, 43.1 13.2, 15.8 8.8, 9.4 60.8, 61.6
Adult 88.5
"
2.51 84.0
"
2.13 35.3
"
1.61 32.0
"
1.01 14.3
"
0.79 6.3
"
0.51 21.6
"
1.45 42.5
"
2.12 14.9
"
0.82 9.3
"
0.56 61.5
"
2.26
83.8–92.6 79.8–86.5 33.0–38.4 30.6–33.8 13.1–15.3 5.3–6.8 19.8–25.1 39.4–45.8 14.1–16.5 8.8–10.5 58.7–64.7
ns12 ns12 ns10 ns11 ns8ns7ns12 ns8ns9ns8ns7
E. major
Older subadult 92.1 89.8 35.9 32.2 14.5 5.7 25.6 45.6 15.2 10.2 65.0
Young adult 96.0 93.4 35.8 31.8 14.3 6.0 23.8 48.5 17.4 10.0 67.6
Adult 92.9, 97.0 91.3, 92.1 38.3, 39.4 32.7, 33.5 14.9, 17.0 4.7, 5.7 23.9, 26.5 46.5, 46.7 15.2, 16.0 10.0, 11.6 67.2, 67.7
For definitions of age classes and measurement acronyms see materials and methods section. Sample sizes of three or more specimens are presented as mean
"
standard deviation, mini-
mum–maximum measurements, and number of specimens.
ples from specimens collected in the future or utilizing tech-
niques for DNA extraction from museum skins and skulls
(Helgen et al. 2008, 2009) are recommended to examine
patterns of geographical variation between and within both
Eupleres taxa; to test our hypothesis that E. goudotii and E.
major each represent relatively deeply divergent evolutionary
lineages; and to revisit the phylogenetic relationship of
Eupleres with other Malagasy carnivoran genera (Yoder et
al. 2003).
Geographical distribution
We present 40 records of Eupleres spp. from different
regions of Madagascar based on a detailed survey of pub-
lished and unpublished literature, as well as collections in
different museums. Given the considerable amount of field-
work and inventories conducted on the island over the past
decades, it is clear that members of this nocturnal–
crepuscular genus are notably rare or at least reclusive in
their habits.
On the basis of specimen and sight records, Eupleres has
a much broader distribution than previously understood
(Albignac 1973a). Eupleres goudotii was previously known
from the northern half of the eastern humid forest (Albignac
1973a, Figure 5), but its distribution extends the complete
length of the eastern portion of the island to Andohahela
(Figure 2, Appendix 1). The westernmost known record of
this species in northern Madagascar is the montane forest of
Montagne d’Ambre. Its documented elevation distribution is
from approximately 50 to 1200 m. Eupleres major was con-
sidered by Albignac (1973a) to be restricted to the Sambi-
rano region, i.e., the transitional humid-dry deciduous forests
of northwestern Madagascar (Gautier and Goodman 2008).
In fact, it also has a broader distribution in the west, reaching
the middle portion of the central lowland zone, at least to
the Parc National de Baie de la Baly, near Soalala (Figure
2). It has been documented across an elevational range from
approximately 10 to 1500 m.
The northern portion of the island, within the Antsiranana
Province, is a zone where the two Eupleres replace one
another. Eupleres major lives in the transitional northwestern
forests and associated marshlands east to the foothills of the
Tsaratanana Massif (based on specimens with well-docu-
mented collection data). However, there is a specimen of E.
major in the MCZ (45963) with the locality ‘‘Diego Suarez’’
wsAntsirananax and collected by ‘‘M. Drouhard, inspecteur
du Eaux et Foreˆts de Mad. nov. 1934’’. Grandidier (1934)
described two species of shrew tenrec (Microgale, family
Tenrecidae) collected by M. Drouhard from ‘‘des environs
de Diego-Suarez’’. As with the Eupleres specimen under dis-
cussion, these type specimens are also part of the Grandidier
Collection now housed in the MCZ (Helgen and McFadden
2001). On the basis of extensive small mammal inventories
in the general vicinity of Antsiranana in both dry and humid
forests (Raxworthy and Nussbaum 1994, Goodman et al.
