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Family Sciuridae G. Fischer, 1817

January 2015

January 2015

DOI:10.7208/chicago/9780226169606.001.0001

In book: Mammals of South America Volume 2, Rodents (pp.1-48)
Chapter: Family Sciuridae G. Fischer, 1817
Publisher: University of Chicago Press
Editors: James L. Patton, Ulyses F.J. Pardiñas, Guillermo D'Elía

Authors:

Mario de Vivo

University of São Paulo

Ana Paula Carmignotto

Universidade Federal de São Carlos

A taxonomic review of the South American squirrels of the Family Sciuridae, Subfamily Sciurinae

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3′. Infraorbital foramen positioned on side of rostrum an-

terior to zygomatic plate; cheek teeth number four, ei-

ther hypsodont, cylindrical, and planar (Geomyidae) or

brachydont, bunodont, and cuspidate (Heteromyidae)

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Castorimorpha

Suborder Sciuromorpha Brandt, 1855

Infraorder Sciurida Carus, 1868

Family Sciuridae G. Fischer, 1817

Mario de Vivo and Ana Paula Carmignotto

Squirrels are ubiquitous inhabitants of all forest biomes

throughout South America except the temperate forests,

with all species adapted for life in the canopy, even though

some forage on the ground. Specialized adaptations for

climbing include elongate bodies, forefeet with four long

toes with claws, hindfeet with ﬁ ve long toes with claws,

large plantar pads on all feet, broad heads, large eyes, short

ears, generally soft fur, long bushy tails, and elongated legs.

The ankle joint is ﬂ exible, allowing the hindfoot to rotate

sufﬁ ciently for headﬁ rst descent of vertical tree trunks. Most

species have enlarged, long incisors accompanied by corre-

spondingly large jaw muscles, which enable them to gnaw

through the hardest nuts. Some species specialize on hard-

shelled seeds of tropical trees such as palms, but most are

more omnivorous, feeding on a range of food types such

as nuts, fruits, insects, fungi, and sometimes leaves, ﬂ ow-

ers, and bark (Emmons and Feer 1997). All South Ameri-

can squirrels are diurnal; as such, individuals can be easily

Class Mammalia Linnaeus, 1758

Order Rodentia Bowdich, 1821

new world rodents

James L. Patton

KEY TO THE SUBORDERS OF RECENT SOUTH

AMERICAN RODENTS:

1. Well- deﬁ ned zygomatic plate anterior to zygomatic arch

present; infraorbital foramen small and either piercing

zygomatic plate or positioned on side of rostrum anterior

to zygomatic plate; lower jaw sciurognathous, with ori-

gin of angular pro cess directly ventral to sheath of lower

incisor. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

1′. No zygomatic plate; infraorbital foramen greatly en-

larged; lower jaw hystricognathous, with the root of

the angular pro cess deﬂ ected lateral to sheath of lower

incisor. . . . . . . . . . . . . . . . . . . . . . . . . Hystricomorpha

2. Zygomatic plate oriented anteriorly; infraorbital fora-

men small, transmitting only nerves and blood vessels;

maxillary teeth number four or ﬁ ve . . . . . . . . . . . . . . 3

2′. Zygomatic plate oriented more laterally; infraorbital

foramen enlarged, narrow, but typically expanded dor-

sally, transmitting medial masseter muscle in addition to

nerves and blood vessels; maxillary teeth number three

or fewer . . . . . . . . . . . . . . . . . . . . . . . . . . Myomorpha

3. Infraorbital foramen pierces zygomatic plate; cheek

teeth number four or ﬁ ve, with molars brachydont, bu-

nodont, with three transversely oriented crests . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sciuromorpha

Taxonomic Accounts

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2 Mammals of South America

basal offshoot that diverged very early in the history of the

family. Although Moore (1959) did recognize the unique-

ness of Sciurillus by placing it in its own subtribe Sciuril-

lina in the tribe Sciurini, molecular studies clearly docu-

ment that Sciurillus has no phyloge ne tic afﬁ nity to other

tree squirrels. Consequently, this genus is now uniformly

placed in its own subfamily, the Sciurillinae (Steppan, Storz,

and Hoffmann 2004; Thorington and Hoffmann 2005;

Thorington et al. 2012). The basal position of Sciurillus

in the family, moreover, suggests great age for its lineage,

which Roth and Mercer (2008) estimated at about 30 mya,

a divergence event only slightly younger than the 36- mya

age for the family. In contrast, the entire radiation of all

remaining tree squirrels in South America, now placed in

the subfamily Sciurinae (Thorington and Hoffmann 2005;

Thorington et al. 2012), appears to have descended from a

single lineage, one that arrived on the continent subsequent

to the closure of the Panamanian land bridge (Roth and

Mercer 2008; Pecnerová and Martínková 2012).

The last revision of South American representatives was

J. A. Allen’s classic monograph Review of the South Amer-

ican Sciuridae, published in 1915 and thus now nearly a

century old. Allen recognized nine genera, a number re-

duced to three or four by subsequent reviewers (e.g., Ell-

erman 1940, three genera [Sciurillus, Microsciurus, and

Sciurus, divided into three subgenera]; Moore 1959, four

genera [Guerlinguetus, Microsciurus, Sciurillus, and Syn-

theosciurus]; Cabrera 1961, three genera [Microsciurus,

Sciurillus, and Sciurus, divided into two subgenera]; R.

S. Hoffmann et al. 1993 and Thorington and Hoffmann

2005, three genera [Microsciurus, Sciurillus, and Sciurus,

divided into three subgenera]). The number of species rec-

ognized in various treatments over the past 50 years has

also differed, sometimes substantially. For example, J. A.

Allen (1915a) divided South American squirrels into 25

species, Cabrera (1961) into 13, Emmons and Feer (1997)

described and mapped 11, and R. S. Hoffmann et al. (1993)

and Thorington and Hoffmann (2005; see also Thoring-

ton et al. 2012) listed 15. In part, differences in species

boundaries recognized by these and other authors attest

to the extreme variability often present geo graph i cally in

color and color pattern. The concordance of both generic

membership and categorical level across these treatments is

limited, a fact also attesting to both the extensive variation

in coloration that characterizes most of these taxa and the

lack of critical reviews.

The lack of any comprehensive or even regional analysis

of character variation for nearly all species of South Ameri-

can squirrels prompted us to examine most available speci-

mens, including holotypes, to assess both genus- and species-

level boundaries. As a result, we recognize seven genera and

seen and often readily heard as they move about or gnaw

on large nuts. They build round, ball- like nests of leaves

and twigs in vine tangles or on tree limbs, or they nest in

tree holes. Capuchin monkeys, Cebus [now Sapajus] apella

(Galetti 1990), and both medium and large felids (Emmons

1978b) are known predators of South American squirrels.

Body size can be arranged into three broadly separable

classes, with one pygmy squirrel (maximum head and body

length about 115 mm, tail length 120 mm, and mass 45 g),

many dwarf squirrels of intermediate size (head and body

length 160– 300 mm, tail length 150– 280 mm, and mass

130– 520 g), and a few large squirrels (head and body length

240– 300 mm, tail length 240– 340 mm, and mass 570–

900 g; Thorington et al. 2012). The skull retains the same

generalized shape in all South American species: brain-

case domed and rounded; interorbital region wide; post-

orbital pro cesses on both frontals and jugals prominent;

rostrum relatively short but broad; zygomatic arches deep

and stout; auditory bullae evenly rounded and relatively

large but rarely excessively inﬂ ated, and divided by sev-

eral transverse bony septa; incisive foramina short; cheek

teeth brachydont and bunodont, with prominent cusps and

transverse crests usually on molars; and dental formula I

1/1, C 0/0, PM 2– 1/1, and M 3/3 = 20– 22, with PM3 re-

duced or absent. Angular pro cesses of the mandible are

slightly inﬂ ected. A baculum is present in the phallus of

males of all species.

Peterka (1937) and Bryant (1945), among others, have

provided detailed descriptions of sciurid morphology. Only

four taxa have been karyotyped: Guerlinguetus brasiliensis

brasiliensis (Lima and Langguth 2002, 2n = 40, FN = 76),

G. b. ingrami (Fagundes et al. 2003, 2n = 40, FN = 74),

Hadrosciurus spadiceus (Lima and Langguth 2002,

2n = 40, FN = 76), and Notosciurus granatensis (Nadler

and Hoffmann 1970, 2n = 42, FN = 78).

Despite their broad range throughout much of the con-

tinent and local ubiquity in all forest types, South Ameri-

can squirrels remain poorly known in nearly all aspects of

their life history and, in par tic u lar relevance to this volume,

their evolutionary relationships and diversiﬁ cation history.

There have been no revisionary studies published in the

last half- century or longer. Furthermore, and in contrast to

virtually all other South American rodent groups, students

of molecular phyloge ne tics have largely bypassed the Sci-

uridae, except for the inclusion of a few species in the phy-

letic delineation of the major lineages in this nearly cosmo-

politan family (Mercer and Roth 2003; Steppan, Storz, and

Hoffmann 2004; Pecnerová and Martínková 2012). These

studies have unambiguously identiﬁ ed two lineages among

the South American representatives of the family. One of

these contains the monotypic Sciurillus, which forms a

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Suborder Sciuromorpha: Family Sciuridae 3

Genus Sciurillus Thomas, 1914

The genus Sciurillus is a tiny rainforest squirrel with three

pairs of mammae, distributed in the Guianas and Amazon

Basin. This is the smallest of the New World tree squirrels,

with a head and body length averaging 113 mm, tail length

114 mm, hindfoot length 30 mm, and body mass 44 g (N = 5).

The ears are small and round, up to 14 mm long. The skull

has a strongly convex dorsal proﬁ le, with the inﬂ exion point

at the middle of the frontals; the rostrum is long, more than

one- third the skull length; the frontals are short, as wide

or wider than their length; the braincase is relatively large

and globose; the temporal part of the orbitotemporal fossa

is completely obliterated, with the postorbital pro cess ex-

tending posteriorly to almost coincide with a vertical plane

at the posteriormost part of the squamosal zygomatic pro-

cess; the anterior portion of the zygoma is nearly vertical,

situated posterior to the ﬁ rst upper molar; a distinct semi-

circular crest, originating at the infraorbital foramen, runs

dorsally along the line of contact between the premaxilla and

maxilla on each side; the anterior opening of the infraorbital

foramen is positioned midway between the anteriormost part

of the zygoma and the incisors, but the infraorbital canal is

not conspicuously elongated; the parapterygoid fossa is deep,

well delimited, and sometimes perforated by bilateral or uni-

lateral vacuities; the posterior aperture of the alisphenoid ca-

nals is small, maximally one- third the width of the foramen

ovale; the aperture of the transverse canal is distinctly visible

in ventral view, and not covered by the posterior border of

the pterygoid encircling the foramen ovale; a sphenopalatine

foramen is situated above the third upper molar or, at most,

over the region between the second and third molars; and

only one bullar septum is present. Very short coronoid pro-

cesses characterize the mandible. Temporal muscles are cor-

respondingly unusually small for a squirrel, but the anterior

deep masseters are enlarged and oriented to assist in retrac-

tion of the jaw (Ball and Roth 1995).

The dental formula is I 1/1, C 0/0 PM 2/1, M 3/3 = 22.

Upper incisors are proodont. PM3 has one root; all re-

maining permanent cheek teeth have three roots. In oc-

clusal view, the maxillary toothrows form a more or less

rounded to slightly oval arrangement. PM3 is small, with a

simple, unicuspidate occlusal surface. PM4 is molariform,

only slightly smaller than M1; M2 is the largest cheek

tooth. PM4 through M2 lack a paraconule, metaconule,

and ectostyle; the trigon of all maxillary teeth is shallow.

The lower cheek teeth lack a paraconid and ectostylid, and

a trigonid is obsolete or absent.

synonyms:

[Myoxus]: Shaw, 1801:171; part (included “le petit guer-

linguet” of Buffon 1789); not Myoxus Zimmerman, 1780.

a total of 19 species. These taxonomic hypotheses differ

from previous treatments of South American squirrels, the

most recent of which is that of Thorington et al. (2012),

and they are ripe for testing by phyloge ne tic and phylogeo-

graphic analyses employing molecular data and methods.

