A new species of the Eliurus majori complex (Rodentia, Muroidea, Nesomyidae) from south-central Madagascar, with remarks on emergent species groupings in the genus Eliurus

A t american museum

Novitates

PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY
CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024
Number 3547, 21 pp., 7 figures, 2 tables January 15, 2007

A New Species of the Eliurus majori Complex
(Rodentia: Muroidea: Nesomyidae) from
South-central Madagascar, with Remarks on
Emergent Species Groupings in the Genus Eliurus

MICHAEL D. CARLETON 1 AND STEVEN M. GOODMAN 2

ABSTRACT

A new species of Eliurus, E. danieli (Nesomyidae: Nesomyinae), is described from the Parc
National de l’lsalo in south-central Madagascar. Although geographically intermediate to eastern
and western forms, diagnostic traits convincingly relate the new species to E. majori and E.
penicillatus, forms distributed primarily in eastern humid forest; it is morphometrically most
similar to E. penicillatus. In view of the increase in number of Eliurus species (11 now described),
discussion is devoted to emerging species associations (5 are identified) and to taxonomic and
distributional problems introduced by the accumulation of larger series and new localities over the
past decade.

RESUME

Une nouvelle espece d ’’Eliurus, E. danieli (Nesomyidae: Nesomyinae), provenant du Parc
National de l’lsalo, situe au centre-sud de Madagascar est decrite ci-dessous. Geographiquement,
c’est un taxon intermediate entre l’Est et l’Ouest. Les caracteres determinants de cette nouvelle
espece sont similaires a ceux de E. majori et E. penicillatus dont la distribution primaire se situe
dans la foret humide de l’Est. Morphometriquement elle se rapproche plus de E. penicillatus que de

1 Research Associate, Division of Vertebrate Zoology (Mammalogy), American Museum of Natural History; and
Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, DC
20560-0111 (carleton@si.edu).

2 Field Museum of Natural History, 1400 South Lake Shore Drive, Chicago, IL 60605, and World Wide Fund for
Nature, BP 738, Antananarivo (101), Madagascar (sgoodman@fmnh.org, sgoodman@wwf.mg).

Copyright © American Museum of Natural History 2007

ISSN 0003-0082

2

AMERICAN MUSEUM NOVITATES

NO. 3547

E. majori. En vue du nombre croissant des especes d Eliurus (11 actuellement decrites), une
discussion est consacree a l’association des especes emergeant (5 sont identifiees) et aussi sur les
problemes de distribution et taxonomiques engendres par l’accumulation des grandes series de
nouvelles localites durant cette derniere decade.

INTRODUCTION

Most species of Madagascar’s tufted-tailed
rats, genus Eliurus (Nesomyidae: Nesomyinae
sensu Musser and Carleton, 2005), are known
to inhabit the eastern humid forest biome
(Carleton, 1994, 2003; Carleton and Good¬
man, 1998). However, diversity of Eliurus in
the island’s central and western landscapes
has been historically underappreciated, as
evidenced by the recent description of E.
antsingy and discovery of extralimital forms
that demonstrate affinity to eastern species
(Jansa et al., 1999; Carleton et al., 2001). The
taxonomic status and relationships of one of
those outlying forms, from the Parc National
de l’lsalo in south-central Madagascar, are
treated herein. First known from a specimen
collected in April 1995 by an inventory team
led by A. F. A. Hawkins (Hawkins, 1995),
Carleton sketched (26 July 1996) his impres¬
sions as “An Eliurus to be sure but probably
new .... In its robust molars, overall shape
and mandibular conformation, skull sugges¬
tive of the majori-penicillatus complex.”
Tissues of the specimen had been preserved,
and molecular data closely affiliated the
individual (called “sp. A”) with E. majori
Thomas (1895) among the eight Eliurus
species studied (Jansa et al., 1999). In
December 2002, Goodman returned to the
area and obtained two additional specimens.
These exhibit the same unique combination
of morphological traits and sustain descrip¬
tion of the Isalo specimens as a new species.
In addition, we provisionally arrange the 11
currently recognized species of Eliurus into
five species groups, present new range records
for certain species ( E . antsingy Carleton,
Goodman, and Rakotondravony, 2001; E.
myoxinus Milne Edwards, 1885; E. petteri
Carleton, 1994; E. penicillatus Thomas, 1908),
and highlight taxonomic problems that have
emerged coincidentally as sample sizes and
geographic inventory of nesomyine rodents
have markedly improved over the past de¬
cade.

Materials and Methods

Specimens reported herein consist princi¬
pally of study skins and skulls from the
following museum collections: the American
Museum of Natural History, New York City
(AMNH); the Natural History Museum,
London (BMNH, formerly British Museum
of Natural History); Field Museum of Natural
History, Chicago (FMNH); Merseyside
County Museums, Liverpool (LMCM);
Museum of Comparative Zoology, Harvard
University, Cambridge (MCZ); Museum
National d’Histoire Naturelle, Paris (MNHN);
Departement de Biologie Animale, Universite
d’Antananarivo, Antananarivo (UADBA);
National Museum of Natural History,
Smithsonian Institution, Washington, D.C.
(USNM, formerly U.S. National Museum);
Universitets Zoologisk Museum, Copenhagen
(UZMC); and Zoologisches Forschungsinstitut
und Museum Alexander Koenig, Bonn
(ZFMK). Coordinates used to compose the
range map are those given by collectors for
recent collections (see appendix 1) or those
estimated by Carleton and Schmidt (1990) for
older ones.

Six external and 18 craniodental variables
were recorded in millimeters (mm) for each
specimen examined. The external variables, as
extracted from collectors’ skin tags or field
catalogs, include: total length of body and tail
(TOTL); head and body length (HBL); length
of tail vertebrae (TL); hind-foot length (HFL,
usually without claw for recent collections);
ear (pinna) length (EL); and weight in grams
(WT). The 16 cranial and two dental variables
(all measured by Carleton) were recorded to
0.01 mm using handheld digital calipers and
while viewing the skull under a microscope.
According to the anatomical landmarks de¬
fined and illustrated in Carleton (1994), these
include: breadth of the braincase (BBC);
breadth across both incisive foramina (BIF);
breadth of the bony palate across the first
upper molars (BMls); breadth across the
occipital condyles (BOC); breadth of the

2007

CARLETON AND GOODMAN: ELIURUS MAJORI COMPLEX

3

rostrum (BR); breadth of the zygomatic plate
(BZP); depth of the auditory bullae (DAB);
interorbital breadth (IOB); length of bony
palate (LBP); length of diastema (LD); length
of incisive foramen (LIF); coronal length of
maxillary toothrow (LM1-3); length of ros¬
trum (LR); occipitonasal length (ONL); pos¬
terior breadth of the bony palate (PPB);
postpalatal length (PPL); width of the first
upper molar (WM1); and zygomatic breadth
(ZB). Morphological description of the mur-
oid skull generally follows the terms defined
and/or illustrated in Hershkovitz (1962),
Carleton (1980), and Voss (1988).

The new species to be described requires
closest comparison with E. antsingy and
species of the E. majori complex ( E . majori,
E. penicillatus ). The degree of its morphomet¬
ric differentiation was broadly assessed
against samples of these three species as
grouped into the operational taxonomic units
(OTUs) identified below. Catalog numbers,
full locality data, and museum sources of all
specimens analyzed are provided in appendix
1. We use the following abbreviations for the
different types of Malagasy protected areas:
PN, Parc National; RNI, Reserve Naturelle
Integrate; RS, Reserve Speciale.

Eliurus antsingy : (1) PN de Bemaraha, N = 3; (2)
PN (formerly RNI) de Namoroka, N = 10; (3)
RS d’Ankarana, N = 9; (4) Daraina, N = 4.
Eliurus majori : (5) PN de la Montagne d’Ambre, N = 27;
(6) RS de Manongarivo, N = 5; (7) RS

d’Anjanaharibe-Sud, N = 16; (8) PN de Marojejy, N
= 17; (9) RS du Pic d’lvohibe, N = 4; (10) PN
(formerly RNI) d’Andringitra, N = 17; (11) PN
(formerly RNI) d’Andohahela, N = 4.

Eliurus, new species: (12) PN de l’lsalo, N = 4.

Eliurus penicillatus : (13) Ampitambe, N = 17.

Univariate descriptive statistics (mean,
range, and standard deviation) and multivar¬
iate analyses were derived for specimens of
adult age classes (young, full, and old) defined
on grades of dental wear. Juveniles and
subadults, identified by their complete or
partial gray coats and newly erupted or little
worn third molars, were excluded. Sample
statistics for external dimensions are reported
in tabular comparisons as a general index of
size differences among species, but these
variables were not employed in multivariate
analyses. Computation of canonical variates
and principal components was based on the 18

craniodental measurements, all of which had
been first transformed to natural logarithms.
Loadings are expressed as Pearson correlation
coefficients of the extracted factors with logs
of the original skull variables. All analytical
results were obtained using Systat for
Windows (2002, version 10.2).