1996, 1997, S.M. Goodman unpublished data), it can be con-
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180 S.M. Goodman and K.M. Helgen: Species limits of Eupleres (Eupleridae)
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Figure 1 Views of skulls and mandibles of Eupleres spp. from Madagascar: left – E. goudotii (MCZ 45959) from the Foreˆt Sihanaka,
right – E. major (MCZ 45963) from ‘‘Diego Suarez’’. Views include (from top to bottom) dorsal view of cranium, ventral view of cranium,
lateral view of cranium, lateral view of mandible, and dorsal view of mandible. Notable differences between these two taxa include the
shape of the cranium and rostrum, as well as the size of the auditory bullae.
Table 3 Dental measurements (in mm) of Eupleres specimens based on age classes.
LM1 WM1 LM2 WM2 M1–M1 C–C C–M2 Wm1 Lm2 Lp3
E. goudotii
Older 3.81, 4.23 3.61, 3.85 2.80, 3.18 3.20, 3.76 17.62, 18.77 8.20, 8.51 33.1, 32.0 1.92, 2.02 5.50, 4.83 3.20, 3.56
subadult
Adult 3.91
"
0.13 3.52
"
0.26 2.62
"
0.06 3.28
"
0.18 16.52
"
0.61 7.84
"
0.30 32.1
"
1.15 1.80
"
0.12 5.13
"
0.35 2.92
"
0.26
3.75-4.11 3.25–3.96 2.56–2.74 2.98–3.50 15.31–17.15 7.46–8.25 30.7–33.9 1.64–1.97 4.85–5.87 2.54–3.32
ns7ns6ns7ns7ns7ns8ns8ns6ns7ns7
E. major
Adult 4.2
"
0.28 3.8
"
0.07 2.6, 3.6 3.6, 3.7 18.3, 19.3 8.6
"
0.38 35.6
"
0.89 2.0
"
0.03 5.9
"
0.20 3.4
"
0.18
3.9–4.5 3.8–3.9 8.2–8.9 34.8–36.5 2.0–2.1 5.7–6.1 3.3–3.6
ns3ns3ns3ns3ns3ns3ns3
Some examined specimens are missing teeth or are damaged. See Materials and methods section for definitions of measurement acronyms.
Sample sizes of three or more specimens are presented as mean
"
standard deviation, minimum–maximum measurements, and number of
specimens.
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Figure 2 Map of the known distribution of Eupleres. The details for each numbered site can be found in Appendix 1. Sites with well-
documented records are partially colored in black and those with vague or questionable records are completely in white. Other localities
mentioned in the text are also shown on the map.
cluded that the type specimens of these two shrew tenrecs
were almost certainly obtained in the humid forests on the
upper slopes of Montagne d’Ambre (Jenkins et al. 1997).
Hence, we presume that the E. major specimen in the Gran-
didier collection (MCZ 45963) could have come from the
same zone of Montagne d’Ambre. Given this supposition,
both E. goudotii and E. major could occur on the slopes of
this massif, although, direct sympatry cannot be confirmed.
The lower slopes of this massif have dry deciduous forest
and at around 900 m there is an ecotone to humid forest.
Eupleres goudotii is known from this massif at 1000 m
(Appendix 1; Sites 13 and 14). If indeed the Drouhard spec-
imen of E. major was obtained on Montagne d’Ambre, it
might have been obtained at a lower elevation in less mesic
forest habitat. Clarifying the nature of geographical and eco-
logical overlap or abutment between E. goudotii and E.
major at the interface of their distributions remains an excit-
ing priority for further research.
Discussion
It was previously thought that the two Eupleres taxa, here
recognized as distinctive species, have parochial habits with
regard to habitat requirements, with E. goudotii being found
in dense humid forests and associated marshlands and E.
major in the Sambirano region, with its transitional humid
and deciduous forests and marshlands (Albignac 1973a).
Based on the distributional records tabulated here, we find
that these characterizations do not always hold. For example,
the habitat associated with the southernmost record of E.
goudotii, along the Mandrare River (Site 40), is notably dif-
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182 S.M. Goodman and K.M. Helgen: Species limits of Eupleres (Eupleridae)
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ferent from that of the majority of other sites where this
species has been recorded from, being a mixture of spiny
bush and gallery forest that receives less than 500 mm of
annual precipitation and experiences a notable dry season
(Chaperon et al. 1993). One of the principal branches of the
Mandrare River has its origin in the Anosyenne Mountains,
a zone to the east that receives at least 1500 mm of rain per
year and which experiences a less pronounced dry season.