KEY TO THE SUBFAMILIES OF SOUTH AMERICAN

SCIURIDAE (FROM MOORE 1959:199– 200):

1. Lateral wall of cranium with squamosal extending dor-

sally to a point less than or equal to halfway from base

of zygomatic pro cess of squamosal to base of postor-

bital pro cess of frontal; typically two transbullar septa;

well- developed masseteric tubercle; maxilla contribut-

ing much less than half to the lateral side of rostrum;

robust zygoma with a well- developed superior pro cess

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sciurinae

1′. Lateral wall of cranium with squamosal extending dor-

sally notably more than halfway between base of zygo-

matic pro cess of squamosal and base of postorbital pro-

cess of frontal; one transbullar septum; no masseteric

tubercle; maxilla contributing much more than half of

the lateral side of rostrum; slender zygoma lacking supe-

rior pro cess. . . . . . . . . . . . . . . . . . . . . . . . . Sciurillinae

Subfamily Sciurillinae Moore, 1959

The subfamily Sciurillinae contains only the single genus

and species Sciurillus pusillus. This is the smallest of the

New World Sciuridae and one of the two most ancient lin-

eages in the family, the other being the Ratuﬁ nae, or gi-

ant tree squirrels of southern Asia. In erecting this taxon,

Moore (1959:180) delineated 12 diagnostic characters of

the skull, including those listed in the key.

Cladistic analyses of both morphological (Roth 1996)

and molecular characters (allozymes: M. S. Hafner et al.

1994; mitochondrial and nuclear DNA sequences: Mer-

cer and Roth 2003; Herron et al. 2004; Roth and Mer-

cer 2008) support the basal position of the Sciurillinae in

the Sciuridae and refute the hypothesized phyletic link be-

tween Sciurillus and the Asian (Nannosciurus) and African

(Myosciurus) pygmy squirrels proposed by G. S. Miller

and Gidley (1918). Mercer and Roth (2003) estimated the

split between the lineage leading to Sciurillus and all other

sciurids (ﬂ ying, ground, and tree squirrels) at about 36

mya, in the late Eocene. These authors also argued that the

deep phyloge ne tic separation between Sciurillus and other

South American sciurids, which formed a monophyletic

assemblage, indicated separate invasions of the continent

by squirrels. Because Sciurillus has no close relatives in

the family, its place of origin and timing of entry to South

America remain enigmas.

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4 Mammals of South America

molaires [= cheek teeth] sont tout-à- fait semblables à celles

des écureuils,” thus making them essentially equal in that

respect (although “le grand guerlinguet” [= Sciurus aestu-

ans] actually has a distinctive dentition that lacks third up-

per premolars).

Later, when F. Cuvier published a comprehensive ac-

count of mammalian dentition (1823c:161– 162), he erected

the genus Macroxus, characterized by (among other fea-

tures) the presence of two upper premolars, and he included

as species “le guerlinguet” and “le toupaye” (the latter

an oriental squirrel). If his diagnosis is interpreted strictly

and if reference is made to his 1812 publication, “le guer-

linguet” would mean only the “petit guerlinguet.” How-

ever, as he clearly considered (incorrectly) that both “guer-

linguets” had similar dentition (F. Cuvier 1819), it can be

safely assumed that by “guerlinguet” Cuvier meant both

pusillus and aestuans. Husson (1978:382– 383) recognized

this possible confusion and speciﬁ cally designated the spec-

imen described and ﬁ gured by Buffon (1789:261, plate 66:

“le petit guerlinguet”) as the lectotype of Sciurus pusillus

É. Geoffroy St.- Hilaire. Earlier, Thomas (1898g:933) had

designated Sciurus aestuans Linnaeus, 1766 as the type

species of Macroxus F. Cuvier, 1823, thus making this

name unavailable for Sciurillus.

The remaining generic names associated with Sciurillus

pusillus (É. Geoffroy St.- Hilaire) clearly belong either to

other rodent groups (e.g., Myoxus Zimmermann, Gliridae)

or to other groups of sciurids (e.g., Sciurus Linnaeus) and

need not be discussed further here.

Sciurillus pusillus (É. Geoffroy St.- Hilaire, 1803)

Neotropical Pygmy Squirrel

synonyms:

Sciurus pusillus É. Geoffroy St.- Hilaire, 1803:177; type lo-

cality “Cayenne,” French Guiana.

Sciurus guianensis Goldfuss, 1809:122; based on Buffon’s

(1789:261, plate 66) “le petit guerlinguet.”

Sciurus olivascens Illiger, 1815:69; nomen nudum.

Sciurus pusillus Desmarest, 1817c:109; renaming of Sci-

urus pusillus É. Geoffroy St.- Hilaire, 1803; based on

Buffon’s (1789:261, plate 66) “le petit guerlinguet.”

Sciurus olivascens Olfers, 1818:208; part; not olivascens

Illiger, 1815, ﬁ de Hershkovitz, 1959b:345.

Macroxus pusillus: Lesson, 1842:111; name combination.

[Funambulus Pucheranii] pusillus: Fitzinger, 1867a:487;

part; name combination.

Macroxus kuhlii Gray, 1867:433; type locality “Brazil,”

restricted by Thomas (1928c) to Pebas, Loreto, Peru.

Sciurus leucotis Gray, 1867:433; unavailable name pub-

lished as a ju nior synonym and attributed to the French

naturalist François de Caumont LaPorte, Comte de

Castelnau (see Hershkovitz 1959b:345).

Sciurus: É. Geoffroy St.- Hilaire, 1803:177; part (inclusion

of pusillus); not Sciurus Linnaeus, 1758.

Sciurus: Desmarest, 1817c:109; part (inclusion of pusillus);

not Sciurus Linnaeus, 1758.

Macroxus: F. Cuvier, 1823b:119; 1823c:161; part (included

“le guerlinguet” of Buffon, 1789).

[Funambulus]: Fitzinger, 1867a:487; part (inclusion of pu-

sillus); not Funambulus Lesson, 1835.

Sciurus (Macroxus): Liais, 1872:503; part (inclusion of pu-

sillus); not Sciurus Linnaeus, 1758.

Sciurus (Microsciurus): E. W. Nelson, 1899b:32; part (in-

clusion of pusillus and kuhli); not Microsciurus J. A. Al-

len, 1895.

Microsciurus: E.- L. Trouessart, 1904:329; part (inclu-

sion of pusillus and kuhli); not Microsciurus J. A. Al-

len, 1895.

Sciurillus Thomas, 1914f:36 [1914f:415]; type species Sci-

urus pusillus Desmarest, 1817 (= Sciurus pusillus É. Geof-

froy St.- Hilaire, 1803), by original designation.

remarks: Buffon (1789:261, plates 65 and 66, re-

spectively) recognized the existence of two “squirrel- like”

animals in the Guiana region, “le grand guerlinguet” (=

Sciurus aestuans Linnaeus) and “le petit guerlinguet” (=

Sciurillus pusillus É. Geoffroy St.- Hilaire). Buffon used the

vernacular French term “guerlinguet” rather than “écu-

reuil” because he was not convinced that the animals were

“true” squirrels.

When É. Geoffroy St.- Hilaire (1803) named “le petit

guerlinguet” as Sciurus pusillus, he implicitly allied the spe-

cies with other known tree squirrels. Desmarest (1817c:109)

redescribed Geoffroy St.- Hilaire’s Sciurus pusillus and, be-

cause many subsequent authors regarded the taxa proposed

by É. Geoffroy St.- Hilaire (1803) as unavailable manuscript

names, much of the nineteenth- and twentieth- century lit-

erature credited Sciurus pusillus to Desmarest (J. A. Allen

1915a; Ellerman 1940; Cabrera 1961; R. S. Hoffmannet

al. 1993). Desmarest (1822:337) even coined a new com-

mon name for the species, calling it “l’Écureuil nain” or

“dwarf squirrel,” probably to emphasize its similarity to

the Eu ro pe an forms and to abolish the distinctiveness sug-

gested by “guerlinguet.” However, some authors contin-

ued to distinguish between “guerlinguets” and squirrels. F.

Cuvier, in a series of papers dealing with the dentition of

mammals, described and ﬁ gured the dentition of “le petit

guerlinguet” (F. Cuvier 1812:277, plate 15, Fig. 1), the dis-

tinctive dental characters being the size of the cheek teeth

depicted and the presence of a diminutive upper third pre-

molar, a feature present only in Sciurillus pusillus and not

in Sciurus aestuans. Indeed, F. Cuvier did not mention “le

grand guerlinguet” in his 1812 paper, but in a later publi-

cation (F. Cuvier 1819:248), he provided brief descriptions

of the “grand” and “petit” guerlinguets, stating that “leurs

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Suborder Sciuromorpha: Family Sciuridae 5

listed this species in their faunal compilations of Colom-

bia but without locality or voucher specimen citations; Al-

berico et al. (2000) listed S. pusillus from the department

of Caquetá in southeastern Colombia without providing

an exact locality but referring to specimens in the IAvH

and ICN in Bogotá. Jessen, Gwinn, and Koprowski (2013)

included southeastern Colombia and Venezuela south of

the Río Orinoco in the mapped range, without either lit-

erature or specimen documentation. The species has not

been recorded from the fauna of either Ec ua dor (Tirira

2007) or Venezuela (O. J. Linares 1998).

selected localities (Map 2): BRAZIL: Amapá,

Amapá (MPEG 1541), Oiapoque, upper Rio Oiapoque

(MZUSP 20353); Amazonas, Andira, near Villa Bella da

Imperatriz (AMNH 93154), Estirão do Equador, Rio Ja-

vari (MPEG 1557); Mato Grosso, São Manoel, Rio Teles

Pires (type locality of Sciurillus pusillus hoehnei Miranda-

Ribeiro); Pará, Boim, Rio Tapajós (MCZ 30216), Igarapé

Amorim (AMNH 95727), Igarapé Brabo (= Bravo), left

bank Rio Tapajós (AMNH 94743). FRENCH GUIANA:

Cayenne (type locality of Sciurus pusillus É. Geoffroy St.-

Hilaire), Inini, Arataye River (USNM 548447), Paracou

(Voss et al. 2001), Tamanoir, Mana River (FMNH 21788).

GUYANA: Upper Demerara– Berbice, Great Falls of De-

merara River (type locality of Sciurillus pusillus glauci-

nus Thomas). PERU: Loreto, Estación Biológica Que-

brada Blanco (Heymann and Knogge 1997), Pebas (BM

28.7.21.79), Quebrada Orán (M. S. Hafner et al. 1994),

Santa Cecília, Río Maniti (FMNH 87182), Sarayacu, Río

Ucayali (AMNH 76185). SURINAM: Sipaliwini, Emma-

keten [= Emma Keten] (Husson 1978), Frederik Willem IV

Falls, Corentyne River (FMNH 48419).

subspecies: Two (J. A. Allen 1915a) or three (An-

thony and Tate 1935; Cabrera 1961; Thorington and

Hoffman 2005) subspecies have generally been recognized.

These include the nominotypical form (the Guianas and

northeastern Brazil), glaucinus Thomas (the Amazon Ba-

sin of central Brazil and interior Guyana), and kuhlii Gray

(northeastern Peru and probably western Brazil and south-

western Colombia).

Anthony and Tate (1935), who studied specimens from

French Guiana, Peru, and the region of the Rio Tapajós,

Brazil, believed that three subspecies could be distinguished

on the basis of color, having found no signiﬁ cant differ-

ences in cranial mea sure ments, speciﬁ cally: S. p. pusillus—

ears black; head decidedly reddish when compared with

the grayish dorsum; S. p. glaucinus— ears the same color

as the head, which was not very distinct in color from the

rest of the grayish dorsum; and S. p. kuhlii— like glaucinus

but more “saturated,” with “belly even more cinnamon”

(p. 8) than in pusillus and with a darker back. However,

our microscopic examination of the pinnae revealed

Sciurus (Macroxus) pusillus: Liais, 1872:503; name

combination.

Sciurus aestuans var. aestuans: J. A. Allen, 1877:756, part;

not Guerlinguetus aestuans (Linnaeus, 1766).