Eliurus danieli, new species
figures 1-3; table 1

Holotype: Field Museum of Natural
History number 175934, an adult male pre¬
pared as skin, skull, and partial skeleton,
collected 10 December 2002 by Steven M.
Goodman (original number SMG 13,425).

The round skin and skull are in fine
condition. External data recorded on the skin
tag include: TOTL, 335 mm; HBL, 150 mm;
TL, 179 mm; HFL, 30 mm; EL, 26 mm; and
WT, 91 gm. The animal was noted as having
scrotal testes (12 X 5 mm) with convoluted
epididymides; tissues were preserved in EDTA
(ethylene diamine tetraacetic acid).

Type Locality: Madagascar, Fianarantsoa
Province, Parc National de l’lsalo, 28 km SE
Berenty-Betsileo, along Sahanafa River near
foot of Bevato Mountain, 650 m; geographic
coordinates: 22°19.0'S, 45°17.6'E.

Diagnosis: A species of Eliurus character¬
ized in combination by a gray dorsal pelage,
contrasting grayish-white venter, and a distinc¬
tive bright-white tail tuft (fig. 1). Cranial size
moderately large (ONL ca. 37-41 mm; figs. 2,
3), molar rows absolutely and relatively long
(LM1-3 ca. 6.0-6.5 mm), incisive foramina
relatively long and wide (LIF/LD ca. 56%),
and ectotympanic bullae large for the genus.
Alveolus of lower incisor short, not forming
a capsular process.

Referred Specimens: Three additional
specimens are known, all from the PN de
l’lsalo: FMNH 175933, a young adult male
also collected by Goodman (SMG 13,387) in
the same month (5 Dec) from 24 km W
Ranohira bas, Andranohavo (Canyon des
Rats), 700 m, 22°28.9'S, 45°22.9'E; UADBA
10483, a young adult female obtained 2 April
1995, A. F. A. Hawkins (FH 3), from a site
within 300 m of where the holotype was
obtained (Georges Rabeony, personal com-
mun.); and UADBA 46805 obtained between

4

AMERICAN MUSEUM NOVITATES

NO. 3547

Fig. 1. Photograph of live Eliurus danieli (FMNH 175933), a young adult male from Andranohavo
(Canyon des Rats) in the PN de l’lsalo. The new species is characterized by a bright white caudal tuft that is
accentuated by the concentration of intensely black hairs along the middle portion of the tail. Photograph
taken by Harald Schutz.

2 and 9 February 1995 (Hawkins, 1995), by A.
F. A. Hawkins (FH 23) in the Canyon des
Rats.

Distribution: To date known only from
the PN de lTsalo, south-central Madagascar.

Description: Fur texture soft and fine.
Cover hairs 9-10 mm in length over middle
rump. Basal four-fifths of dorsal cover hairs
bright plumbeous gray, distal one-fifth medi¬
um buff, some hairs tipped with a third band
of faint dusky brown. Guard hairs dark brown
to blackish, only slightly longer than the cover
hairs except over the rump. General effect of
upperparts a pale brownish gray; forehead,
face, and lower legs a purer medium gray.
Boundary of dorsal and ventral pelage clearly
demarcated along sides. Ventrum variable in
coverage of white hairs: entirely bright buff
white (UADBA 10483); or dull buff white
over the chin, throat, and chest with some
basal gray thinly showing over the lower

abdomen and inguinum (FMNH 175934); or
wholly white hairs confined to the chin and
throat, cover hairs over the chest and abdo¬
men bicolored (basal half gray, distal half
white), and the underparts overall light
grayish white (FMNH 175933). Medial dusky
brown band extends over the tarsus and
proximal metatarsus; remainder of metatarsus
and toes, as well as tops of forefeet, covered
with bright white hairs; ungual tufts at base
of claws well developed.

Caudal tuft prominent over distal half of
tail length, tuft hairs becoming gradually
longer toward the tip and measuring about
12-15 mm long over the terminal portion
(fig. 1). Terminal one-third of tail length
covered with pure white hairs, forming a bril¬
liant white tuft accentuated by the intensely
black hairs of the proximal tuft (distal white
segment = 35 mm in UADBA 10483, 45 mm
in FMNH 175934, and 75 mm in FMNH

2007

CARLETON AND GOODMAN: ELIURUS MAJORI COMPLEX

5

Fig. 2. Dorsal (upper row) and ventral (lower row) views (ca. 1.75X) of adult crania of species
composing the Eliurus majori species group as recognized herein (see Discussion): left pair, E. penicillatus
Thomas (USNM 49672; Fianarantsoa Province, Ampitambe, ca. 900 m; ONL = 35.6 mm); middle pair, E.
majori Thomas (FMNH 151663; Fianarantsoa Province, PN d’Andringitra, 40 km S Ambalavao, along
Volotsangana River, 1210 m; ONL = 38.1 mm); and right pair, E. danieli new species (FMNH 175934, type
specimen; Fianarantsoa Province, Parc National de l’lsalo, 28 km SE Berenty-Betsileo, along Sahanafa
River near foot of Bevato Mountain, 650 m; ONL = 40.6 mm).

6

AMERICAN MUSEUM NOVITATES

NO. 3547

Fig. 3. Lateral view (ca. 1.75X) of adult crania of species composing the Eliurus majori species group as
recognized herein (same specimens as depicted in fig. 2): top, E. danieli new species; middle, E. majori ; and
bottom, E. penicillatus.

175933). Proximal half of tail clothed with
short black hairs that do not mask the caudal
scales; scalation moderately textured; epider¬
mis pale gray to whitish toward the base of the
tail, lighter underneath, but paleness visually
obscured by the black hairs.

Hind feet short and broad. Six plantar pads
arranged as characteristic of the genus; pads
not so bulbous and crowded as in E. antsingy.
Pinnae medium-sized, more rounded, not so
elongate as E. tanala Major (1896); color pale
dusky brown externally, somewhat lighter
internally, and sparsely clothed with fine
brown hairs inside and outside.

Cranium moderately large (ONL = 37-
41 mm; figs. 2, 3), agreeing with largest geo¬
graphic examples of E. majori (e.g., from PN
de Marojejy), smaller than E. antsingy and
large-bodied E. tanala (e.g., from RS
d’Anjanaharibe-Sud). Dorsal profile of skull
arched (fig. 3), sloping gradually toward the
rostrum and more steeply toward the occiput,

not so flat as in E. tanala of similar size.
Rostrum proportionally long (LR/ONL =
36.2%) for the genus, comparable to relative
length in the E. tanala complex but longer
than E. myoxinus or E. majori. Braincase
rounded and smoothly contoured; subsqua-
mosal fenestra patent and medium-sized,
exposing the interior braincase and dorsal
periotic; hamular process of the squamosal
stout but well delineated. Zygomatic arches
almost parallel-sided in the three specimens;
zygomatic plate medium in width for the
genus, forming a shallow but distinct dorsal
notch, more incised than in E. penicillatus and
E. majori. Incisive foramina bluntly pointed
on their posterior ends, more acute at the
anterior; wide and relatively long (LIF/LD =
56%) for the genus, in proportion resembling
E. antsingy and the E. majori complex (fig. 2).
Posterior palatine foramina oval, situated
within the maxillary-palatine suture at the
level of the Ml-M2; supernumerary palatal

2007

CARLETON AND GOODMAN: ELIURUS MAJORI COMPLEX

7

foramina occurring in the palatine bones, but
minute in size and irregularly formed; slitlike
palatal vacuities absent. Posterior margin of
the bony palate terminates just short of the
ends of the third molars; anterior expanse of
mesopterygoid fossa broad and horseshoe¬
shaped, its sides gently incurved. Alisphenoid
strut present on both sides in all three
individuals examined. Auditory bullae rela¬
tively large for the genus, the ectotympanic
covering nearly all of the periotic in ventral
view, even more so than in E. antsingy and E.
myoxinus. Mandible stocky in form, ascending
ramus deep.

Enamel face of upper and lower incisors
dull yellowish-orange in color. Alveolus of
lower incisor relatively short, terminating
posteriorly at the level of the coronoid process
and well below the sigmoid notch; tip of
incisor root indicated by slight mound on
mandibular ramus but not forming a project¬
ing capsular process. Toothrows long (LM1-3
= 6.0-6.5 mm) relative to cranial size, in
proportion like those of E. majori and E.
penicillatus’, molars robust with laminate
occlusal configuration as per the genus, higher
crowned than most Eliurus but again approx¬
imating E. majori and E. penicillatus in this
detail; upper and lower third molars about
equal in length to second molars or only
slightly shorter.

Other external, cranial, and dental features
as described for the genus (Carleton, 1994;
Carleton and Goodman, 1998).

Morphological and Morphometric Com¬
parisons: The cardinal traits of E. danieli
implicate the eastern species E. majori and E.
penicillatus or western E. antsingy as its closest
relatives. All four are relatively large species of
Eliurus and possess long incisive foramina that
span over half the diastema (fig. 2) and short
lower incisors that lack a prominent capsular
process. In combination, these qualitative
features serve to distinguish the four from all
other species of Eliurus (see Carleton, 1994,
Carleton and Goodman, 1998, and Carleton
et al., 2001, for discussion of interspecific
variation in these characters).