Hence, the marshlands associated with gallery forest of the
middle section of this river basin would remain mesic
throughout the year. In the Foreˆt d’Andampy (Site 12), at
the northern end of the distribution of this species, the natural
forest habitat would be closer to a dry deciduous type, but
with more mesic conditions than the Mandrare River basin;
the nearby town of Vohemar receiving over 1600 mm on
average of annual rainfall (Chaperon et al. 1993).
Across the known range of Eupleres major there is also a
notable cline in annual rainfall and length of the dry season.
This ranges from notably moist montane forest at 1500 m
on the slopes of the Tsaratanana Massif (Site 4), where the
syntypes of E. major were collected, to the largely dry decid-
uous forest at the Baie de Baly (Site 11) with annual rainfall
of slightly more than 1000 mm per year and a notable dry
season (Chaperon et al. 1993). Within the southern portion
of range of E. major, there are considerable marshlands,
which could be the mainstay of this species.
Members of the enigmatic genus Eupleres, which show
remarkable adaptive and ecological parallels to the African
aardwolf Proteles, a termite specialist (Matsebula et al.
2009), and remains one of the most poorly known carnivo-
rans of Madagascar. Herein, using morphological data, we
provide some details on the species limits of Eupleres, divid-
ing the former monospecific genus into two separate species,
which have broader distributions and more ecologically
diverse habitat requirements than previously realized. Further
fieldwork to collect tissue samples for molecular phylogeo-
graphical studies should provide further insight into the
patterns of speciation in this genus. On the basis of site
observations and specimen records, E. major appears to have
the most limited distribution of the two species; it is currently
only known from five specimens (all in the MCZ). Eupleres
goudotii sensu lato is considered by the IUCN as Near
Threatened (Hawkins 2008). With the taxonomic changes
proposed here associated with splitting E. goudotii into two
separate species, both the level of molecular divergence
between them and their conservation status will need to be
reassessed.
Acknowledgements
We are grateful to the following curators and collection managers
for allowing access to specimens in their care: Robert Asher, Judith
Chupasko, Christiane Denys, Paula Jenkins, Darrin Lunde, Eileen
Westwig, Robert Voss, Eric Sargis, and Kristof Zyskowski. We also
thank Jean-Louis Camicas, Luke Dollar, Scott Cardiff, Lucienne
Wilme´ for information on distributional records, Lauren Helgen for
photography and other assistance with our manuscript, Lucienne
Wilme´ for producing Figure 2, and two anonymous reviewers for
comment on an earlier version of the paper.
Appendix 1
Localities from which Eupleres goudotii and E. major have
been collected or reported. The individual sites have separate
numbers, which are plotted in Figure 2, and are presented
here by province (north to south), with supporting informa-
tion and associated comments. Abbreviations for protected
areas are: PNsParc National, RNIsRe´serve Naturelle Inte´-
grale, RSsRe´serve Spe´ciale. In a few cases, supplementary
information is presented on unacceptable records.
Eupleres major
Province d’Antsiranana
Dollar (2000) noted this genus in the RS d’Ankarana; this
record is incorrect and associated with a printing error (L.
Dollar personal communication).
Site 1 In the MCZ (45963) there is a specimen collected
near ‘‘Diego Suarez’’ (sAntsiranana). This might well have
been taken from the slopes of Montagne d’Ambre. A note
on the specimen label has inscribed on it ‘‘don de M. Drou-
hard, inspecteur du Eaux et Foreˆts de Mad. nov. 1934’’.
There are other cases of Drouhard specimens bearing the
same collection locality and are probably from Montagne
d’Ambre (see text).
Site 2 Albignac (1973a) mapped a record of this species
approximately halfway between Ambilobe and Ambanja
(estimated at 138279S, 488409E), but no precise detail is
presented.
Site 3 Seven individuals were obtained from the region of
Ambanja (Albignac 1973a, p. 85); the exact sites are not
explicitly stated and this region is estimated to be located at
138419S, 488279E.
Site 4 Several authors have noted that this species might
occur in the RS de Manongarivo (central point is 138599S,
488229E) (Schreiber et al. 1989, Dollar 2000, Conservation
Breeding Specialist Group 2002). We are unaware of any
definitive proof of its occurrence within this reserve despite
several mammal expeditions to the site (Raxworthy and
Rakotondraparany 1988, Goodman and Soarimalala 2002).
Site 5 The syntypes of E. g. major (MCZ 45961, 45962)
were described by Lavauden (1929) based on specimens in
the Grandidier collection that were subsequently transferred
to the MCZ (Helgen and McFadden 2001, Helgen 2002).