Macroxus aestuans: E.- L. Trouessart, 1897:428, part (in-

clusion of pusillus and kuhlii as synonyms); not Guer-

linguetus aestuans (Linnaeus, 1766).

Sciurus (Microsciurus) pusillus: E. W. Nelson, 1899b:32;

name combination.

Sciurus (Microsciurus) kuhli: E. W. Nelson, 1899b:32;

name combination and incorrect subsequent spelling of

Macroxus kuhlii Gray.

Microsciurus pusillus: E.- L. Trouessart, 1904:329; name

combination.

Microsciurus kuhli: E.- L. Trouessart, 1904:329; name

combination and incorrect subsequent spelling of Mac-

roxus kuhlii Gray.

Microsciurus kuhlii: J. A. Allen, 1914a:162; name

combination.

[Sciurillus pusillus] kuhli Thomas, 1914b:575; incorrect

subsequent spelling or invalid emendation of Macroxus

kuhlii Gray.

Sciurillus pusillus: Thomas, 1914b:575; ﬁ rst use of current

name combination.

Sciurillus pusillus glaucinus Thomas, 1914b:575, type local-

ity “Great Falls of Demerara River, British Guiana,” =

Ororo Marali, Upper Demerara– Berbice, Guyana.

Sciurillus pusillus pusillus: J. A. Allen, 1915a:197; name

combination.

S[ciurillus]. kuhlii: Thomas, 1928c:291; name combination.

Sciurus pusilus Olalla, 1935:426; incorrect subsequent

spelling of Sciurus pusillus É. Geoffroy St.- Hilaire.

Sciurillus pusillus kuhlii: Cabrera and Yepes, 1940:192;

name combination.

Sciurillus pusillus hoehnei Miranda- Ribeiro, 1941:10; type

locality “Rio Teles Pires (São Manoel), northern Mato

Grosso,” Brazil (page number incorrectly given as 139

by Hershkovitz 1959b:346).

Sciurus aestuans: Rode, 1943:385; part; not Sciurus aestu-

ans Linnaeus.

Microsciurus pusillus hoehnei: C. O. da C. Vieira, 1955:

410; name combination.

Sciurillus guajanensis: Hershkovitz, 1959b:345; not guja-

nensis of Gmelin, 1788, nor guajanensis of Kerr, 1792.

Sciurillus pusilus Moore, 1959:204; incorrect subsequent

spelling of Sciurus pusillus É. Geoffroy St.- Hilaire.

description: As for the genus.

distribution: Sciurillus pusillus is known from

scattered localities in Guyana, Surinam, French Guiana,

Amazonian Brazil in Amapá, Amazonas, Pará, and Mato

Grosso states, and northeastern Amazonian Peru. Cuervo

Díaz et al. (1986) and Rodríguez- Mahecha et al. (1995)

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6 Mammals of South America

Knogge (1997) also suggested that a solitary lifestyle was

the dominant mode of sociospatial or ga ni za tion. Peres et al.

(2003) reported four sightings, each averaging 1.5 indi-

viduals, in the headwaters of the Rio Arapiuns, Pará state,

Brazil, and estimated their density at 0.023 individuals/ha.

Dalecky et al. (2002) stated that home range size in French

Guiana was small, which facilitated individual per sis tence

in habitat islands created by ﬂ ooding. Olalla (1935), Hey-

mann and Knogge (1997), and Voss et al. (2001) observed

this species only in noninundated or terra ﬁ rme forest, and

Olalla (1935) reported a litter size of two.

remarks: Due to the scarcity of specimens of S. pu-

sillus from the time of its discovery in the late eigh teenth

century until the 1930s, the taxonomic history of this

taxon has presented certain per sis tent problems. Most of

the trouble relates to confusion between Sciurillus pusillus

(or some ju nior synonym) and several other South Ameri-

can squirrels, usually Sciurus aestuans Linnaeus (herein

Guerlinguetus aestuans). Because both of these species

were ﬁ rst known from Guianan specimens, their respec-

tive taxonomic histories are entangled. We comment on

all names, available or not, that have been associated

with Sciurillus pusillus; however, new combinations and

changes in speciﬁ c versus subspeciﬁ c status are not dis-

cussed because they can be easily followed in the synon-

ymy provided earlier.

Sciurillus pusillus was the second squirrel to be known

from the Guianan region, the ﬁ rst being Sciurus aestu-

ans, described by Linnaeus in 1766. Ten years later, Buf-

fon (1776:146– 147) published his ﬁ rst report on a Gui-

anan squirrel. The essential parts of Buffon’s report are

translated as: “M. de la Borde, Physician of the King in

Cayenne, said that there is only one species of squirrel in

the Guyanne, that it lives in the forests, that its fur is red-

dish, that it is not larger than the Eu ro pe an rat . . . I am

not quite certain that this animal from the Guyanne, of

which M. de la Borde speaks, is a true squirrel, because

the latter are hardly found in excessively hot climates, such

as that of the Guyanne” (emphases ours). This is a very

uninformative account, but it is clear that Buffon did not

have specimens to describe in 1776, only an oral or written

account by de la Borde. It is also clear that Buffon believed

that there was a single species of “squirrel- like” mammal

in the Guiana region, and that it was not larger than the

Eu ro pe an Rattus. Such reference to size is vague, but it

can be interpreted to mean an animal about the size of a

Norway or black rat; because Sciurillus is so much smaller,

we believe this feature would not go unnoticed even by an

amateur such as de la Borde. Also, Sciurillus pusillus can

hardly be described as having “reddish” fur, a character

that could roughly be applied to Sciurus aestuans. There-

fore, it is plausible that the animal reported by de la Borde

that the epithelium is pigmented (thus appearing black on

dried skins) in specimens from the entire range of the ge-

nus. Moreover, the color of the head is variable in the se-

ries from the Rio Tapajós, as is the intensity of the orange

mixed with gray in the belly. The number of specimens we

have examined from Peru, the putative region of occur-

rence of kuhlii, is small, but a good series from Estirão do

Equador, Rio Javari, Amazonas, which is geo graph i cally

close to the Peruvian localities, cannot be distinguished

from the larger series from the Rio Tapajós, and which has

been ascribed to glaucinus. We have not seen the holotype

of Sciurillus pusillus hoehnei from northern Mato Grosso,

but nothing in the description supports it as a plausible

valid taxon.

Mercer and Roth (2003) reported substantial sequence

divergence in both mitochondrial and nuclear genes be-

tween two specimens from opposite ends of the geographic

range of this species (USNM 578014 from French Guiana,

and LSUMZ 27994 from Loreto department, Peru), posit-

ing a late Miocene divergence between these samples. De-

spite the extensive level of divergence and apparent deep

age of the species, we are unable to ﬁ nd signiﬁ cant, geo-

graph i cally partitioned variation among the few samples

available from this very large area; we provisionally regard

Sciurillus pusillus as a monotypic species.

natural history: Jessen, Gwinn, and Koprowski

(2013) synthesized the few data available on ecol ogy, be-

havior, and life history. Emmons and Feer (1990, 1997),

Ball and Roth (1995), Roth (1996), Heymann and Knogge

(1997), Voss et al. (2001:76), and Youlatos (2011) de-

tailed observations of solitary individuals in trees in well-

drained primary forest, primarily in Inga (French Guiana)

and Parkia (Peru), both in the subfamily Mimosoideae. In

both geographic areas, animals were observed gnawing on

the bark, probably feeding either on cambium or on exu-

dates (sap or gum), incising pieces of the upper bark layer,

and leaving the trunk densely pockmarked. Heymann and

Knogge (1997) recorded large amounts of bark chips that

accumulated on the ground under one such tree. Emmons

and Feer (1990:176) noted that individuals forage on

trunks from near the ground to high in the canopy, “hop-

ping and ﬂ itting from branch to branch,” and traveling

from tree to tree high in the canopy.

Their small size makes individuals difﬁ cult to spot, but

they are not wary of humans and can be easily observed.

Youlatos (2011) observed ﬁ ve individuals using wide verti-

cal substrates in the canopy, with locomotion primarily by

claw climbing. Velhagen and Roth (1997) suggested that a

decrease in bite force with decreasing size might account

for a shift in diet of the pygmy squirrels, including Sciuril-

lus, from hard fruits and nuts to gleaning bark. Because

most sightings were of solitary individuals, Heymann and

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Suborder Sciuromorpha: Family Sciuridae 7

both acknowledged that his usage derived from Gmelin

and decided to keep the name Sciurus guajanensis (his dif-

ferent spelling of the name either is an unjustiﬁ ed emen-

dation or a lapsus calami) for de la Borde’s animal, while

creating the new name Sciurus bancrofti for the other “spe-

cies.” As documented earlier, these two names have nothing

to do with Sciurillus pusillus and do not belong in its syn-

onymy. Hershkovitz (1959b:345) also stated that Turton

(1802:94) employed the name Sciurus guajanensis based

on the “petit guerlinguet” of Buffon. There is, however, no

mention in Turton’s work of the French naturalist or of the

“petit guerlinguet.” Indeed, Turton’s General System is only

an updating of Kerr’s, which is entirely referable to Gmelin

(1788).

Finally, Hershkovitz (1959b:345) cited yet another au-

thor (Goldfuss 1809:122) who used the name Sciurus guia-

nensis for the “petit guerlinguet,” a publication that can

be viewed in the same perspective as those by Kerr and

Turton. However, as neither of us has personally seen the

work by Goldfuss, we herein treat his Sciurus guianensis as

a possible synonym of Sciurillus pusillus.

The ﬁ rst scientiﬁ c name unequivocally associated with

“le petit guerlinguet” is Myoxus guerlingus Var. (?), coined

by Shaw (1801:171), which actually encompasses both

Guianan squirrels, Sciurus aestuans Linnaeus and Sciuril-

lus pusillus Thomas. Because we ﬁ x the type specimen of

Myoxus guerlingus as the same as that of Sciurus aestuans

(guerlingus had no type, being a name for both of Buffon’s

guerlinguets; see the account for Guerlinguetus aestuans),

Shaw’s name cannot be employed as the valid one for “le

petit guerlinguet” and is not listed in the synonymy of S.

pusillus as a result.

É. Geoffroy St.- Hilaire (1803:177) created the name

Sciurus pusillus exclusively for “le petit guerlinguet,” mak-

ing it the valid one for the species. He stated, however, that

de la Borde had collected the specimen he studied. This

made Husson (1978:382) question whether Geoffroy St.-

Hilaire’s specimen was the same as that seen by Buffon,

who obtained his material from Sonini de Manoncourt.

The holotype of Sciurus pusillus É. Geoffroy St.- Hilaire in

the Paris museum (MNHN 312, a mounted skin), however,

has an old label bearing de la Borde’s name attached to its

support. In our opinion, this is probably the same speci-

men seen by Buffon (1789), and de la Borde’s name is on

the label because Buffon believed it represented the “guer-

linguet” he reported. Husson (1978:382) never questioned

that the holotype is really what is currently called Sciurillus

pusillus, in spite of its somewhat deteriorated condition,

but he felt the need to select a lectotype, which he indi-

cated (1978:383) to be the same specimen described and

ﬁ gured by Buffon in 1789 (p. 261, plate 46). Husson’s act,

however, was unnecessary, as the specimen (MNHN 312)

was Sciurus aestuans, by far the most common species

found in the Guianas.

Later, Buffon (1789) was ﬁ nally provided with speci-

mens from French Guiana, which had been collected by M.

Sonini de Manoncourt, and he was able to give ﬁ rsthand

information on the entire squirrel fauna of the region. The

essential aspects of this later contribution are, ﬁ rst, that

Buffon (1789:261– 264, plates 45 and 46) believed the ani-

mals were not real squirrels (e.g., the Eu ro pe an “écureuil”

or Sciurus vulgaris), conﬁ rming his earlier prediction, and

he called them “guerlinguets”; and, second, that he con-

sidered them two species, “le grand” and “le petit guer-

linguet.” He then proceeded to reinterpret his own 1776

account of one species to accommodate the two he now

had in front of him. Thus, the “petit guerlinguet” became

the species “not larger than a rat,” and the “grand” one

was compared in size to Sciurus vulgaris. Obviously, any-

thing the size of Sciurillus is “not larger than a rat,” but

then neither are its ﬂ eas. Buffon’s awkward adaptation

of his earlier, vague size reference to the actual specimens

collected by Sonini de Manoncourt allowed for some mis-

interpretation, mainly because other naturalists were also

busy reporting and cata loging the Guianan sciurid fauna. It

is now important to devote attention to a number of pub-

lications dealing with Guianan squirrels, which appeared

before É. Geoffroy St.- Hilaire (1803) coined the name Sci-

urus pusillus for the “petit guerlinguet” of Buffon, because

Hershkovitz (1959b) considered some of these names to

have priority over pusillus.