Other character differences associate E.
danieli more closely to E. majori and E.
penicillatus instead of E. antsingy. Foremost,
specimens of E. danieli have long, robust

molar rows in proportion to cranial size
(LM1-3 ca. 16-17% of ONL), a configuration
that agrees with E. majori and E. penicillatus
(Carleton, 1994). In contrast, examples of E.
antsingy have shorter toothrows in relation to
cranial size (LM1-3 ca. 13-14% of ONL),
resembling in this regard all other species of
Eliurus (Carleton et ah, 2001). A simple
bivariate plot underscores the segregation of
the two groups by dental proportion (fig. 4);
only one stray individual (an E. danieli )
overlaps the otherwise clearly delineated
data-point constellations. The morphogenetic
determination of these dental shape contrasts
is such that molar variables (LM1-3, WM1)
always contribute most heavily to group
separation on the first factor extracted in all
ordinations of our cranial measurements,
whether principal component (not illustrated)
or canonical variate (fig. 5, table 2). Eliurus
danieli, together with E. majori and E.
penicillatus, also differ from E. antsingy in
fur texture (not so fine and soft), plantar pads
(not as large, bulbous, and close-set), size of
upper and lower third molars (not smaller
than second molars), and molar hypsodonty
(not as low-crowned). These impressionistic
differences are not so easily quantifiable as is
the disparity in toothrow proportions.

Eliurus danieli can be distinguished from E.
majori and E. penicillatus by means of the
following traits. Although similar in size to E.
majori, specimens of E. danieli possess a con¬
spicuous white caudal tuft. This trait is
unknown in nearly all geographic samples of
E. majori, in which the tuft is wholly dark
brown or black to the tip (Carleton, 1994;
Goodman and Carleton, 1996). In the PN de
Marojejy, however, Carleton and Goodman
(2000) reported that nine of the 20 E. majori
specimens collected possess a white caudal tip
of variable extent. The white tail tip in those
particular E. majori, however, is not accentu¬
ated by a contiguous band of dense black hairs
as found in E. danieli. Furthermore, in other
features and in morphometric analyses, the
Marojejy sample unambiguously accords with
typical E. majori (Carleton and Goodman,
2000; also see next), not E. danieli. Although
possessing a white caudal tuft as diagnostic of
E. penicillatus, specimens of E. danieli are
conspicuously larger in nearly all variables

AMERICAN MUSEUM NOVITATES

NO. 3547

• E. majori + E. danieli

Fig. 4. Scatter plot of length of the upper molar row (LM1-3) versus greatest length of skull (ONL) for
intact specimens of E. antsingy, E. danieli, E. majori , and E. penicillatus. Note that examples of the latter
three species possess longer toothrows in relation to cranial size compared with those of E. antsingy.

measured (figs. 2, 3; table 1); in view of that
great size disparity, the two species are
strongly circumscribed in morphometric space
(fig. 5). Finally, in contrast to E. majori and E.
penicillatus, E. danieli has gray dorsal pelage,
bright in tone (in both eastern species, dorsum
blackish gray to blackish brown and somber
colored); its venter is bright buff white or
grayish white and clearly delineated from the
dorsum (in both eastern species, venter paler
but similarly colored to the dorsum and
dorsal-ventral pelage contrast less well
marked); its ectotympanic bulla is absolutely
larger (table 1), obscuring nearly all of the
periotic in ventral view (in both eastern
species, ectotympanic less inflated, exposing
posteromedial wedge of the periotic); and its
zygomatic plate projects forward to form
a distinct dorsal notch (in both eastern species,
zygomatic plate narrower and dorsal notch
weakly incised).

In projections of the first three canonical
variates (cumulative variation explained =
85.5%) extracted from 13-group discriminant
function analysis, the three specimens from
PN de l’lsalo are wholly isolated in morpho¬
metric space from all other OTUs and species
(fig. 5). As commented above, the two molar
variables (LM1-3, WM1) most strongly in¬
fluence separation of groups along the first
canonical variate (table 2). Other dimensions
that load moderately along this factor include
those that reflect palatal breadth (BMls,
PPW) and interorbital width (IOB); compared
with E. antsingy, the palate is relatively
broader and interorbit narrower in E. danieli,
E. majori, and E. penicillatus (table 1). Size as
a general factor is expressed along the second
canonical variate, as indicated by the positive
and moderately strong correlations (Pearson’s
R = 0.6-0.8, P < .001) of most variables with
this factor (table 2). In general size, specimens

2007

CARLETON AND GOODMAN: ELIURUS MAJORI COMPLEX

9

TABLE 1

Selected External and Craniodental Measurements for the Type Series of E. danieli and Samples of E. majori,

E. penicillatus, and E. antsingy

(Sample statistics include the mean, ± 1 SD, and the observed range)

Variable

E. danieli Isalo
(N = 4)

E. majori Andringitra
(V = 17)

E. penicillatus Ampitambe
(N = 17)

E. antsingy Ankarana
(N = 9)

HBL

150, 152

158.3 ± 7.2

_

141 ± 9.1

145-171

133-153

TL

179, 195

189 ± 11.1

—

166 ± 12.2

170-207

145-183

HFL

30, 32

28.8 ± 1.3

—

28.3 ± 1.4

27-31

25-30

EL

26, 28

19.6 ± 0.7

—

23.7 ± 1.1

18-21

22-25

WT

91, 100

101.0 ± 12.9

—

86.4 ± 9.9

77-122

72-99

ONL

38.9 ± 1.4

37.6 ± 1.0

36.2 ± 0.9

39.3 ± 1.0

37.1-40.6

35.5-39.4

32.9-37.4

37.8-10.5

ZB

18.5 ± 0.9

19.1 ± 0.5

18.0 ± 0.9

19.0 ± 0.7

17.4-19.2

18.1-19.8

15.9-19.4

18.2-20.1

BBC

14.9 ± 0.2

14.3 ± 0.4

13.6 ± 0.4

14.7 ± 0.5

14.6 15.1

13.4-14.8

12.7 14.3

13.9-15.4

IOB

5.4 ± 0.2

5.3 ± .02

5.1 ± 0.2

5.7 ± 0.2

5.3-5.7

4.9-5.7

5.1 5.8

5.3-6.0

LR

14.1 ± 0.8

13.1 ± 0.5

12.4 ± 0.6

13.3 ± 0.4

13.0-14.9

11.6-14.0

10.8-13.2

12.7 13.9

BR

7.1 ± 0.3

7.1 ± 0.2

6.9 ± 0.4

6.9 ± 0.4

6.6-7.5

6.8-7.6

6.1-7.4

6.2-7.4

PPL

13.7 ± 0.9

13.4 ± 0.5

12.9 ± 0.7

14.2 ± 0.5

12.7 14.2

12.4-14.6

11.5-14.2

13.4-15.0

LBP

7.5 ± 0.2

7.5 ± 0.4

6.8 ± 0.3

7.6 ± 0.3

13-1.1

6.9-8.6

6.3-7.4

7.1-8.0

LIF

5.8 ± 0.4

6.3 ± 0.3

5.5 ± 0.3

5.7 ± 0.3

5.4-6.1

5.6-6.7

5.0-5.9

5.3-6.2

BIF

2.7 ± 0.3

2.7 ± 0.2

2.7 ± 0.2

2.6 ± 0.2

2.4-2.9

2.4-3.1

2.4-2.9

2.4-2.9

LD

10.2 ± 0.6

10.8 ± 0.4

9.9 ± 0.6

11.0 ± 0.5

9.6-10.9

10.0-11.8

8.5 10.7

10.2-11.7

BMls

7.6 ± 0.1

8.1 ± 0.3

7.3 ± 0.3

7.4 ± 0.4

13-1.1

7.1-8.6

6.8-7.9

6.9-8.0

DAB

5.7 ± 0.1

5.2 ± 0.1

5.4 ± 0.2

5.3 ± 0.2

5.6-5.9

5.1-5.5

5.0-5.6

5.1 5.6

BZP

3.6 ± 0.5

3.5 ± 0.2

2.9 ± 0.3

3.5 ± 0.3

2.9-3.8

2.9-3.9

2.5-3.4

3.1—4.1

BOC

8.3 ± 0.3

8.5 ± 0.3

8.0 ± 0.3

8.8 ± 0.3

7.9-8.6

8.0-9.0

7.4-8.4

8.2-9.2

LM1-3

6.25 ± 0.20

6.65 ± 0.24

6.02 ±0.15

5.61 ± 0.20

6.00-6.46

6.13-7.15

5.74-6.27

5.24-5.90

WM1

1.66 ± 0.12

1.76 ± 0.07

1.58 ± 0.05

1.53 ± 0.09

1.48-1.74

1.66-1.88

1.49 1.68

1.41-1.65

10

AMERICAN MUSEUM NOVITATES

NO. 3547

• E. majori + E. danieli

A E. penicillatus O E. antsingy

-6 -5 -4 -3 -2 -1 0 1 2 3 4 5 6 7 8 9

CV 1 (52.0%)