The material was collected north of Beangona, in the Upper
Sambirano River Valley and at the foot of the Tsaratanana
Massif at a site estimated to be 148019S, 488479E, 1500 m.
Province de Mahajanga
Site 6
There is a report by Schwitzer (2005) of this animal
on the Sahamalaza Peninsula (central position of the asso-
ciated forest block is at 148069S, 478429E).
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Site 7 Albignac (1973a) noted that an individual was
found in the region of Port Berge´(sBoriziny), estimated to
be near 158339S, 478409E.
Site 8 Kaudern (1915) observed an individual at Andra-
nolava (approximately at 158409S, 488089E, 200 m).
Site 9 An individual was found by Kaudern (1915) in a
zone between Ste. Marie and Marovoay (approximately
168079S, 468439E).
Site 10 There are reports from Ste. Marie de Marovoay
(Kaudern 1915) in an area estimated to be located at 168089S,
468589E, 50 m. Albignac (1973a, Figure 5) incorrectly
reported this record from I
ˆ
le Ste. Marie, an eastern near shore
island, based on Kaudern (1915).
Site 11 An animal in the MCZ (45964) was obtained 115
SE of Mahajanga (estimated to be 168239S, 468569E, 50 m),
which would be in the region near Maevatanana.
Site 12 An individual of Eupleres was observed in degrad-
ed deciduous forest in the PN de la Baie de Baly by Hawkins
(1994) at 168049S, 458169E, 10 m.
Eupleres goudotii
Province d’Antsiranana
Site 13
There are numerous observations and photographs
of this animal in close vicinity of the Station Forestie`re des
Roussettes in the PN de la Montagne d’Ambre
(12831937.30S, 49810919.10E, 1000 m) (Nicoll and Langrand
1989, Projet ZICOMA 1999, Dollar 2000, Goodman 2009).
Site 14 A specimen (AMNH 100461) was collected at
Montagne d’Ambre, probably in close proximity to Site 13
(estimated at 128319S, 498109E, 1000 m).
Site 15 In the MNHN (1912.129) there is a fluid preserved
specimen collected by G. Grandidier in the ‘‘Foreˆt d’Andam-
py, coˆte NE de Madag’’. The inferred coordinates for this
locality are 138309S, 498519E, approximately 100 m.
Site 16 Safford and Duckworth (1990) noted that local
guides living close to the RNI de Marojejy (now a PN)
reported a Eupleres from near Manantenina (148299S,
498499E, approximately 350 m).
Site 17 There are numerous reports of Eupleres on the
Masoala Peninsula, which is a PN (Schreiber et al. 1989, R.
Albignac personal communication). A portion of this reserve
falls within the Province de Toamasina.
Province de Toamasina
Site 18
Golden (2009) has stated that people living in the
Makira Forest (approximately centered at 158149S, 498319E)
hunt Eupleres for bushmeat, although details of the specific
sites were not cited.
Site 19 A specimen in the ZMB (44572) and another in
the AMNH (188211) were collected at Ambatond’Radama
(sAmbatondradama or Col de Radama) by H. Blu¨ntschli
(158179S, 508049E, 600 m).
Site 20 Two skin specimens in the BMNH (1932.3542,
35.1.8.310) were obtained by Rand at a site 40 km NW of
Maroantsetra, near the village of Bevato and at approxi-
mately 158179S, 498349E, between 400 and 500 m (L. Wilme´
personal communication).
Site 21 A specimen in the Grandidier Collection (MCZ
45965) was obtained at Fahampanambo, which is located at
158229S, 498389E and at approximately 100 m.
Site 22 A specimen was collected in 1878 (RMNH 34449)
in the vicinity of Mahalevona at approximately 158269S,
498579E and at 1100 m (L. Wilme´ personal communication),
in probably lowland areas.
Site 23 Rand collected two individuals (AMNH 100484,
100462), 20 km SW of Maroantsetra estimated to be
158329S, 498379E, approximately 50 m. The site was along
the coastal plain and in close proximity to the village of
Manombia (Rand 1936).
Site 24 A specimen (RMNH 34450) was collected in 1876
by J. Audebert near Mananara (estimated locality at 168239S,
498449E). A number of modern reports of this taxon have
come from the PN de Mananara-Nord, presumably in the
same general area (Albignac 1973a, Nicoll and Langrand
1989, Projet ZICOMA 1999).