The ﬁ rst of these is Sciurus gujanensis, coined by Gme-

lin (1788:152) for two squirrels from different reports con-

cerning the Guianan region. One of these reports is that by

de la Borde, which we discussed earlier and associated with

Sciurus aestuans Linnaeus. The other report commented

on by Gmelin was Bancroft’s (1769:143), taken from his

account of the natural history of Guiana, where he states

that the animal was close to the common En glish squirrel

in size and morphology, and that it had a pale yellowish

brown dorsum, white under parts, small white streaks on

the sides of the body, and a tail similar in color to the dor-

sum but “variegated” with white and black. The presence

of lateral white streaks does not ﬁ t with the color pattern

of any known South American squirrel, and Sciurus aestu-

ans does not have white under parts. Whichever squirrel

Bancroft had in mind, however, both his description of the

color pattern and size completely eliminate the possibility

that his animal was Sciurillus pusillus.

Hershkovitz (1959b:345) ignored Gmelin’s name and

its complete lack of correspondence with Sciurillus pusil-

lus. He proposed that the valid name for the dwarf squirrel

should be Sciurillus guajanensis (Kerr) and then or ga nized

a synonymy with that in mind. However, Kerr (1792:265)

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8 Mammals of South America

came from the same region in Brazil. He later restricted the

type locality of kuhlii Gray to Pebas, Loreto, Peru (Thomas

1928c:291). Because we consider Sciurillus pusillus to be

a monotypic species, the matter of the correct type local-

ity of Macroxus kuhlii, a ju nior synonym, is not critical.

However, because kuhlii has been considered by Anthony

and Tate (1935) to be a valid subspecies (and may even be

a species separate from pusillus, as suggested by the deep

molecular divergence of western and eastern Amazonian

samples; Mercer and Ross 2003), validation of Thomas’s

restriction of the type locality to Peru is warranted.

J. A. Allen (1877:756– 757), without examining speci-

mens of the dwarf squirrel, placed both pusillus and kuhlii

as ju nior synonyms of his excessively broad Sciurus aes-

tuans var. aestuans, believing they could be young speci-

mens of the larger animal. Alston (1878:670) immediately

corrected Allen, considering Sciurus pusillus a valid species

with Macroxus kuhlii as a ju nior synonym. Most subse-

quent authors were convinced of the validity of pusillus but

placed the species in the genus or subgenus Microsciurus

(e.g., E. W. Nelson 1899b; E.- L. Trouessart 1904; J. A. Al-

len 1914a). Any remaining doubts regarding the distinc-

tiveness of pusillus were ﬁ nally dispelled when Thomas

(1914b,f) erected the genus Sciurillus, completely divorcing

it from the larger squirrels of the genus Microsciurus.

Thomas (1914b:575) described Sciurillus pusillus glauci-

nus from Guyana (former British Guiana), thus adding a

third species group taxon to the picture. Anthony and Tate’s

(1935) review of Sciurillus took advantage of the ﬁ rst rela-

tively large series of specimens available at any museum;

they recognized three subspecies, namely, Sciurillus p. pu-

sillus, S. p. glaucinus, and S. p. kuhlii. They restricted the

is the holotype of Sciurus pusillus É. Geoffroy St.- Hilaire,

as the name is applied today, and is characterized by the

small and rounded pinnae and the small size of the feet. It

is not the young of a Sciurus aestuans as proposed by Rode

(1943:385).

Another early name resurrected by Hershkovitz

(1959b:345) was Sciurus olivascens, placed in the syn-

onymy of Sciurillus guajanensis (Kerr) [= Sciurillus pusil-

lus], which he attributed to Olfers, 1818. According to

Hershkovitz (1959b:345), Olfers erected the name after a

manuscript usage by Illiger, purportedly based on “le petit

Guerlinguet” of Buffon, but of this Olfers himself was not

completely sure because he added a question mark after

the French vulgar name. Illiger (1815:69) published the

name Sciurus olivascens in a list of mammals from South

America without diagnosis or description, and he made no

subsequent reference to this name in his brief discussion of

South American squirrels (pp. 74– 75). It thus seems clear

that olivascens is unavailable from Illiger (1815) in the

meaning of the International Code of Zoological Nomen-

clature (ICZN 1999). The same name was again mentioned

in yet another list of South American mammals (Minding

1829:89), also unavailable for the same reasons. Given that

Olfers (1818:208) only questionably associated “Sc. oliva-

scens Ill.” to Buffon’s “le petit guerlinguet,” and that both

Illiger’s (1815) original usage of this name and the subse-

quent usage by Minding (1829) are nomina nuda, it seems

to us that the name continues to be unavailable. But even if

Olfers’s usage were judged available, olivascens would re-

main a ju nior synonym of Sciurus pusillus, being antedated

by E. Geoffroy St.- Hilaire’s name by 15 years.

For some de cades, no new specimens other than the

one(s) in the Paris museum were reported until Gray

(1867:433) listed and described Macroxus pusillus and

Macroxus kuhlii, the latter a new species said to have been

collected by Castelnau in “Brazil.” The name Sciurus leu-

cotis was also given by Gray (1867:433) in the synonymy

of kuhlii and attributed to Castelnau (manuscript); leucotis,

having never been employed as valid, is not available under

the International Code of Zoological Nomenclature (ICZN

1999). Gray’s (1867:433) descriptions are not good, but

one of us (MdV) has seen the holotype of Macroxus kuhlii

(BM 51.7.3.7, skin and skull); the skin is in bad shape, with

extensive loss of hair in the region immediately posterior to

the crown, legs, tail, throat, and chest, and with small losses

in other scattered areas. Nevertheless, what is left of the

animal is very similar to the specimen from Cayenne (BM

45.5.1.3) that probably served as basis for Gray’s account

of Macroxus pusillus. Thomas (1920f:276) assigned two

specimens from Villa Braga on the Rio Tapajós, Pará, Bra-

zil, to S. pusillus but noted that these were “quite like” the

type of Macroxus kuhlii and suggested that the latter likely Map 2 Selected localities for Sciurillus pusillus (●). Contour line = 2,000 m.

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Suborder Sciuromorpha: Family Sciuridae 9

nearly equal- sized orbital and temporal fossa; always

with only one upper premolar . . . . . . . . . . . . . . . . . . 2

2. Mean head and body length of adults between 160 and

180 mm; ears small and rounded . . . . . Syntheosciurus

2′. Mean head and body length of adults >160 mm; ears

relatively large and pointed. . . . . . . . . . . . . . . . . . . . . 3

3. Adult females with three pairs of mammae; distinctive

postauricular patch of different color than that of top of

head usually present . . . . . . . . . . . . . . . . . Notosciurus

3′. Adult females with four pair of mammae; postauricular

patch may or may not be present . . . . . . . . . . . . . . . . 4

4. Pelage coarse; distribution west of the Andes in Ec ua dor

and Peru; pinnae hirsute, with somewhat long hairs, but

not quite forming a tuft . . . . . . . . . . . . . . .Simosciurus

4′. Pelage normal to soft; distribution east of the Andes;

pinnae hirsute, covered with very short hairs . . . . . . . 5

5. Size medium, mean head and body length between 175

and 180 mm . . . . . . . . . . . . . . . . . . . . . . Guerlinguetus

5′. Size large, mean head and body length >215 mm . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hadrosciurus

Genus Guerlinguetus Gray, 1821

Guerlinguetus is a tropical tree squirrel genus occurring

from eastern Colombia, Venezuela, and the Guianas, south

throughout the lowland Amazonian rainforest, then through

the Atlantic Forest of eastern and southern Brazil to north-

eastern Argentina. It is not known to occur in neighboring

Paraguay and Bolivia, where the genus is replaced by the

morphologically very similar Notosciurus. Although wide-

spread in Amazonian Brazil, Guerlinguetus is not known

from south of the Rio Solimões, between the Rio Javari and

Rio Madeira. It is absent from all but the broad gallery forests

at the periphery of the Cerrado biome in central Brazil and

mesic forest enclaves in the Caatinga of eastern Brazil (Car-

mignotto et al. 2012). Species of Guerlinguetus, as is true of

all other South American squirrels, are exclusive forest dwell-

ers. The genus is mostly parapatric with Notosciurus, except

in the general region of the Río Marañón in Peru.

Guerlinguetus is easily distinguished from other South

American squirrels by its size (larger than Microsciurus,

Sciurillus, and Syntheosciurus, but smaller than Hadro-

sciurus and Simosciurus). In comparison with Notosci-

urus, which has an approximately similar size range,

Guerlinguetus is unique in the possession of four pairs

of mammae instead of three and typically by the absence

of a differentially colored postauricular patch. Cranially,

the genus can be distinguished by an orbitotemporal fossa

that is equally divided by a well- developed postorbital

pro cess and an upper second molar with two parallel,

buccolingual crests perpendicular to the anteroposterior

axis of the skull.

nominotypical subspecies to French Guiana, glaucinus to

British Guiana (= Guyana) south to the lower Rio Tapajós,

Pará, Brazil, and kuhlii to northeastern Amazonian Peru.

Another subspecies, Sciurillus pusillus hoehnei, was posthu-

mously published by Miranda- Ribeiro (1941), from north-

ern Mato Grosso, Brazil. For reasons discussed earlier, we

consider Sciurillus pusillus to be a monotypic species, with

kuhlii, glaucinus, and hoehnei as ju nior synonyms.

Subfamily Sciurinae G. Fischer, 1817

Species in the subfamily Sciurinae are the “typical” tree

squirrels, with members distributed on all continents save

Antarctica and Australia. South American representatives

include taxa allocated by Moore (1959) to his subtribes

Sciurina and Microsciurina. Both groups are now collec-

tively placed in the Sciurinae (Thorington and Hoffmann

2005). Species in some genera (e.g., Hadrosciurus, Micro-

sciurus, Notosciurus) are characterized by an ear patch

colored differently than the top of the head, neck, and

shoulders; typically, this patch is continuous from the dis-

tal part of the medial surface of the pinna to just behind

the ear (herein referred to as a postauricular patch). In a

few cases, the colored hairs of the patch protrude above

the rim of the ear. In species of Simosciurus, some hairs

may extend from the dorsal edge of the pinnae, but none of

these resemble a tuft, such as that characterizing the Eu ro-

pe an Sciurus vulgaris (see Thorington et al. 2012).

Phyloge ne tic relationships among the genera and spe-

cies we recognize are very incompletely known, as few of

these taxa have been included in any of the surprisingly

small number of molecular phyloge ne tic studies to date.

The most comprehensive analysis of sciurid phylogeny

(Mercer and Roth 2003) included only three South Ameri-

can taxa of “Sciurus” and Microsciurus. Assuming an en-

try into South America at or subsequent to the closure of

the Panamanian seaway some 3– 4 mya, the net rate of di-

versiﬁ cation of tree squirrels was exceedingly rapid, more

than twice that for any other continental assemblage (Roth

and Mercer 2008).

KEY TO THE GENERA OF SOUTH AMERICAN

SCIURINAE:

1. Mean head and body length of adults <160 mm; orbito-

temporal fossa of skull, in dorsal view, unevenly divided

by the postorbital pro cess of the frontals, resulting in

nearly obliterated temporal fossa; generally with two

upper premolars, the anteriormost very small. . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Microsciurus

1′. Mean head and body length of adults >160 mm; orbi-

totemporal fossa of skull, in dorsal view, evenly divided

by the postorbital pro cess of the frontals, resulting in

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10 Mammals of South America

cept of Guerlinguetus, a view that is very close to ours

herein.