Fig. 5. Results of discriminant function analysis performed on 18 log-transformed craniodental
variables, as measured on 130 specimens representing 13 OTUs of Eliurus (see Materials and Methods). Top,
projection of individual scores onto the first and second canonical variates extracted. Bottom, projection of

2007

CARLETON AND GOODMAN: ELIURUS MAJORI COMPLEX

11

TABLE 2

Results of 13-group Discriminant Function Analysis

(OTUs represented include four of E. antsingy, N =
24; one of E. danieli, N = 3; seven of E. majori, N =
90; and one of E. penicillatus, N = 13. See Materials
and Methods and fig. 4)

Correlations

Variable

CV 1

CV 2

CV 3

ONL

0.37

0.75

-0.20

ZB

-0.05

0.72

-0.23

BBC

0.27

0.73

-0.30

IOB

0.58

0.43

-0.01

LR

0.18

0.69

-0.13

BR

-0.23

0.48

0.01

PPL

0.41

0.66

-0.14

LBP

-0.12

0.53

-0.39

LIP

-0.08

0.71

-0.09

BIP

0.24

0.46

0.43

LD

0.15

0.55

-0.39

BMls

-0.48

0.59

0.01

PPB

-0.49

0.60

-0.17

DAB

0.36

0.68

0.38

BZP

0.07

0.66

-0.42

BOC

0.46

0.45

-0.41

LM1-3

-0.82

0.46

0.15

WM1

-0.71

0.44

0.08

Canonical

correlations

0.96

0.90

0.87

Eigenvalue

10.9

4.4

3.0

% Variance

51.1

20.3

14.1

of E. danieli match the larger samples of E.
majori and E. antsingy and conspicuously
exceed the diminutive E. penicillatus in most
dimensions quantified (table 1). Between -
group variation summarized by the third
canonical variate conveys the morphometric
association of E. danieli and E. penicillatus
(fig. 5). The few cranial variables (BIF, LD,
LBP, DAB, BZP) that account for this
association constitute proportional subtleties
that are not readily apparent in side-by-side
skull comparisons of the two species due to
their gross size disparity; notable among these
are the relatively spacious incisive foramina

(BIF) and short diastema (LD). Although
visually unimpressive, such proportional fea¬
tures nevertheless influence the phenetic pair¬
ing of the E. danieli and E. penicillatus samples
among all OTUs represented, as indexed by
Mahalanobis distances between group cen¬
troids (fig. 6). The multivariate level of di¬
vergence between the two (Mahalanobis D 2 =
52.3) surpasses that derived for the most
strongly differentiated samples within either
E. majori or E. antsingy.

Ecological Notes: The PN de l’lsalo is
located on the Isalo Massif, a vast area of
badlands composed of ruiniform Jurassic
sandstone. Much of the park includes rocky
habitat with sparse short scrub or completely
lacking any plants. Many deep canyons found
within this formation contain permanent
rivers that are in part responsible for the
steeply cut valley sides, and the precipitous
facies have a unique rupicolous plant commu¬
nity. Gallery forest can be found along river
margins at the base of these canyons and at
their mouths. An evergreen sclerophyllous
forest formation dominated by the genus
Uapaca (Euphorbiaceae) covers more gently
sloping hillsides. Although the region receives
around 850 mm of rain per year and has
a pronounced dry season (Nicoll and
Langrand, 1989), the water-fed canyons retain
moist conditions and support vegetation
whose floral affinity is transitional between
humid and deciduous formations (Gautier and
Goodman, 2003). Within these moist canyons
occur numerous animals that represent taxa or
species groups otherwise known from the
eastern humid forests.

Microhabitats where specimens of E. danieli
were captured are noted on skin tags or in
original specimen catalogs as follows: “On
rocky cliff about one-half way up Bevato Mt.
in zone with little woody vegetation. Trapped
on ground in small hole leading into rock fall”
(FMNH 175934, holotype); “on the rocky
mountain of Sahanafa” [translated from the

individual scores onto the first and third canonical variates extracted. The maximally inclusive polygons
enclose specimen scores around an OTU’s centroid; in view of extensive overlap and visual confusion, the
sample polygons for E. antsingy (4 OTUs) and E. majori (7 OTUs) were collapsed to enclose maximal
dispersion around the species’ grand centroid. See table 2 for variable correlations and percent variance
explained.

12

AMERICAN MUSEUM NOVITATES

NO. 3547

100

E antsingy

E. major!
Group

E. penicillatus

E. danieli

E, majori

i

50

MahalanobisD 2

o.o

Daraina

Ankarana

Namoroka

Bemaraha

Ampitambe

Isafo

Mt d'Ambre

Andohahela

Marojejy

Manongarivo

Anjanaharibe

Ivohibe

Andringitra

Fig. 6. Phenogram produced from clustering (UPGMA) of Mahalanobis’ distances derived from
discriminant function analysis of 13 OTUs representing four species of Eliurus (see fig. 5).

French] (UADBA 10483); “In canyon with
vegetation at base of rock face and along river
banks—sandy soil. Trap on ground under
opening of large boulder at river edge and
surrounded by ferns and dense rupicolous
vegetation” (FMNH 175933); and “in the
interior of the Canyon des Rats, a site of
exposed rock and at a level more than 30 m
above the water [= river] level” [translated
from the French] (UADBA 46805). This
information suggests that the species is at
least in part terrestrial and lives outside forest
formations.

On the basis of considerable field experience
with members of Eliurus, Goodman has never
captured an individual in an ecological setting
similar to that of the E. danieli holotype. The
site at the foot of Mt. Bevato is a broad
expanse of exposed pseudosteppe, on a steep
rocky slope, and approximately 500 m from
the nearest forest. According to M. George

Rabeony, an employee of the park service who
has accompanied numerous researchers in the
PN de l’lsalo, this is also the locale where
UADBA 10483 was trapped in April 1995. In
late December 2002, Goodman returned to
this site and placed thirty traps (10 Nationals
and 20 Shermans) on the ground under large
rock outcrops between the forest edge and the
middle portion of Mt. Bevato. The individual
(FMNH 175934) was captured in one of the
highest-elevation traps set along the Bevato
cliff face. Other mammals captured in this trap
line include the introduced Rattus rattus and
the large tenrecid Tenrec ecaudatus. A few
individuals of E. myoxinus were trapped
within the Sahanafa Forest, about 1.5 km
from where the holotype of E. danieli was
obtained. In the Andranohavo Forest
(Canyon des Rats), E. myoxinus and R. rattus
were common and captured within a short
distance of E. danieli.

2007

CARLETON AND GOODMAN: ELIURUS MAJORI COMPLEX

13

Taxonomic and Biogeographic Rema¬
rks: The phenetic union of E. danieli and
E. penicillatus and their joint possession of
a white tail tuft, largely to the exclusion of E.
majori, offer a provisional hypothesis of sister-
group relationship. Nonetheless, we are not
persuaded that this cognate relationship will
be necessarily sustained by other data sources,
such as gene-sequence studies. First, with only
four specimens at hand, the sample size of E.
danieli is too small to adequately represent
variational bounds in the species population.
Second, as enumerated above, E. majori and
E. penicillatus do share a number of morpho¬
logical similarities not found in E. danieli. In
view of their distribution in eastern humid
forest, the possibility that E. majori and E.
penicillatus form a clade vicariantly related to
E. danieli in west-central Madagascar remains
as viable an interpretation (and see below).

Compared with other Eliurus that have
a white caudal tuft (E. grandidieri, E. penicil¬
latus, E. tanala), that possessed by E. danieli
is the most visually striking. As we currently
understand interspecific affinities (see Dis¬
cussion) and assuming that an all-dark tuft
represents the primitive state, a terminal white
tuft has evolved at least three times within the
genus. The significance of these contradistinc-
tive caudal flags to social behavior of the
species or in interspecific encounters deserves
investigation.

Although the Isalo Massif today receives
only moderate amounts of rainfall per year
and experiences a pronounced dry season,
organisms identified from regional paleonto¬
logical sites supply ample biotic evidence that
local climatic conditions were distinctly more
mesic during the Late Pleistocene and
Holocene (Burney, 1997; Goodman and
Rakotozafy, 1997; Godfrey et al., 1999).
Relicts from this more mesic phase of late
Quaternary climate persist in the modern flora
of the Isalo area, most notably in the deep
canyons where many plants show phytogeo¬
graphic affinities to the eastern humid forests.
The same relictual biogeographic pattern has
also emerged from the study of certain birds,
reptiles, and amphibians that inhabit the Isalo
Massif (e.g., Raxworthy and Nussbaum, 1996,
1997; Fjeldsa et al., 2000). Two kinds of
taxonomic evidence have been identified

among those vertebrate groups: (1) the same
species is found in common between Isalo
and sites in the eastern humid forests; or
(2) closely related taxa occur in these two
areas and are presumed to have originated
through east-west vicariance. The presence of
E. danieli in the Isalo region and its proposed
sister-group relationship to E. majori and/or
E. penicillatus exemplify the second line of
evidence for a former east-west continuity of
humid forest.