Site 25 A considerable number of specimens have been
collected in the Sihanaka (sSianaka) Forest (BMNH
1932.3540, 1932.3541, 35.1.8.308, 35.1.8.30935.12.10.2,
1962.16021962.2105; FMNH 30492; MCZ 27830, 45957,
45959; ZMB 43127), a region to the east of Lac Alaotra.
Most, if not all, of these specimens were obtained by Hers-
hell Chauvin, who collected many vertebrate specimens in
the region. Numerous birds obtained by Chauvin are labeled
‘‘foreˆt de Sihanaka’’ and have secondary locality informa-
tion making reference to the villages of Didy (18879S,
488339E) and Fito (188059S, 488549E). Hence, the Sihanaka
Forest (sensu Chauvin) probably falls within the region
between these two villages.
Site 26 Hoogstraal and Camicas (1977) noted that speci-
mens of the tick Haemaphysalis eupleres (described by
Hoogstraal et al. 1965) were collected at Manakambahiny
Est, Andranomalaza (estimated coordinates at 178389S
0488389E) and within the RNI de Zahamena. It is presumed
that the ectoparasites were obtained from an individual of
Eupleres, but this detail cannot be confirmed (Jean-Louis
Camicas, personal communication).
Site 27 There are observations of this taxon from within
the RNI de Zahamena at Volotsanganana (178429S, 488469E,
850–1000 m) (Anonymous 1995).
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184 S.M. Goodman and K.M. Helgen: Species limits of Eupleres (Eupleridae)
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Site 28 Anonymous (1995) observed this species from
within the RNI de Zahamena at Namarafana (178449S,
48858.59E, 400–500 m).
Site 29 A specimen (BMNH 71.4.3.8) was collected by
C.S. Webb at ‘‘12 mls NW Lohariandava’’, which has been
interpreted as being at 188429S, 488359E, 450 m (Carleton
and Schmidt 1990).
Site 30 The holotype of E. goudotii was obtained by M.
Goudot from local people ‘‘de Tamatave’’, who indicated
that this animal is found in areas with sand, where they dig
burrows (Doye`re 1835). The precise location of the collec-
tion site is unknown and is placed in close proximity to the
city of Toamasina (sTamatave) at 188099S, 498239E.
Site 31 There is a specimen in the MCZ (45958) collected
near Rogez (sAmbatovola) (estimated at 188489S, 488379E,
450 m).
Site 32 The Conservation Breeding Specialist Group
(2002) lists this species for the PN de Mantadia (centroid at
188509S, 488289E, 930–1040 m); the basis for this record is
not known.
Site 33 There are numerous reports from the Pe´rinet area
(Albignac 1973a, Nicoll and Langrand 1989, Dollar 2000),
which are presumably within or in close proximity to the RS
d’Analamazaotra (188569S, 488269E, approximately 900 m).
A specimen was collected in the region of Pe´rinet on 6 May
1972 by R. Albignac (Hoogstraal and Camicas 1977).
Site 34 Albignac (1973a) plotted a record just to the south
of Pe´rinet and on the southern side of Route National 2,
linking Antananarivo to Toamasina. Based on the charted
point, the site could be close to or within the Maromizaha
Forest (188579S, 488279E, 1000–1200 m).
Province de Fianarantsoa
Site 35
Observed at Vatoharanana (218169S, 478269E,
approximately 1125 m) within the PN de Ranomafana (Dol-
lar 1999).
Site 36 Observed at Talatakely (218149S, 478269E, approx-
imately 850 m) within the PN de Ranomafana (Dollar 1999).
Site 37 An individual was observed in the RNI d’Andrin-
gitra (now a PN) (228139400S, 478009130E, 810 m) (Good-
man 1996).
Site 38 There is an undocumented report by Nicoll and
Langrand (1989) of this species in the PN de Befotaka-
Midongy du Sud (238459S, 478079E).
Province de Toliara
Site 39
There is one record of Eupleres in parcel 1 of the
RNI d’Andohahela (now a PN) (46844.19S, 24835.09 E,
1200 m) (Goodman and Pidgeon 1999), as well as other
reports from this reserve pre-dating this record (Nicoll and
Langrand 1989, Durban in Hawkins 1994).
Site 40 A specimen in the MCZ (45737) was obtained at
Ambovombe (Androy) by Raymond Decary (estimated site
details are 258109S, 468059E, 135 m). Decary (1950, p. 39)
provides some further details on this record; he notes that
this species occurs within the Androy, specifically the middle
portion of the Mandrare River.
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