KEY TO THE SPECIES OF GUERLINGUETUS:

1. Ventral coloration orange, or, if mostly orange, only the

throat has white or grayish- white hairs; tail color lined

with orange . . . . . . . . . . . . . . . Guerlinguetus aestuans

1′. Ventral coloration heavily grizzled with white or

grayish- white, either along the entire venter or else only

in throat, chest, and inguinal areas; tail color either

lined with orange or grayish white . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . Guerlinguetus brasiliensis

Guerlinguetus aestuans (Linnaeus, 1766)

Guianan Squirrel, Quatipuru

synonyms:

[Sciurus] aestuans Linnaeus, 1766:88; type locality

“Surinam.”

Sciurus bancrofti Kerr, 1792:265; type locality “Guiana.”

Sciurus guajanensis Kerr, 1792:265; type locality “Cayenne.”

Myoxus Guerlingus Shaw, 1801:171; type locality “Gui-

ana”; based on Buffon’s (1789:261, plate 65) “le grand

guerlinguet.”

Sciurus brasiliensis: Lichtenstein, 1818:216; part; remain-

der under Guerlinguetus brasiliensis.

Sc[iurus]. guianensis Lichtenstein, 1818:216; type locality

not given; name proposed as a synonym of S. brasil-

iensis and based on Bancroft’s report of a squirrel from

(British) Guiana; name became available when Peters

(1864) recognized it as a valid taxon.

Sciurus guerlinguetus Gray, 1821:304; nomen nudum;

name attributed to Linnaeus.

aesciurus stuans: F. Cuvier, 1823c:255; typographic error.

Macroxus aestuans: Lesson, 1827:238; name combination.

Sc[iurus]. gilvigularis Wagner, 1843c:43; nomen nudum.

Sciurus gilvigularis Wagner, 1845a:148; type locality

“nördlichen Brasilien”; name attributed to Johann Nat-

terer (Wagner 1850:283).

S[ciurus]. aestuans: Cabanis, 1848:778; name combination.

Sciurus aestuans var. Guianensis Peters, 1864:655; type lo-

cality “Guiana” = Guyana.

[Funambulus] gilvigularis: Fitzinger, 1867a:486; name

combination.

Sciurus gilviventris Fitzinger, 1867a:486; incorrect subse-

quent spelling of Sciurus gilvigularis Wagner; unavail-

able name cited under [Funambulus] gilvigularis.

[Funambulus] aestuans: Fitzinger, 1867a:486; part; name

combination.

Macroxus aestuans Var. 1: Gray, 1867:432; name combi-

nation.

[S. aestuans] var. aestuans: J. A. Allen, 1877:672; part;

name combination.

We recognize only two species in Guerlinguetus: G. aes-

tuans, a Guianan and Amazonian species, and G. brasilien-

sis, from eastern Amazonia to eastern and southern Brazil

and northern Argentina. Although there are more publica-

tions on aspects of the natural history of these species than

for any other South American squirrels, details of ecol ogy,

behavior, and other aspects of population biology remain

unknown. Guerlinguetus appears to rely heavily on small

palm fruits, using several palm genera through its distri-

bution. Several regional Brazilian vernacular names are

used for these squirrels, without distinction as to species,

including esquilo, serelepe, caxinguelê, and quatipuru, the

last used exclusively in the Brazilian Amazon. In Argentina,

where only G. brasiliensis occurs, the local name is ardilla

gris or ardilla misionera.

synonyms:

Sciurus: Brisson, 1762:107; part (inclusion of brasiliensis);

not Sciurus Linnaeus, 1758; rejected for nomenclatural

purposes (ICZN 1998:Opinion 1894).

Myoxus: Shaw, 1801:171; part (included “le grand guer-

linguet” of Buffon as species); not Myoxus Zimmer-

mann, 1780.

Guerlinguetus Gray, 1821:304; type species Sciurus guer-

linguetus, by tautonomy (“le grand guerlinguet” of Buf-

fon, 1789 = Myoxus Guerlingus Shaw, 1801 = Sciurus

guerlinguetus Gray, 1821 = Sciurus aestuans Linnaeus,

1766. Gray [1821] writes “Gerlinguet, Guerlinguetus,

sciurus guerlinguetus. Lin.”).

Macroxus F. Cuvier, 1823c:161; type species Sciurus aestu-

ans Linnaeus designated by Thomas (1898g:933).

[Funambulus]: Fitzinger, 1867a:486; part (listing of aestu-

ans); not Funambulus Lesson, 1835.

[Sciurus] (Guerlinguetus): Ellerman, 1940:322, 340; as

subgenus.

remarks: The ﬁ rst available generic name for this

group of medium- sized South American squirrels is Guer-

linguetus Gray, 1821, with type species Sciurus guer-

linguetus. Gray attributed S. guerlinguetus to Linnaeus,

a name that is actually a nomen nudum because Gray’s

usage was not followed by any description or plate. How-

ever, Gray’s generic name Guerlinguetus is clearly identiﬁ -

able to the French vernacular “guerlinguet,” which Buf-

fon (1789:261, plates 65, 66) used for the squirrels of the

Guianas, for which he recognized two varieties, a larger

“le grand guerlinguet” [= Guerlinguetus aestuans (Lin-

naeus)], and a smaller “le petit guerlinguet” [= Sciurillus

pusillus (E. Geoffroy St. Hilaire)]. Since Gray (1821), the

name Guerlinguetus has been either ignored or variously

employed for a full genus or subgenus, but at either cat-

egorical level it broadly encompassed most medium- sized

South American squirrels, some now placed in other gen-

era. J. A. Allen (1915a) was the ﬁ rst to restrict the con-

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Suborder Sciuromorpha: Family Sciuridae 11

Sciurus (Guerlinguetus) gilvigularis gilvigularis: C. O. da

C. Vieira, 1955:407; name combination.

Sciurus [(Guerlinguetus)] aestuans poaiae: Cabrera,

1961:361; name combination.

Sciurus [(Guerlinguetus)] aestuans quelchii: Cabrera,

1961:361; name combination.

Sciurus (Guerlinguetus) aestuans georgihernandezi Barriga-

Bonilla, 1966:492; type locality “Margen derecha del

río Vaupés, cerca de Mitú, Vaupés, Colombia, altura

240 m.s.n.m.”

Sciurus (Guerlinguetus) aestuans: Eisenberg, 1989:331;

name combination.

Sciurus (Guerlinguetus) gilvigularis: Eisenberg, 1989:331;

name combination.

description: Adult body weight averages 174 g

(N = 28), head and body length 176 mm (N = 235), tail length

177 mm (N = 205), hindfoot length 47 mm (N = 245), and

ear length 22 mm (N = 83). Dorsal color uniformly brown

to olive brown, heavily streaked with orange throughout,

including the upper parts of the legs; ventral color highly

variable, with throat orange to grayish white and remain-

ing parts orange, sometimes with dark gray hairs inter-

spersed; inner parts of forelimbs colored similarly to chest

and belly; inner parts of hindlimbs darker than belly; area

immediately circumscribing mammae not distinct from re-

mainder of venter, and tail color indistinct from dorsum.

Postauricular patches absent, except for some specimens

from Guianan highlands.

distribution: Guerlinguetus aestuans occurs

throughout Amazonia, in northeastern Peru, eastern Colom-

bia, Venezuela south of the Río Orinoco, and Brazil north of

the Rio Amazonas and west of the Rio Xingu, and through-

out the Guianan region.

selected localities (Map 3): BRAZIL: Amapá,

Cachoeira de Santo Antônio, Rio Jari (MPEG 21804),

Mazagão, Rio Vila Nova (MPEG 438), Vila Velha do Cassi-

poré, Oiapoque (MPEG 6742); Amazonas, Igarapé Auará,

Rio Madeira (AMNH 91831), Santo Isidoro, Tefé (AMNH

78949); Mato Grosso, Cachoeira Dardanelos, right bank

Rio Aripuanã (MPEG 8786), Estação Ecológica Serra das

Araras (MPEG 15272); Pará, Aramapucu, Rio Parú do

Oeste (MPEG 10019), Cussary (AMNH 37453), Foz do

Rio Curuá, right bank Rio Amazonas (MZUSP 4601), Rio

Jamanxim, right bank Rio Tapajós (MPEG 453); Rondônia,

Cachoeira Nazaré, left bank Rio Ji- Paraná (MPEG 20825),

Ouro Preto d’Oeste (MPEG 20529). COLOMBIA: Vau-

pés, La Providencia, Río Apaporis (Barriga- Bonilla 1966),

right bank Río Vaupés, near Mitú (Barriga- Bonilla 1966).

FRENCH GUIANA: River Iracoubo (MNHN 1983/350),

River Oyapock (MNHN 1983/346). GUYANA: Essequibo

Islands– West Demerara, Supinaam River (BM 10.5.4.12);

Upper Takutu– Upper Essequibo, Kanuku Mountains

Sciurus gilviventris Pelzeln, 1883:59; type locality “Borba,”

Amazonas, Brazil (see Goeldi 1893:78).

[Macroxus] gilvigularis: E.- L. Trouessart, 1897:429; name

combination.

Sciurus Quelchii Thomas, 1901e:147; type locality “Ka-

nuku Mountains, British Guiana,” Upper Takutu– Upper

Essequibo, Guyana.

Sciurus Macconnelli Thomas, 1901e:148; type locality “Mt.

Roraima, near its base,” Cuyuni– Mazaruni, Guyana.

[Guerlinguetus] gilvigularis: E.-

L. Trouessart, 1904:328;

name combination.

Sciurus aestuans gilvigularis: J. A. Allen, 1910c:146; name

combination.

Sciurus (Guerlinguetus) aestuans gilvigularis: J. A. Allen,

1911:244; name combination.

G[uerlinguetus]. aestuans: Cabrera, 1912:89; part (also in-

cluded species of Notosciurus); ﬁ rst use of current name

combination.

Guerlinguetus aestuans aestuans: J. A. Allen, 1915a:256;

name combination.

Guerlinguetus aestuans gilvigularis: J. A. Allen, 1915a:256;

name combination.

Guerlinguetus aestuans macconnelli: J. A. Allen, 1915a:

259; name combination.

Guerlinguetus aestuans quelchii: J. A. Allen, 1915a:259;

name combination.

Guerlinguetus aestuans venustus J. A. Allen, 1915a:260; type

locality “Boca Sina (altitude 440 ft.), Rio Cunacunumá

(southern base of Mount Duida), [Amazonas,] Venezuela.”

Guerlinguetus Aestuans Gilvicularis O. M. O. Pinto,

1931:290; incorrect subsequent spelling of Sciurus gil-

vigularis Wagner.

Sciurus [(Guerlinguetus)] aestuans aestuans: Ellerman,

1940:342; name combination.

Sciurus [(Guerlinguetus)] aestuans quelchi: Ellerman,

1940:342; name combination, and incorrect subsequent

spelling of Sciurus quelchii Thomas.

Sciurus [(Guerlinguetus)] aestuans venustus: Ellerman, 1940:

342; name combination.

Guerlinguetus gilvigularis gilvigularis: Moojen, 1942:6;

name combination.

Guerlinguetus poaiae Moojen, 1942:11; type locality “Mata

da Poaia, Tapirapoã, Chapada, Mato- Grosso,” Brazil.

G[uerlinguetus]. gilvigularis gilvigularis: Melo- Leitão, 1943:

356; name combination.

Sciurus (Guerlinguetus) poaiae: W. P. Harris, 1944:11;

name combination.

Sciurus gilvigularis gilvigularis: Moojen, 1952b:25; name

combination.

Sciurus poaiae: Moojen, 1952b:26; name combination.

Sciurus aestuans quelchii: Moojen, 1952b:27; name

combination.