Etymology: We name this new species to
honor Professor Daniel Rakotondravony,
Departement de Biologie Animale, Universite
d’Antananarivo, especially for his contribu¬
tions to our knowledge on Malagasy rodents
and for helping to foster zoological research
on Madagascar for numerous national and
foreign scientists.

DISCUSSION

Carleton based his 1994 revision of Eliurus
on approximately 200 specimens that were
then available in the world’s museum hold¬
ings. With the resurgence of intensive system¬
atic fieldwork over the past decade (see
Andriamialisoa and Langrand, 2003), that
number has climbed to nearly 900 specimens,
a vastly improved empirical base that boasts
both larger locality series and denser geo¬
graphic inventory. The new specimens have
provided a fresh and more rigorous under¬
standing of nongeographic and geographic
variation within Eliurus species, as well as the
discovery of new forms (Carleton and
Goodman, 1998; Carleton et al., 2001; this
paper). Coincidentally, or perhaps inevitably,
the greater sample sizes and proliferation of
intermediate localities have also disclosed
problematic variants that raise questions
about definitions of species limits, distribu¬
tions, and interspecific relationships. We here
review the 11 Eliurus species currently recog¬
nized, highlight populations and geographic
areas of taxonomic uncertainty, and arrange
those discussions around five emergent species
assemblages. These species groups are princi¬
pally based on morphological similarity—
notably as judged by size, cranial and dental
conformation, morphometric cohesion, and
pelage traits. Soft anatomical systems, such as

14

AMERICAN MUSEUM NOVITATES

NO. 3547

the character-rich reproductive tract, have yet
to be comprehensively surveyed. Gene-se¬
quence evidence (cytochrome b ) supplies
phylogenetic perspective that supports the
monophyly of certain of these groups (Jansa
et al., 1999), but not all species nor the
problematic populations discussed below have
yet to be evaluated using such molecular
information.

Eliurus majori Group {E. majori Thomas,
1895; E. penicillatus Thomas, 1908; E.

DANIELI CARLETON AND GOODMAN, 2006): As
reviewed above, these three species constitute
a strongly circumscribed morphologic subset
within the genus. They are moderately large in
size and the hind feet comparatively short. The
tail is well furred, but the caudal hairs
gradually lengthen toward the tip rather than
abruptly elongate as an expansive terminal
tuft. Two of the three species bear a white
caudal tip (E. danieli, E. penicillatus ), but
certain northern populations of the third ( E.
majori ) vary in the incidence of white tipping
(PN de Marojejy-Carleton and Goodman,
2000). Their molar rows are exceptionally
robust with the third molars subequal to the
second, the incisive foramina are wide and
long (> 50% of diastemal length), and the
lower incisor is short and does not form
a protruding capsular process (see Carleton,
1994: fig. 20). Carleton (1994) thought that
E. majori and E. penicillatus retained several
plesiomorphic traits and his cladistic interpre¬
tation represented the two as a basal clade
within Eliurus; however, the molecular results
of Jansa et al. (1999) indicate that, although E.
majori (and E. danieli, reported as “sp. A”)
subtended the E. tanala and E. myoxinus
groups, E. grandidieri-Voalavo composed the
most basal cladistic position among eliurine
Nesomyinae.

The species complex is principally confined
to montane and sclerophyllous montane forest
settings in the eastern humid forest (Carleton,
2003). Size within E. majori is fairly uniform
over much of its geographic range, from the
southern Anosyenne Mountains in the south
to the Northern Highlands, but the northern¬
most population on the isolated peak
Montagne d’Ambre is diminutive. With skulls
arrayed side-by-side, the Montagne d’Ambre
animals “look” like a separate species, but

shape features and genetic data clearly affiliate
this montane outlier with E. majori proper
(Carleton and Goodman, 1998; Jansa et al.,
1999). Extralimital populations have been also
uncovered in remnant montane forests of the
Central High Plateau (Jansa et al., 1999;
Carleton et al., 2001). That from the PN de
l’lsalo is herein diagnosed as a new species
based on recently collected material. Still other
western populations of uncertain status pos¬
sess mq/on'-like features, namely those from
the RS d’Ambohijanahary and RS d’Ambo-
hitantely (appendix 1); these specimens have
an all-dark tail tuft and approximate the
Montagne d’Ambre sample of E. majori in
their smaller size.

Carleton (1994) doubtfully retained E.
penicillatus as distinct from E. majori, but
the accumulation of larger samples of the
latter species, now available over a broad
north-south geographic swath, continues to
underscore the clear-cut morphometric sepa¬
ration of the two (Carleton and Goodman,
2000; this study). Jansa et al.’s (1999)
cytochrome-/? genetic evidence divulged close
relationship of the Isalo species, E. danieli
(called “sp. A”), and E. majori, whereas our
morphometric evaluation identified E. penicil¬
latus as an alternative possibility of cognate
affinity. Jansa et al. (1999) lacked tissues for
E. penicillatus, which until recently has been
known only from the type locality,
Ampitambe, and all specimens collected by
Forsyth Major in 1895-1896 (see Jenkins and
Carleton, 2005). A lone specimen (UADBA
16209) collected in 2000 from Ankerana,
about 35 km NE of Fandriana, represents
only the second locality that we consider
definitely referable to E. penicillatus, but it
supplies evidence that the species still survives.
Ampitambe is about 45 km SE Fandriana (see
Carleton and Schmidt, 1990, for discussion).
Perhaps the distribution of E. penicillatus, like
E. petteri, is notably localized along the
eastern flank of the Central High Plateau.
Renewed search for the species and preserva¬
tion of fresh material to investigate its status
and relationship with regard to E. majori and
E. danieli are warranted.

Eliurus ant sin gy Group ( E . ant sin gy
Carleton, Goodman, and Rakotondravony,
2001): Eliurus antsingy is the most recent of

2007

CARLETON AND GOODMAN: ELIURUS MAJORI COMPLEX

15

the morphological themes within Eliurus to be
formally recognized. A large species with
a dark tail tuft and extremely bulbous plantar
pads, E. antsingy possesses some character¬
istics that intimate relationship with the E.
majori group: namely, small and rounded
pinnae; a short lower incisor that terminates
in an indistinct incisive capsule; and long and
wide incisive foramina. However, unlike
species of the E. majori group, its molars
are decidedly less robust in relation to cranial
size (fig. 4), and the upper and lower third
molars are smaller than the second; according
to these features, E. antsingy resembles other
species of Eliurus. Until evaluated with other
data sources, we isolate E. antsingy in its own
group.

The distribution of E. antsingy is highly
disjunct, so far known from three widely
separate regions (fig. 7). As conveyed by the
specific epithet, the first examples identified,
from the PN de Bemaraha and PN de
Namoroka, appeared to be restricted to
limestone tsingy physiography and associated
forest habitat. Recent collecting has extended
the range northward, including other sites in
tsingy in the RS d’Ankarana but also near
massive metamorphic rock outcrops at the
edge of transitional dry-humid forest in the
Foret d’Ambilondambo, near Daraina (see
appendix 1 for detailed localities). Collecting
sites with known elevation are all relatively
low, 50-550 m. The animals from the PN de
Namoroka, as noted by Carleton et al.
(2001), are slightly smaller and possess all-
white venters compared with those from the
PN de Bemaraha. In ventral pelage color,
those from Ankarana and Daraina recall the
series from PN de Namoroka but are again
smaller by an increment comparable to that
between the Bemaraha and Namoroka series.
The overall amount of craniodental differen¬
tiation among the three regions approximates
that derived among samples of the broadly
distributed E. majori (fig. 6), as the latter is
currently understood, and they marginally
overlap in morphometric space (not illustrat¬
ed). We continue to recognize these regional
samples as the one species E. antsingy , but
stress the need for collections from interme¬
diate localities, wherever potential habitat
exists, and for the application of genetic data

to assess levels of divergence and phylogeo-
graphic structure.

Eliurus myoxinus Group ( E. myoxinus
Milne Edwards, 1885; E. minor Major,
1896): Affiliation of these two species in
a single species group at first seems incongru¬
ous given that each was conventionally
accepted as one of the only two valid species
of Eliurus (Ellerman, 1949; Petter 1972, 1975;
Honacki et al., 1982; Corbet and Hill, 1991).
In light of current research, the polytypic
construct of E. myoxinus that so long masked
the distinctive morphologies and genetic di¬
vergence of forms such as E. majori and E.
tanala almost defies comprehension. Unen¬
cumbered by identification with these species
and others, the morphological similarity of E.
myoxinus and E. minor is more readily
appreciable: size small ( E. minor ) to medium
( E. myoxinus ); skull stockily built with a rela¬
tively broad interorbit, short rostrum, and
incisive foramina medium in length; dorsal
pelage color dominated by brown tones;
caudal pilosity dense and tuft long, covering
one-half to two-thirds of the tail length.
Although no special affinity was appreciated
by Carleton (1994), Jansa et al. (1999)
presented molecular data that lend confidence
in the sister-group relationship of E. myoxinus
and E. minor.