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12 Mammals of South America

are no qualitative cranial distinctions along the entire dis-

tribution; in fact, the species is cranially indistinct from

G. brasiliensis.

We recognize two subspecies. The nominotypical form

occurs in the Guianas, adjacent Venezuela, Amazonian

Brazil, in the states of Roraima, Amazonas (to the north

of the Rio Solimões– Amazonas axis), Amapá, Pará (ex-

cept on the south bank of the Rio Amazonas, to the east

of the Rio Xingu, where the other species of Guerlinguetus

occurs), in northern Peru from scattered localities along

the Río Marañón, and in Colombia near its border with

Brazil. By contrast, G. a. gilvigularis (Wagner, 1845) oc-

curs in Brazil, in the Rio Tapajós and Rio Madeira basins,

and from there southward to the states of Rondônia and

western Mato Grosso. These two taxa can be distinguished

by the color of their ventral surfaces, aestuans with a gray-

ish white throat but orange or yellowish chest and belly,

and gilvigularis with an entirely orange or yellowish venter.

Specimens from the Rio Negro region in Amazonas, Brazil,

exhibit a ventral color pattern characteristic of both sub-

species, suggesting that this area is a zone of intergradation

between aestuans and gilvigularis. Dorsally, all animals are

very similarly colored.

One important aspect of morphological variation in

the distribution of G. a. aestuans is that some animals col-

lected in the highlands of the Guianas and Venezuela pos-

sess postauricular patches whereas in the adjacent lowlands

such patches are either inconspicuous or absent altogether.

The presence of a patch is associated with the names guia-

nensis Peters, 1864, and macconnelli Thomas, 1901, both

of which we regard as synonyms to Guerlinguetus aestuans

aestuans (Linnaeus, 1766).

(type locality of Sciurus Quelchii Thomas). PERU: Loreto,

Orosa, Río Amazonas (AMNH 73917), Sarayacu, Río

Ucayali (AMNH 75277). SURINAM: Para, Zanderij (BM

52.1102). VENEZUELA: Amazonas, Monduapo, right

bank upper Río Orinoco (BM 99.9.11.29– 30); Bolívar, Ci-

udad Bolívar, Suapure, Río Caura valley (AMNH 16128),

El Yagual, lower Río Caura (AMNH 16948).

subspecies: We recognize two subspecies, namely,

G. a. aestuans (Linnaeus, 1766) and G. a. gilvigularis

(Wagner, 1845). Distributions and coat color variation of

each are discussed in the Remarks section.

natural history: There are very few published

ﬁ eld observations of the ecol ogy or behavior of this spe-

cies in the wild. It ranges throughout lowland rainfor-

est and, marginally, into the dry forests of Mato Grosso

state, Brazil. In French Guiana, and likely throughout

this broad range, seed- caching behavior plays an impor-

tant role in the demography of understory palms such as

Astrocaryum sciophilum (see Sist 1989) and other large-

seeded plants such as Carapa procera (Forget 1996).

Forget (1991) reported that this species did not forage at

ground level even when seeds were available. In French

Guiana, this squirrel may persist on small and medium-

sized islands produced by ﬂ ooding after dam construc-

tion, which suggests that their home range size is relatively

small (Dalecky et al. 2002). Peres et al. (2003), from a site

in the headwaters of the Rio Arapiuns, Pará state, Brazil,

estimated density of 0.444 individuals/ha, with sightings

of seven solitary individuals. Voss et al. (2001) observed

animals perched in trees at heights of 3– 30 m above the

ground, with most individuals solitary. Individuals these

authors captured came primarily from primary forest at

both well- drained and swampy sites as well as from some-

what disturbed habitats.

remarks: Use of the name Sciurus aestuans Lin-

naeus by almost all authors from the late eigh teenth and

early nineteenth centuries (e.g., Gmelin 1788:151; Tur-

ton 1802:93; Illiger 1815:75; Thunberg 1823:2; Minding

1829:29; Schinz 1840:144; P. Gervais 1841:37, 1854:308;

Gray 1843b, 1867; Chenu 1850:34) to the present (e.g.,

Emmons and Feer 1997; Eisenberg and Redford 1999;

Thorington and Hoffmann 2005) includes both G. aestu-

ans, as we restrict this species herein, and G. brasiliensis

(next section) or even species we allocate to Notosciurus

(e.g., Alston 1878:668).

Geographic variation of coat color across the distri-

bution of G. aestuans has led authors to name distinct

taxa based on slight variations, all of which names we

regard either as applying to subspecies or as synonyms.

Here, we summarize the geographic variation of charac-

ters to justify our recognition of a single species. There Map 3 Selected localities for Guerlinguetus aestuans (●). Contour line = 2,000 m.

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Suborder Sciuromorpha: Family Sciuridae 13

Guerlinguetus ingrami ingrami: Moojen, 1942:14; name

combination.

Guerlinguetus ingrami henseli: Moojen, 1942:16; name

combination.

G[uerlinguetus]. g[ilvigularis]. paraensis: Melo- Leitão, 1943:

356; name combination.

Sciurus ingrami ingrami: Moojen, 1952b:27; name combi-

nation.

Sciurus ingrami henseli: Moojen, 1952b:28; name combi-

nation.

Sciurus garbei: Moojen, 1952b:28; name combination.

Sciurus (Guerlinguetus) ingrami ingrami: C. O. da C. Vie-

ira, 1953:131; name combination.

Sciurus (Guerlinguetus) ingrami henseli: C. O. da C. Vie-

ira, 1955:407; name combination.

Sciurus (Guerlinguetus) garbei: C. O. da C. Vieira, 1955:

407; name combination.

Sciurus (Guerlinguetus) alphonsei: C. O. da C. Vieira, 1955:

407; name combination.

Sciurus (Guerlinguetus) gilvigularis paraensis: C. O. da C.

Vieira, 1955:407; name combination.

Sciurus gilvigularis paraensis: C. T. Carvalho, 1959:461;

name combination.

Sciurus [(Guerlinguetus)] aestuans alphonsei: Cabrera, 1961:

359; name combination.

Sciurus [(Guerlinguetus)] aestuans henseli: Cabrera, 1961:

360; name combination.

Sciurus [(Guerlinguetus)] aestuans ingrami: Cabrera, 1961:

360; name combination.

Sciurus [(Guerlinguetus)] aestuans paraensis: Cabrera, 1961:

361; name combination.

Sciurus ingrami sebastiani P. Müller and Vesmanis, 1971:

378; type locality “São Sebastião island [= Ilha de São

Sebastião], São Paulo, Brazil.”

Sciurus aestuans ingrami: Avila- Pires, 1977:23; name com-

bination and typographical error.

Sciurus [(Guerlinguetus)] aestuans: Honacki, Kinman, and

Koeppl, 1982:362; name combination; part.

Sciurus aestuans ingrami: Vaz, 1983:34; name combination.

Guerlinguetus alphonsei: J. A. Oliveira and Bonvicino,

2006:348; name combination.

Guerlinguetus henseli: J. A. Oliveira and Bonvicino,

2006:349; name combination.

Sciurus aestuans sebastiani: Hutterer and Peters, 2010:14;

name combination.

description: Adult mass averages 193 g (N = 33),

head and body length 180 mm (N = 175), tail length 183 mm

(N = 171), foot length 46 mm (N = 177), and ear length

22 mm (N = 143). Dorsal color uniform brown to olive

brown, heavily streaked with orange throughout body, in-

cluding upper parts of forelegs and hindlegs; ventral color

Guerlinguetus brasiliensis (Gmelin, 1788)

Brazilian Squirrel, Caxinguelê, Serelepe, Esquilo

synonyms:

[Sciurus] brasiliensis Gmelin, 1788:151; part; based on

“Sciurus brasiliensis” of Marcgraf (1648), cited as syn-

onym; type locality “Brasilia et Guiana,” but restricted

herein to Recife, Pernambuco, Brazil (see Remarks).

Sc[iurus]. guianensis: Lichtenstein, 1818:216; part; re-

mainder under G. aestuans.

Sc[iurus]. brasiliensis Lesson, 1838:468; type locality

“Brésil.”

Sciurus Ingrami Thomas, 1901d:368; type locality “Tun-

nel, Southern Minas Gerais: alt. 1200 m,” Brazil.

Sciurus Roberti Thomas, 1903d:463; type locality “S. Lou-

renço [= São Lourenço], near Pernambuco, Alt. 50 m,”

Pernumbuco, Brazil.

Sciurus aestuans var. paraensis Goeldi and Hagmann,

1904:70; type locality “Mates do Pará,” Pará, Brazil.

[Guerlinguetus] ingrami: E.- L. Trouessart, 1904:328; name

combination.

[Guerlinguetus] roberti: E.- L. Trouessart, 1904:328; name

combination.

Sciurus Alphonsei Thomas, 1906c:442, replacement name

for S. Roberti Thomas, 1903, preoccupied by Sciurus

thaiwanensis roberti Bonhote, 1901:166.

Guerlinguetus alphonsei alphonsei: J. A. Allen, 1915a:261;

name combination.

Guerlinguetus alphonsei paraensis: J. A. Allen, 1915a:261;

name combination.

Guerlinguetus ingrami: J. A. Allen, 1915a:262; name

combination.

Guerlinguetus Aestuans Paraensis: O. M. O. Pinto, 1931:

291; name combination.

Guerlinguetus Aestuans Alphonsei: O. M. O. Pinto, 1931:

292; name combination.

Guerlinguetus Aestuans Garbei O. M. O. Pinto, 1931:

294; type locality “Villa Collatino, Espírito Santo,”

Brazil.

Sciurus [(Guerlinguetus)] ingrami: Ellerman, 1940:343;

name combination.

Sciurus [(Guerlinguetus)] aestuans garbei: Ellerman, 1940:

343; name combination.

Sciurus [(Guerlinguetus)] alphonsei alphonsei: Ellerman,

1940:34; name combination.

Sciurus [(Guerlinguetus)] alphonsei paraensis: Ellerman,

1940: 34; name combination.

Sciurus aestuans henseli Miranda- Ribeiro, 1941:10; type

locality “Porto Feliz, Rio Uruguay, Rio Grande do Sul,”

Brazil.

Guerlinguetus gilvigularis paraensis: Moojen, 1942:9;

name combination.

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14 Mammals of South America

nambuco, Alagoas, Sergipe, and Bahia; G. b. paraensis

(Goeldi and Hagmann) is distributed in eastern Brazilian

Amazonia, in the states of Pará, the south of the Rio Ama-

zonas from the Rio Xingu basin eastward to eastern Ma-

ranhão, Tocantins, and eastern Mato Grosso states; and

G. b. ingrami (Thomas), which is found in coastal Brazil

from Espírito Santo state south to Rio Grande do Sul, and

extending to Misiones in Argentina. See Remarks for com-

ments on the geographic variation of these subspecies.

natural history: Although the distribution sug-

gests that G. brasiliensis inhabits several distinct biomes in

eastern South America, from Amazonian rainforest to the

central Brazilian savannas (Cerrado), northeastern Brazil

dry forests (Caatinga), and Atlantic rainforest, we believe

this species is restricted to forest habitats and does not oc-

cur in open vegetation communities. In the Amazonian and

Atlantic rainforests, this species is common, and specimens

are well represented in collections. In contrast, the species

appears to be rare in the Cerrado and Caatinga, where it is

restricted to marginal gallery forests and isolated pockets

of mesic vegetation, respectively. One of us (MdV) has ob-

served solitary individuals or groups of up to four animals

in the wild, but there are literature reports of associations

of up to eight animals (Solórzano- Filho 2006).

As with other tree squirrels, G. brasiliensis is an arboreal

specialist, shot, trapped, or observed typically at 5– 12 m in

the midstratum layer (Grelle 2003) where it builds nests

made of moss, twigs, leaves, and plant ﬁ bers on the out-

side and exclusively of ﬁ bers internally; nest circumference

averages 56 cm, and nests are usually placed at midcanopy

(Alvarenga and Talamoni 2005). The diurnal activity pat-

tern was represented by two peaks, one in the morning and

other during the afternoon (Bordignon and Monteiro- Filho

2000). Fixed routes between the nests and food resources

have been observed, and animals mark territories with

urine and head- gland rubbing against the substrate. Foods

include lichens, bryophytes, mushrooms, leaves, fruits, in-

sects, and bird eggs (L. F. Ribeiro et al. 2009). At some sites,

coconuts may be the most consumed item, comprising up to

70% of the diet; foods also include exotic plant seeds such

as Pinus, demonstrating this species’ ability to occupy al-

tered habitats (Bordignon and Monteiro- Filho 1999; Alva-

renga and Talamoni 2006). Several studies have discussed

the role of G. brasiliensis as either a seed predator and/or

disperser, especially of palm species with large seeds and

hard endocarps (e.g., Paschoal and Galetti 1995; G. S. Silva

and Tabarelli 2001; Pimentel and Tabarelli 2004; Donatti

et al. 2009).

Alvarenga and Talamoni (2005) and T. G. Oliveira et al.

(2007) presented evidence of reproduction associated with

the onset of the rainy season, but Bordignon and Monteiro-

Filho (2000) captured pregnant females during two peaks,

highly variable, with throat always white to grayish white,

with white hairs sometimes extending to chest and/or belly

and/or inner sides of arms and legs; area immediately around

mammae lighter than rest of venter; and dorsal color of tail

either indistinct from dorsum or, if distinct, grizzled with

whitish bands on hairs. Postauricular patches absent.

distribution: This species is distributed in Ama-

zonian Brazil east of the Iriri and Xingu rivers and south

of the lower Amazon in Pará, Maranhão, Tocantins, and

Mato Grosso states, and extends throughout northeastern

and southwestern Brazil from the states of Ceará to Rio

Grande do Sul and from there to Misiones in Argentina.

The species appears to occupy two disjunct areas, one cor-

responding to the Amazonian subspecies G. b. paraensis

and the other to the two subspecies G. b. brasiliensis and

G. b. ingrami, which occur continuously from northeast-

ern to southern Brazil and Argentina.

selected localities (Map 4): ARGENTINA:

Misiones, Eldorado (BM 26.2.11.1), Santa Ana (FMNH

49885). BRAZIL: Alagoas, Usina Sinimbú, Mangabeiras

(Manimbu) (MZUSP 7532); Bahia, Ilhéus (MZUSP 3501),

Macaco Seco, Monte Andaraí, Rio Paraguaçu (FMNH

20424), Senhor do Bonﬁ m [= Villa Nova] (MZUSP 2608);

Ceará, Serra do Castelo (BM 20.7.1.7); Espírito Santo, Co-

latina, right bank Rio Doce (type locality of Guerlingue-

tus aestuans garbei O. M. O. Pinto), Rio Itaúnas (MZUSP

7071), Rive [= Engenheiro Reeve] (BM 3.9.4.44); Mara-

nhão, Aldeia Gurupiúna, Reserva Indígena Alto Turiaçú

(MPEG 21981), Estreito, right bank Rio Tocantins (MPEG

2444), São Bento (BM 25.5.21.6); Mato Grosso, Serra

do Roncador, 264 km N of Xavantina (BM 79.203– 207);

Minas Gerais, Rio das Velhas, near Lagoa Santa (FMNH

20733); Pará, Belém (MPEG 710), Gradaús, right bank Rio

Fresco (MZUSP 20399), mouth of Rio Bacajá, right bank

Rio Xingú (MZUSP 25455), Portel, Rio Procupi (MPEG

606), Rio Riosinho, left bank Rio Fresco (MPEG 1039),

Santa Júlia, Rio Iriri (BM 20.7.1.6), Urumajó, Arabóia,

east of Bragança, Estrada de Ferro de Bragança (MZUSP

20398); Paraíba, Mamanguape, Camaratuba (MZUSP

8241); Pernambuco, São Lourenço (type locality of Sciurus

Roberti Thomas); Rio de Janeiro, Sepetiba (NMW B1393),

Terezópolis (NMW 1693); Rio Grande do Sul, Colônia do

Mundo Novo (Ihering 1892:107); Santa Catarina, Joinville

(BM 13.7.8.1); São Paulo, Ilha de São Sebastião (type lo-

cality of Sciurus ingrami sebastiani P. Müller and Vesma-

nis), Ilha do Cardoso (MZUSP 3905), Porto Cabral, Rio

Paraná (MZUSP 6021), Presidente Epitácio, left bank Rio

Paraná (MZUSP 3733), Santos (MZUSP 3699), Serra da

Juréia (MZUSP 12835).

subspecies: Guerlinguetus brasiliensis is polytypic:

G. b. brasiliensis (Gmelin) occurs in northeastern Brazil,

in the states of Ceará, Rio Grande do Norte, Paraíba, Per-

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Suborder Sciuromorpha: Family Sciuridae 15

south of this area, intermediate forms between the nomi-

notypical subspecies and G. b. ingrami appear in the state

of Espírito Santo (the form to which O. M. O. Pinto [1931]

gave the name garbei), with specimens having less white

and more yellowish under parts but retaining grizzled tails.

Farther to the south as well as west of Espírito Santo, ani-

mals show white only at the throat and chest, sometimes

also at the posterior and inguinal parts of the venter; the

tail is never grizzled and is always punctuated with orange

or red. The third subspecies, G. b. paraensis from eastern

Brazilian Amazonia, resembles ingrami but has less gray

on the belly; individuals also appear to have shorter pelage

and are slightly smaller. Current rec ords suggest that G. b.

paraensis is not contiguous with the other two subspecies

because G. b. brasiliensis is known only from forest pock-

ets in the otherwise open formation of the Caatinga and G.

b. ingrami occurs in the Atlantic Forest, which is mostly

separated from Amazonia by open savannas of the Cer-

rado, a biome without any squirrel rec ords (Carmignotto

et al. 2012).

Genus Hadrosciurus J. A. Allen, 1915

Hadrosciurus contains the largest of the South American

squirrels, distributed in Venezuela south of the Orinoco,

Colombia, Ec ua dor, and Peru east of the Andes, western

Brazilian Amazonia, and western central Brazil, Bolivia,

and northern Paraguay.

This genus can be distinguished from most other South

American sciurid genera by their size, except for the similar-

sized Simosciurus, which differs from Hadrosciurus by

its distinctly coarser pelage, ears covered by longer hairs,

shorter skull with a broader rostrum, and a trans- Andean

distribution. Individuals of some species of Notosciurus

reach sizes that overlap the lower size limits of Hadrosci-

urus, but the two genera can be distinguished because the

latter exhibits four pair of mammae instead of three.

Hadrosciurus, like other South American squirrels, ex-

hibits considerable variation in external morphology and

especially in color. Mass varies from 450 to 750 g, head and

body length from 215 to 320 mm, tail length from 220 to

330 mm, hindfoot length from 50 to 75 mm, and ear length

from 20 to 40 mm. A well- developed postorbital pro cess

equally divides the orbitotemporal fossa of the skull.

The genus includes three polytypic species, namely,

Hadrosciurus igniventris, with three subspecies, Hadro-

sciurus pyrrhinus, with two subspecies, and Hadrosciurus

spadiceus, with three subspecies. These three species pres-

ent an interesting pattern of sympatry, with their large-

scale ranges overlapping along the lower Solimões and

lower Purús rivers, the only region where museum speci-

mens document the presence of all three. All remaining

one in the winter and the other during the summer; the

latter investigators also estimated a population density of

0.89 squirrels/ha, with males occupying larger areas during

the winter mating period.

remarks: The name for the Brazilian Squirrel is Guer-

linguetus brasiliensis, and its author is Gmelin (1788). The

name was published as a synonym of his Sciurus aestuans

and was attributed to G. Marcgraf (1648). Lesson (1838)

recognized the species as valid, stating that F. Cuvier in

Buffon’s Supplement, Tome I, had provided the name;

however, in this work, F. Cuvier (1831) does not employ

a binomial designation for Marcgraf’s squirrel. Accord-

ing to the International Code of Zoological Nomencla-

ture (ICZN 1999:Arts. 11.6.1 and 50.7), the authorship

of such a name must be attributed to the ﬁ rst publisher of

the name, even if it was given as a ju nior synonym. Gmelin

(1788) gave the type locality as “Brasilia et Guiana,” but

as other authors have argued for the species ﬁ rst reported

by Marcgraf (1648), we restrict it here to Recife, Pernam-

buco, Brazil.

There is considerable geographic variation in the wide

distributional range of this species, and some of this varia-

tion has been recognized at the species or subspecies level.

The material examined does allow us to recognize three

valid subspecies in G. brasiliensis. The typical subspecies,

G. b. brasiliensis, occurs in northeastern Brazil, from the

states of Ceará to Bahia, where the color of the venter is

generally white on the throat, chest, belly, and inner parts

of forelegs and hindlegs, and the tail is grizzled. To the

Map 4 Selected localities for Guerlinguetus brasiliensis (●). The likely disjunct

distributions of Amazonian, Caatinga, and coastal populations are emphasized.

Contour line = 2,000 m.

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16 Mammals of South America

pyrrhinus Thomas as a species— Mesosciurus (Histriosci-

urus) pyrrhinus.

However, Thomas (1927a) suggested that Allen’s genera

were “very intangible” and decided to change the status of

many by placing them as subgenera of Sciurus. In this same

work, Thomas allied pyrrhinus with igniventris, thus, for all

practical purposes, removing pyrrhinus from Mesosciurus.

Shortly thereafter, Thomas (1928b) realized that Sciurus

ﬂ ammifer, the type and only species of Allen’s Hadrosci-

urus, belonged together with all the other large squirrels in

a single taxonomic entity and that, having been described a

few pages earlier than Urosciurus, the former would be the

proper generic name for these squirrels, with Urosciurus as

a ju nior synonym. Except for a few speciﬁ c and subspeciﬁ c

taxa not yet known at the time of Thomas’s publication, the

content he assigned to Hadrosciurus is the same as ours.

Ellerman (1940) did not subscribe to Thomas’s views

and rearranged the speciﬁ c contents of Thomas’s Sciurus

(Hadrosciurus), placing pyrrhinus with most other South

American squirrels in his Sciurus (Guerlinguetus). Moore

(1959) arranged all non- dwarf South American squirrels in

the genus Guerlinguetus, with Hadrosciurus as a subgenus.

Subsequent authors, for example, Thorington and Hoff-

mann (2005), believed that Moore (1959) had addition-

ally recognized Urosciurus J. A. Allen as a subgenus of his

Guerlinguetus. However, Moore (1959:177, 198) included

Sciurus igniventris Wagner, the type species of Urosciurus J.

A. Allen, as Guerlinguetus (Hadrosciurus) igniventris, thus

treating Urosciurus as a ju nior synonym. Because Cabrera’s

(1961) work was published posthumously, he probably

never saw Moore’s (1959) arrangement and so employed

Thomas’s scheme for the genus. The current practice of re-

garding Urosciurus as a valid generic name for the large

South American squirrels has resulted from a misreading of

Moore’s intentions.

Hershkovitz (1959b) expanded the discussion of cranial

characters initiated by J. A. Allen (1915a) by recognizing

two groups of large South American squirrels, which he

treated as distinct species based on a characterization of

the skull as either “short and broad” (Sciurus igniventris)

or “long and narrow” (Sciurus spadiceus). Patton (1984)

performed a morphometric study of Peruvian samples,

agreed with Hershkovitz (1959b) that two distinct skull

morphotypes could be recognized in the area, and resur-

rected Urosciurus as a subgenus of Sciurus.

At present, we cannot ascertain how the three species

and subordinated subspecies that we have recognized in

the genus Hadrosciurus can be distinguished by the cranial

characters described by Hershkovitz (1959b) and Patton

(1984), particularly because the presence of these characters

has not been tested throughout the entire distribution of

the genus. We employ Hadrosciurus as a full genus to keep

sympatry rec ords are for pairs of species, with H. igniven-

tris and H. spadiceus broadly sympatric in Ec ua dor and

northern Peru; H. pyrrhinus and H. spadiceus sympatric

along the middle and lower Rio Juruá in western Brazil;

and H. pyrrhinus and H. igniventris sympatric along the

lower Rio Negro, also in Brazil. It is somewhat surpris-

ing that so few localities have yielded sympatric species of

squirrels, if we consider that, at least potentially, the area

of sympatry at map scale covers much of central Peru and

a vast area of the Brazilian western Amazonia. Patton et al.

(1982) and Lemke et al. (1982) documented sympatry

for H. igniventris and H. spadiceus in northern Peru and

southern Colombia, respectively.

synonyms:

Sciurus: Olfers, 1818:208; part (listing of spadiceus); not

Sciurus Linnaeus, 1758.