Morphological resemblance of the two
species is more convergent among their
northernmost populations. Carleton (1994)
drew attention to the large size of E. minor
that inhabit lowlands around Antongil Bay;
robust proportions also characterize those
obtained from the RS de Manongarivo. Size
variation within E. myoxinus, on the other
hand, trends from larger in the south (Petriky
Forest) to smaller in the northwest (PN de
l’Ankarafantsika), as the species range was
then known to Carleton et al. (2001). Recent
surveys have now documented E. myoxinus
even farther to the north, from the RS de
Manongarivo, Daraina Forest, and the north¬
western sector of the PN de Marojejy in
Antsiranana Province (Goodman and Soari-
malala, 2002; Soarimalala and Goodman,
2003; appendix 1). These northern samples
continue the pattern of size decline, represent¬
ing the smallest specimens of the species we
have so far examined, but they generally

16

AMERICAN MUSEUM NOVITATES

NO. 3547

Antsiranana

a Efiurus antsingy
• Eliurus danieli
o Efiurus majori
n Efiurus penicillatus

Mahajanga

Toliara

i Toamasina

Fianarantsoa ■

Altitude (m)

□ 0-1000
1000-2000
2000 - 2876

O

2007

CARLETON AND GOODMAN: ELIURUS MAJORI COMPLEX

17

conform to the cranial proportions and
external morphology that typify E. myoxinus
populations to the south (unpublished data
and analyses). Sample sizes and geographic
representation of both forms have vastly
improved since Carleton’s (1994) revision
and should enable a critical and much-needed
evaluation of the complex. Careful scrutiny
should be devoted to northern Madagascar
where the ranges of E. myoxinus and E. minor
appear to interdigitate in a complex manner,
unlike the southern facies of their distributions
where biotic transition is dramatically abrupt
and their morphological differentiation is just
as sharply marked (Goodman et al., 1999).

Eliurus petteri Group (E. petteri Car-
leton, 1994; E. grandidieri Carleton and
Goodman, 1998): These two species are
medium in size, possess soft-textured and
short pelage, and have small auditory bullae,
delicately built upper and lower incisors, and
an extremely short lower incisor without
a capsular projection. Their tails are relatively
long and the tail tuft very short and weakly
developed in both (ca. distal one-fourth of tail
length), one with a white-tip ( E. grandidieri)
and the other dark ( E. petteri ). Some of these
conditions are the extreme expression of the
character as so far observed within the genus
(see Carleton and Goodman, 1998: 173).
Genetic investigation to date is limited to E.
grandidieri, which surprisingly is revealed as
most closely allied to Voalavo gymnocaudus
according to these data (Jansa et al., 1999).
These results puzzle us in view of the morpho¬
logical traits that characterize Voalavo and
unambiguously distinguish it from all species of
Eliurus (Carleton and Goodman, 1998: 189—
194). Further, Voalavo possesses a relatively
large midventral gland, unknown in any species
of Eliurus, that produces a distinctly musky
odor. Although the two genus-group taxa are
undoubtedly close kin (Carleton and
Goodman, 1998; Jansa and Weksler, 2004),
the possible paraphyly of Eliurus as indicated
earlier by Jansa et al. (1999) requires further

corroboration and its generic diagnosis and
contents amended if appropriate. Inclusion of
tissues from E. petteri, along with additional
geographic samples of E. grandidieri, may
provide a fresh taxonomic scope to do so.

The range of E. grandidieri is more extensive
than that of E. petteri, the former known in
the Northern Highlands and northern Central
High Plateau, within montane and upper
montane forest (ca. 1200-1900 m), and the
latter restricted to a small area in east-central
Madagascar, at intermediate elevations (ca.
400-1100 m) within Toamasina Province
(Carleton, 2003). Recent recoveries of E.
petteri —from Andriantantely, Saharay, and
Sandranantitra (appendix 1)—originated from
the same localized area and reaffirm the
species’ distinctive morphology.

Eliurus tan ala Group (E. tan ala Major,
1896; E. webbi Ellerman, 1949; E. ellermani
Carleton, 1994): These are large species, E.
ellermani and some northern samples of E.
tanala being fully as robust as E. antsingy, but
in cranial and dental proportions, the three
differ strikingly from that species and from
those of the E. majori group. The molars are
shorter and narrower relative to cranial size,
with the third molars smaller than the second;
upper and lower incisors are stout, and the
lower is long and terminates as a moderately
(E. webbi ) to strongly (E. tanala, E. ellermani )
pronounced capsular process on the upper
mandibular ramus. The lunate incisive foram¬
ina are medium in length for the genus,
supernumerary palatal foramina are common¬
place, often coalesced as elongate vacuities, and
the subsquamosal fenestra is small (most E.
webbi) or absent (E. tanala, E. ellermani). The
pinnae are pliant and elongate, averaging the
longest in the genus, and the caudal tuft is well
developed but confined to the terminal 30-40%
of tail length. Carleton (1994) speculated that
E. webbi is sister species to the clade E. tanala-
E. ellermani, an expectation not sustained by
the few cladistic characters he mustered. Jansa
et al. (1999), using cytochrome b sequences,

Fig. 7. Geographic occurrence of the new species E. danieli, known only from PN de l’Isalo, and its
eastern relatives E. majori and E. penicillatus. Also illustrated is the currently documented range of E.
antsingy based on new records reported herein.

18

AMERICAN MUSEUM NOVITATES

NO. 3547

demonstrated phylogenetic pairing of E. webbi
and E. tanala among the eight species studied.

The E. tanala group presents many taxonom¬
ic problems that require attention. For one, the
status of E. ellermani as a species distinct from
E. tanala merits reexamination. Application of
molecular techniques to fresh material from the
amended type locality of E. ellermani (Carleton
and Goodman, 1998), along with examples of
the large, creamy white-bellied E. tanala from
the Northern Highlands, should provide the
pivotal evidence.

As in the E. majori group, western outliers
of the E. tanala assemblage have been recently
documented (Carleton et al., 2001; appendix
1). Compared with eastern E. tanala proper,
those from the PN de Bemaraha have
a distinctive color pattern—notably the pale
brown dorsum, entirely creamy-white under¬
parts, and the alternately banded tail tuft—
but in cranial shape and their robust size, they
suggest E. ellermani and northern E. tanala.
That from the RS de Maningoza is similar in
size and body color but possesses a monoco¬
lored tail tuft. Such chromatic contrasts,
coupled with the wide geographic hiatus
between the western tanala- like populations
and the eastern range of E. tanala proper, cast
doubt on their conspecific stature. The taxo¬
nomic status and distributional extent of the
western morphs invite critical review.

Morphological variability within both E.
tanala and E. webbi increases in the northern
segments of their ranges and complicates their
identification (Carleton and Goodman, 1998,
2000). Variability is particularly evident in the
invasiveness of all creamy-white hairs on the
ventrum, the constancy of tail tuft coloration,
closure of the subsquamosal fenestra, and
overall size. Across the central and southern
parts of their distributions, such features
appear more stable in their population expres¬
sion and meaningfully contribute to specific
discrimination (Carleton, 1994). The mixture
of traits exhibited by certain northern samples,
such as those from the Daraina region
(appendix 1) and perhaps the RS de
Manongarivo (Goodman and Soarimalala,
2002—listed as E. webbi), is so atypical that
specific allocation is uncertain. These speci¬
mens may constitute a different species alto¬
gether or represent exceptional morphological

convergence of E. webbi toward the E. tanala
morphotype. Teasing apart these hypotheses
will require denser transects within the trou¬
blesome geographic region in question, broad
distributional sampling of typical E. tanala
and E. webbi against which to gauge the
differentiation of the northern samples, and
application of multiple information sources.

In summary, we can be confident that
Eliurus comprises many more than the two
species so long recognized as valid by
Ellerman (1949) and others. Nevertheless, the
genus obviously requires further field, muse¬
um, and laboratory investigation to improve
its alpha-level systematics. Some of the re¬
search challenges outlined above will un¬
doubtedly reveal new species, but of greater
systematic interest, the next phase of Eliurus
investigation should aim to refine relation¬
ships among species and to explore patterns of
their biogeographic origination.

ACKNOWLEDGMENTS

For collection access and/or specimen loans,
we very much appreciate the helpfulness of the
following museum staff: G. G. Musser
(AMNH); P. Jenkins (BMNH); J. Phelps and
W. Stanley (FMNH); M. J. Largen (LMCM);
M. Rutzmoser (MCZ); M. Tranier (MNHN);
C. Smeenk (RMNH); D. Rakotondravony,
V. Soarimalala, and M. Raheriarisena
(UADBA); H. Baggoe and M. Anderson
(UZMC); and R. Hutterer (ZFMK). Field
surveys in the Isalo area were financed by
a grant to SMG from the Volkswagen
Foundation, and Georges Rabeony consider¬
ably helped SMG with the Isalo survey.
Permission to conduct the fieldwork was
granted by the Association Nationale pour la
Gestion des Aires Protegees and the Direction
des Eaux et Forets. We are most grateful to
Harald Schiitz for making available the
photograph of the living animal shown in
Figure 1; John Steiner (NMNH Branch,
Smithsonian Photographic Services), who
photographed the skulls and composed
Figures 2 and 3; and Lucienne Wilme for
producing the map in Figure 7. Lastly, we
appreciate the time devoted and comments
offered by Guy G. Musser and two anony¬
mous reviewers.

2007

CARLETON AND GOODMAN: ELIURUS MAJORI COMPLEX

19

REFERENCES

Andriamialisoa, F., and O. Langrand. 2003.
History of scientific exploration. In S.M.
Goodman and J.P. Benstead (editors), The
natural history of Madagascar: 1-15. Chicago:
University of Chicago Press, xxi + 1709 pp.

Burney, D.A. 1997. Theories and facts regarding
Holocene environmental change before and
after human colonization. In S.M. Goodman
and B.D. Patterson (editors), Natural and
human-induced change in Madagascar: 75-89.
Washington, D. C.: Smithsonian Institution
Press, xiii + 432 pp.

Carleton, M.D. 1980. Phylogenetic relationships in
neotomine-peromyscine rodents (Muroidea)
and a reappraisal of the dichotomy within
New World Cricetinae. University of Michigan
Miscellaneous Publications Museum of
Zoology 157: 1-146.

Carleton, M.D. 1994. Systematic studies of
Madagascar’s endemic rodents (Muroidea:
Nesomyinae): revision of the genus Eliurus.
American Museum Novitates 3087: 1-55.

Carleton, M.D. 2003. Eliurus, tufted-tailed rats. In
S.M. Goodman and J.P. Benstead (editors),
The natural history of Madagascar: 1373-1380.
Chicago: University of Chicago Press, xxi +
1709 pp.

Carleton, M.D., and S.M. Goodman. 1998. New
taxa of nesomyine rodents (Muroidea:
Muridae) from Madagascar’s northern high¬
lands, with taxonomic comments on previously
described forms. Fieldiana: Zoology, new series
90: 163-200.

Carleton, M.D., and S.M. Goodman. 2000.
Rodents of the Parc National de Marojejy,
Madagascar. Fieldiana: Zoology, new series
97: 231-263.

Carleton, M.D., S.M. Goodman, and D.
Rakotondravony. 2001. A new species of
tufted-tailed rat, genus Eliurus (Muridae:
Nesomyinae), from western Madagascar, with
notes on the distribution of E. myoxinus.
Proceedings of the Biological Society of
Washington 114: 972-987.

Carleton, M.D., and D.F. Schmidt. 1990. Sys¬
tematic studies of Madagascar’s endemic ro¬
dents (Muroidea: Nesomyinae): an annotated
gazetteer of collecting localities of known
forms. American Museum Novitates 2987:
1-36.

Corbet, G.B., and J.E. Hill. 1991. A world list of
mammalian species, 3 rd ed. London: Natual
History Publications, vii + 243 pp.

Ellerman, J.A. 1949. The families and genera of
living rodents. Volume 3, Appendix II [Notes
on the rodents from Madagascar in the British

Museum, and on a collection from the island
obtained by Mr. C. S. Webb]. London: British
Museum (Natural History), v + 210 pp.
Fjeldsa, J., S.M. Goodman, T.S. Schulenberg, and
B. Silkas. 2000. Molecular evidence for rela¬
tionships of Malagasy birds. In N.J. Adams
and R.H. Slotow (editors), Proceedings of the
22nd International Ornithological Congress:
3084-3094. Durban: University of Natal.
Gautier, L., and S.M. Goodman. 2003. Intro¬
duction to the flora of Madagascar. In S.M.
Goodman and J.P. Benstead (editors), The
natural history of Madagascar: 229-250.
Chicago: University of Chicago Press, xxi +
1709 pp.

Godfrey, L.R., W.L. Jungers, E.L. Simons, P.S.
Chatrath, and B. Rakotosamimanana. 1999.
Past and present distributions of lemurs in
Madagascar. In B. Rakotosamimanana, H.
Rasamimanana, J.U. Ganzhorn and S.M.
Goodman (editors), New directions in lemur
studies: 19-53. New York: Kluwer Academic/
Plenum, x + 295 pp.

Goodman, S.M., and M.D. Carleton. 1996. The
rodents of the Reserve Naturelle Integrale
d’Andringitra, Madagascar. Fieldiana: Zoo¬
logy, new series 85: 257-283.

Goodman, S.M., and L.M.A. Rakotozafy. 1997.
Subfossil birds from coastal sites in western and
southwestern Madagascar: A paleoenviron-
mental reconstruction. In S.M. Goodman and
B.D. Patterson (editors), Natural and human-
induced change in Madagascar: 257-279.
Washington, D C: Smithsonian Institution
Press, xiii + 432 pp.

Goodman, S.M., and V. Soarimalala. 2002. Les
petits mammiferes de la Reserve Speciale
de Manongarivo, Madagascar. Boissiera 59:
383-401.

Goodman, S.M., M.D. Carleton, and M. Pidgeon.
1999. The rodents of the Reserve Naturelle
Integrale d’Andohahela, Madagascar. Fieldi¬
ana: Zoology, new series 94: 217-249.
Hawkins, A.F.A. 1995. Isalo faunal inventory.
Final report to the Association Nationale pour
la Gestion des Aires Protegees. Unpublished
report, 58 pp.

Hershkovitz, P. 1962. Evolution of Neotropical
cricetine rodents (Muridae) with special refer¬
ence to the phyllotine group. Fieldiana:
Zoology 46: 1-524.

Honacki, J.H., K.E. Kinman, and J.W. Koeppl.
1982. Mammalian species of the world.
Lawrence, KS: Association of Systematic
Collections, ix + 694 pp.

Jansa, S.A., S.M. Goodman, and P.K. Tucker.
1999. Molecular phylogeny and biogeography
of the native rodents of Madagascar (Muridae:

20

AMERICAN MUSEUM NOVITATES

NO. 3547

Nesomyinae): a test of the single-origin hy¬
pothesis. Cladistics 15: 253-270.

Jansa, S.A., and M. Weksler. 2004. Phylogeny of
muroid rodents: relationships within and
among major lineages as determined by IRBP
gene sequences. Molecular Phylogenetics and
Evolution 31: 256-276.

Jenkins, P.D., and M.D. Carleton. 2005. Charles
Immanuel Forsyth Major’s expedition to
Madagascar, 1894 to 1896: beginnings of
modern systematic study of the island’s mam¬
malian fauna. Journal of Natural History
39(20): 1779-1818.

Major, C.I.F. 1896. Descriptions of four additional
new mammals from Madagascar. Annals and
Magazine of Natural History, series 6, 17:
461^163.

Milne Edwards, A. 1885. Description d’une nou-
velle espece de rongeur provenant de
Madagascar. Annales des Sciences Naturelles,
Zoologie et Paleontologie 20: Article 1 bis.

Musser, G.G., and M.D. Carleton. 2005.
Superfamily Muroidea. In D.E. Wilson and
D.M. Reeder (editors), Mammal species of the
World, a taxonomic and geographic reference,
3 rd ed: 894-1531. Baltimore: Johns Hopkins
University Press, Volume 2: xvii +745-2142 pp.

Nicoll, M.E., and O. Langrand. 1989. Madagascar:
revue de la conservation et des aires protegees.
Gland: WWF, xvii + 374 pp.

Petter, F. 1972. The rodents of Madagascar: the
seven genera of Malagasy rodents. In R.
Battistini and G. Richard-Vincard (editors),
Biogeography and ecology in Madagascar:
661-666. The Hague: Dr. W. Junk B. V.
Publishers, xv + 765 pp.

Petter, F. 1975. Family Cricetidae: Subfamily
Nesomyinae. In J. Meester and H.W. Setzer
(editors), The mammals of Africa, an identifi¬

APPENDIX 1

Specimens Examined of Eliurus

Listed below are specimens that formed the basis
for the study’s morphological comparisons, sample
statistics, morphometric analyses, mapped distribu¬
tions, and range extensions. Abbreviations for
protected areas in Madagascar are: PN, Parc
National; RNI, Reserve Naturelle Integrate; RS,
Reserve Speciale.

Eliurus aff. E. majori Group: Mahajanga
Province: RS d’Ambohijanahary, Foret d’Ankazo-

cation manual, Part 6.2: 1^1. Washington, DC:
Smithsonian Institution Press, [not continuous¬
ly paginated, parts issued separately]

Raxworthy, C.J., and R.A. Nussbaum. 1996.
Pattern of endemism for terrestrial vertebrates
in eastern Madagascar. In W.R. Lourengo
(editor), Biogeographie de Madagascar:
369-383. Paris: ORSTOM, x + 588 pp.

Raxworthy, C.J., and R.A. Nussbaum. 1997.
Biogeographie patterns of reptiles in eastern
Madagascar. In S.M. Goodman and B.D.
Patterson (editors), Natural change and human
impact in Madagascar: 124-141. Washington,
D. C: Smithsonian Institution Press, xiii +
432 pp.

Soarimalala, V., and S.M. Goodman. 2003.
Diversite biologique des micromammiferes
non volants (Lipotyphla et Rodentia) dans le
complexe Marojejy-Anjanaharibe-Sud. In S.M.
Goodman and L. Wilme (editors), Nouveaux
resultats d’inventaires biologiques faisant
reference a l’altitude dans la region des massifs
montagneux de Marojejy et d’Anjanaharibe-
Sud. Antananarivo: Centre d’lnformation et de
Documentation Scientifique et Technique,
Recherches pour le Developpement, Serie
Sciences biologiques 19: 231-278.

Thomas, O. 1895. On a new species of Eliurus.
Annals and Magazine of Natural History,
series 6, 26: 164-165.

Thomas, O. 1908. A new species of the Mascarene
genus Eliurus. Annals and Magazine of Natural
History, series 8, 2: 453^154.

Voss, R.S. 1988. Systematics and ecology of
ichthyomyine rodents (Muroidea): patterns of
morphological evolution in a small adaptive
radiation. Bulletin of the American Museum of
Natural History 188: 259^193.

tsihitafototra, 1150 m, 18°15.7'S, 45°25.2'E (FMNH
167546,167555). Antananarivo Province: RS d’Ambo-
hitantely, 24 km NE Ankazobe, Jardin Botanique,
1450 m, 18°10.1', S, 47°16.6'E (FMNH 165505,
176019-176022).

Eliurus aff. E. tan ala Group: Antsiranana
Province: Foret de Binara, 7.5 km SW Daraina,
near Analamazava River, 325-550 m, 13°15.3'S,
49°37.0'E (FMNH 172665-172675). Mahajanga
Province: RS de Maningoza (FMNH 167562).
Mahajanga Province: Bekopaka (MNHN
1966.221); PN de Bemaraha, 2.5 km NE
Bekopaka, Ankidrodroa, 100 m, 19°07.9'S,
44°48.5'E (FMNH 172722); PN de Bemaraha,

2007

CARLETON AND GOODMAN: ELIURUS MAJORI COMPLEX

21

3.5 km E Bekopaka, near Tombeau Vazimba,
100 m, 19°08.4'S, 44°49.7'E (FMNH 172723-
172725).

Eliurus antsingy: Antsiranana Province: Foret
d’Ambilondambo, 5 km N Daraina, Andrafia, 300-
550 m, 13°09.9'S, 49°38.9'E (UADBA 46062, 46072,
46073); Foret de Bobankora, 11 km ESE Daraina,
Beantely, 350-550 m, 13°13.4'S, 49°45.6'E (UADBA
46101); RS d’Ankarana, 2.6 km E Andrafiabe, forest
near Andrafiabe Cave, 50 m, 12°55.9'S, 49°03.4'E
(FMNH 169718-169720); RS d’Ankarana, 7.5 km
NW Mahamasina, Anilotra, 125 m, 12°54.5'S,
49°06.6'E (FMNH 173104-173106, 173109); RS
d’Ankarana, 10 km NW Mahamasina, 100 m,
12°53.2'S, 49°06.6'E (FMNH 173107, 173108).
Mahajanga Province: RNI [now PN] de Namoroka
(FMNH 167563-167566; UADBA 16169); RNI
[now PN] de Namoroka, 26 km NW Andra-
nomavo, Foret d’Ambovonomby, 200 m, 16°28.2'S,
45°20.9'E (FMNH 175909-175911); RNI [now PN]
de Namoroka, 31 km NW Andranomavo, Foret de
Mahabo, 100 m, 16°23.4'S, 45°20.9'E (FMNH
175912, 175913). Mahajanga Province: Antsingy
forest near Bekopaka, ca. 19°07.5'S, 44°49.0'E
(MNHN 1966.2220, 1966.2222); PN de Bemaraha,

3.5 km E Bekopaka, near Tombeau Vazimba,
100 m, 19°08.4'S, 44°49.7'E (FMNH 172721).

Eliurus majori: Antsiranana Province: Montagne
d’Ambre, 1000 m (AMNH 100687, 100854); PN de
Montagne d’Ambre, 12 km SW Joffreville, Grand
Lac, 1325 m, 12°35.8'S, 49°09.6'E (FMNH 154345);
PN de Montagne d’Ambre, 5.5 km SW Joffreville,
1000 m, 12°31.6'S, 49°10.3'E (FMNH 154536-
154539, 154603-154616); RS de Manongarivo,
17.3 km SW Antanambao, 1600 m, 14°01.3'S,
48°25.1'E (FMNH 166211-166213); PN de
Marojejy, 11km NW Manantenina, 1325 m,
14°26.2'S, 49°44.5'E (FMNH 159624, 159709); PN
de Marojejy, 10.5 km NW Manantenina, Antra-
nohofa, 1625 m, 14°26.4'S, 49°44.5'E (FMNH
159625-159637, 159710-159715, UA-SMG 8463);
PN de Marojejy, 11 km NW Manantenina, 1875 m,
14°26.8'S, 49°44.1'E (FMNH 159638); RS

d’Anjanaharibe-Sud, 9.2 km WSW Befingotra,
1260 m, 14°44.7'S, 49°27.7'E (FMNH 154052,
154241-154243, 154258, 154266, UA-SMG 6981,
6998); RS d’Anjanaharibe-Sud, 11km WSW
Befingotra, 1550 m, 14°44.5'S, 49°26.0'E (FMNH
154053, 154244); RS d’Anjanaharibe-Sud, 12.2 km

WSW Befingotra, 1950 m, 14°44.8'S, 49°26.0'E
(FMNH 154054, 154245, 154298; UA-SMG 7090).
Mahajanga Province: Western slope Anjanaharibe-
Sud, 13 km SW Befingotra, 1600 m, 14°45.9'S,
49°25.9'E (FMNH 167452-167457). Antananarivo
Province: 16.2 km SE Tsinjoarivo, Andrindizimbolo
River, Foret d’Ankilahila, 1400 m, 19°42.4'S,
47°50.1'E (FMNH 166162, 166613). Fianarantsoa
Province: Ambohimitombo (BMNH 97.9.1.147;
MCZ 45929); RNI [now PN] d’Andringitra, 38 km
S Ambalavao, ridge east of Volotsangana River,
1625 m, 22°11'39"S, 46°58'16"E (FMNH 151666,
151667, 151732, 151752, 151851-151858); RNI
[now PN] d’Andringitra, 40 km S Ambalavao, along
Volotsangana River, 1210 m, 22°13'22"S, 46°58T8"E
(FMNH 151661-151665, 151730, 151731, 151847-
151849); Andringitra Massif, Anjavidilava, 2030 m
(MNHN 1972.602); RS du Pic d’lvohibe, 1575 m
(FMNH 161891, 162073-162075). Toliara Province:
RNI [now PN] d’Andohahela, Parcel 1,13.5 km NW
Eminiminy, 1200 m, 24°35.0'S, 46°44.1'E (FMNH
156503, 156615); RNI [now PN] d’Andohahela,
Parcel 1, 15.0 km NW E mi ni mi ny, 1500 m,

24°34.2'S, 46°43.9'E (FMNH 156616, 156617,
156658).

Eliurus myoxinus : Antsiranana Province: Foret
de Binara, 7.5 km SW Daraina, near Analamazava
River, 325-550 m, 13°15.3'S, 49°37.0'E (FMNH
172655-172664); RS de Manongarivo, 12.8 km SW
Antanambao, 785 m, 13°58.6'S, 48°25.4'E (FMNH
166252); RS de Manongarivo, 14.5 km SW
Antanambao, 1240 m, 14°00.0'S, 48°25.7'E (FMNH
166253); PN de Marojejy, along Bemanavy River,

11.5 km SE Doany, 810 m, 14°25.6'S, 49°36.5'E
(FMNH 172606-172614, 173207, 173209-173214,
173216, 173220-173224).

Eliurus penicillatus : Fianarantsoa Province:
Ampitambe (BMNH 97.9.1.148-97.9.1.150, 97.9.1.152,
1939.1892; FMNH 18822; LMCM A19.4.98.26;
MCZ 12435, 45932; MNHN 1897.536, 1909.191;
USNM 49672; UZMC 1219, 1224, 7941; ZFMK
82.217, 83.56); Prefecture de Fandriana, Ankerana,
1670 m, 20°03'49"S, 47°40T8"E (UADBA 16209).

Eliurus petteri: Toamasina Province: Andrian-
tantely, 530 m (FMNH 166062, 166063); Saharay,
1000-1145 m, 18°41.7'S, 48°35.24'E (UADBA
19139); Sandranantitra, 450 m (FMNH 16604).

Complete lists of all issues of the Novitates and the Bulletin are available at World Wide
Web site http://library.amnh.org/pubs. Inquire about ordering printed copies via e-mail from
scipubs@amnh.org or via standard mail from: American Museum of Natural History,
Library—Scientific Publications, Central Park West at 79th St., New York, NY 10024. TEL:
(212) 769-5545. FAX: (212) 769-5009.

@ This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).