Macroxus: Gray, 1867:428; part (listing of langsdorfﬁ i);

not Macroxus F. Cuvier, 1823.

Guerlinguetus: Miranda- Ribeiro, 1914:36; part (listing of

langsdorfﬁ i); not Guerlinguetus Gray, 1821.

Mesosciurus (Histriosciurus) J. A. Allen, 1915a:213; type

species Sciurus gerrardi Gray, by original designation;

part; all species- level taxa in this genus belong to No-

tosciurus J. A. Allen except for J. A. Allen’s Mesosciurus

pyrrhinus.

Hadrosciurus J. A. Allen, 1915a:265; type species Sciurus

ﬂ ammifer Thomas, by original designation.

Urosciurus J. A. Allen, 1915a:267, type species Sciurus tri-

color Tschudi, by original designation.

Sciurus (Urosciurus): Thomas, 1926f:635; name combi-

nation.

Sciurus (Hadrosciurus): Thomas, 1928b:259; name combi-

nation.

[Sciurus] (Guerlinguetus): Ellerman, 1940:340; as subge-

nus (among the included species is Sciurus pyrrhinus

Thomas).

Guerlinguetus (Hadrosciurus): Moore, 1959:177; name

combination.

remarks: Our concept of Hadrosciurus mirrors that

of Thomas (1928b), who suggested that Sciurus (Hadrosci-

urus) encompassed all the largest squirrels east of the Andes,

but this concept requires more detailed clariﬁ cation. As new

species of squirrels were added to the roster of taxa from

South America, J. A. Allen (1915a), acting as the ﬁ rst true

reviewer of the family for the continent, recognized several

distinct genera and subgenera, many of which he described

as new. For the largest squirrels, Allen described two new

genera: Hadrosciurus, containing only H. ﬂ ammifer, and

Urosciurus, with several species, including U. langsdorfﬁ i, U.

igniventris, U. pyrrhonotus, and others. He also created the

genus Mesosciurus for many species and subspecies in the

Sciurus granatensis complex, in which he included Sciurus

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Suborder Sciuromorpha: Family Sciuridae 17

in Roraima, Brazil, about 1,000 km to the west. Cláudia

Regina da Silva (pers. comm. to MdV) has conducted ex-

tensive mammal inventories in Amapá and has never en-

countered this species in the wild, nor has she ever heard

locals refer to such a squirrel.

selected localities (Map 5): BRAZIL: Amazo-

nas, Canabuoca, Paraná do Jacaré (SMF 6918), Igarapé

de Alvarães, right bank Rio Solimões (MZUSP 18921),

Paraná do Aiapuá, Rio Purus (MZUSP 20357), Paraná do

Manhãna, between Rio Japurá and Rio Solimões (MZUSP

6141); Roraima, BR 174, frontier between Brazil and Ven-

ezuela, frontier mark 8 (MZUSP 22347), lower Rio Mu-

cajaí, south of Boa Vista (MZUSP 9779); Roraima, Rio

Catrimani (MZUSP 13688). COLOMBIA: Boyacá, Río

Lengupa (AMNH 38407); Caquetá, Florencia, Río Bodo-

quera (AMNH 33691); Cundinamarca, Sabana Grande,

lowlands near Bogotá (type locality of Sciurus igniventris

taedifer Thomas); Meta, La Macarena, Río Yerli (FMNH

87954). EC UA DOR: Loja, El Porotillo San José (MNHN

1934/1207); Morona– Santiago, Chiguaza (FMNH 54287),

Mendéz Sur (MNHN 1960/3790); Orellano, Río Cotapino

(FMNH 47600); Pichincha, Quito (MNHN 1937/893).

PERU: Cajamarca, Chaupe (AMNH 64047); Loreto, Pe-

bas (AMNH 98426), Santa Elena, Río Samiria (FMNH

87158), Santa Rita, Iquitos (FMNH 87148), Yurimaguas,

Río Huallaga (FMNH 19674); San Martín, about 35 mi

W of Moyobamba, Río Negro (AMNH 73214). VEN-

EZUELA: Amazonas, Nericagua (Caño Usate), Río Ori-

noco (BM 99.9.11.26), Raya 32 km SSE of Puerto Aya-

cucho (USNM 409801); Bolívar, Ciudad Bolívar (AMNH

16127), El Yagual, lower Río Caura (AMNH 16940).

subspecies: We recognize three subspecies of H. ig-

niventris. See Remarks for a description of the geographic

variation.

H. i. cocalis (Thomas, 1900)

synonyms:

Sciurus cocalis Thomas, 1900e:138, type locality “Mouth

of Coca River, Upper Rio Napo,” Orellana, Ec ua dor.

Urosciurus igniventris cocalis: J. A. Allen, 1915a:273;

name combination.

Sciurus (Hadrosciurus) igniventris: Thomas, 1928c:289;

part.

Sciurus [(Hadrosciurus)] igniventris cocalis: Ellerman, 1940:

345; name combination.

Sciurus igniventris cocalis: Soukup, 1961:244; name

combination.

Sciurus [(Urosciurus)] igniventris [cocalis]: Thorington and

Hoffmann, 2005:762; name combination.

This subspecies occurs in Ec ua dor east of the Andes, and

in Peru in the departments of Loreto, Cajamarca, and San

Martín.

what seems to us a distinct group of species— distinguished

mainly by their large size from other South American

taxa— and to separate these squirrels from Sciurus, a near

cosmopolitan taxon. We wish to emphasize that the differ-

ences in skull morphology demonstrated by Patton (1984)

are quite substantive, contrary to what some previous au-

thors believed (such as Oldﬁ eld Thomas). Indeed, we have

not found such trenchant character differences in other

South American squirrels, and it seems highly unlikely that

strongly bimodal distribution is populational or merely

geo graph i cal. Hadrosciurus might, with further study, be

justiﬁ ably split into distinct taxonomic groups based on

their skull morphology alone.

KEY TO THE SPECIES OF HADROSCIURUS:

1. Dorsum black or dark brown background streaked with

red, orange, or yellow (dark hairs with red, orange, or

yellow subterminal bands) . . . . . . . . . . . . . . . . . . . . . 2

1′. Dorsum brownish red to reddish orange with little or

no streaking; postauricular patch small, but distinct. . .

. . . . . . . . . . . . . . . . . . . . . . . . Hadrosciurus pyrrhinus

2. Forelimbs uniformly orange or red, differently colored

from dorsum; postauricular patch rarely present . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . Hadrosciurus igniventris

2′. Forelimbs similar to that of anterior dorsum; postau-

ricular patch always absent. . . .Hadrosciurus spadiceus

Hadrosciurus igniventris (Wagner, 1842)

Northern Amazon Red Squirrel, Quatipuru- açú

synonyms: See under subspecies.

description: Large, with head and body length

246– 300 mm (N = 117 adults), tail length 236– 310 mm

(N = 118), hindfoot length 58– 75 mm (N = 111), ear length

29– 40 mm (N = 73), and mass 453– 700 g (N = 12). Dorsum

dark, distinctly punctuated by light subapical bands to hairs,

with background color from dark brown to black, and light

bands from pale yellow to orange and red. Presence of dis-

tinct postauricular patches variable geo graph i cally; if pres-

ent, pale orange in color. Forelimbs, and sometimes the hind

legs, homogeneously orange to red; ventral color from white

to reddish orange or orange yellow; base of tail colored like

dorsum, with the terminal portion distinctly orange. Com-

pletely or partially melanistic individuals common.

distribution: The species occurs in northwestern

Amazonia from the right bank of the Río Orinoco in Ven-

ezuela southward to the Rio Negro and Rio Solimões in

Brazil and westward to Colombia, Ec ua dor, and north-

ern Peru east of the Andes. The record of two specimens

(MPEG 21807, 21808) from the Brazilian state of Amapá

(Cachoeira do Santo Antonio, Rio Jari) is likely in error be-

cause there are no other rec ords between this locality and

the easternmost known distributional limit of the species

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18 Mammals of South America

Sciurus [(Hadrosciurus)] igniventris igniventris: Ellerman,

1940:344; name combination.

Sciurus [(Hadrosciurus)] igniventris taedifer: Ellerman, 1940:

344; name combination.

Sciurus [(Hadrosciurus)] igniventris zamorae: Ellerman,

1940: 345; name combination.

Hadrosciurus igniventris manhanensis Moojen, 1942:24,

type locality “Paraná do Nanhana [= Manhana; Paynter

and Traylor 1991], entre o Rio Japurá e o Amazonas,

Amazonas,” Brazil.

Hadrosciurus pyrrhonotus purusianus Moojen, 1942:31,

type locality “Lago Aiapuá, Baixo Purús, Estado do

Amazonas,” Brazil.

Sciurus [(Hadrosciurus)] igniventris manhanensis: W. P.

Harris, 1944:12; name combination.

Sciurus [(Hadrosciurus)] igniventris purusianus: W. P. Har-

ris, 1944:12; name combination.

Sciurus igniventris igniventris: Sanborn, 1949b:285; name

combination.

Sciurus igniventris manhanensis: Moojen, 1952b:30; name

combination.

Sciurus pyrrhonotus purusianus: Moojen, 1952b:31; name

combination.

Sciurus (Hadrosciurus) pyrrhonotus purusianus: C. O. da

C. Vieira, 1955:408; name combination.

Guerlinguetus (Hadrosciurus) igniventris: Moore,

1959:203; name combination.

Sciurus [(Urosciurus)] igniventris: Patton, 1984:53; name

combination.

The nominotypical subspecies occurs in Colombia east

of the Andes in Cundinamarca, Meta, and Caquetá depart-

ments; along the Río Napo in Loreto, Peru; in Amazonas

department of Venezuela; and in Roraima state in Brazil,

on the left bank of the Rio Branco, and in Amazonas state,

along the Rio Solimões and Rio Negro.

H. i. ﬂ ammifer (Thomas, 1904)

synonyms:

Sciurus ﬂ ammifer Thomas, 1904d:33, type locality “La Union,

Caura district, Lower Orinoco,” Bolívar, Venezuela.

Hadrosciurus ﬂ ammifer: J. A. Allen, 1915a:266; name

combination.

S[ciurus (Hadrosciurus)]. ﬂ ammifer: Thomas, 1928c:290;

name combination.

Guerlinguetus (Hadrosciurus) ﬂ ammifer: Moore, 1959:203;

name combination.

This subspecies is distributed from the Venezuelan de-

partment of Bolívar, south of the Orinoco and west of Río

Caroní, to the Brazilian border at Serra de Pacaraima.

natural history: Emmons and Feer (1997) noted

that individuals are diurnal, arboreal, territorial, and

H. i. igniventris (Wagner, 1842)

synonyms:

Sciurus igniventris Wagner, 1842c:360; type locality “Rio

Negro”; restricted by Wagner (1850:277) to “Marabi-

tanos, überhaupt vom Rio negro” [= Marabitanas],

right bank of the upper Rio Negro, Amazonas, Brazil.

Sciurus Morio Wagner, 1850:275, type locality “Marabita-

nos, überhaupt vom Rio negro” [= Marabitanas], right

bank of the upper Rio Negro, Amazonas, Brazil.

(?) Macroxus fumigatus Gray, 1867:428, type locality

“Brazil, Upper Amazons”; identity uncertain as holo-

type is almost entirely melanistic.

(?) Macroxus brunneo- niger Gray, 1867:429, type locality

“Brazil”; identity uncertain as holotype is almost en-

tirely melanistic.

Sciurus igniventer Gray, 1867:433; incorrect subsequent

spelling of Sciurus igniventris Wagner.

Sciurus igniventris niger Fitzinger, 1867a:479; type locality

“Am. Brasilien, Rio negro, Marabitanas,” Amazonas,

Brazil.

Sciurus igniventris taedifer Thomas, 1903b:487, type lo-

cality “Sabaña Grande, near Bogota,” Cundinamarca,

Colombia.

Sciurus tricolor morio: E.- L. Trouessart, 1904:328; name

combination.

Sciurus brunneo- niger: E.- L. Trouessart, 1904:328; name

combination.

Sciurus duida J. A. Allen, 1914d:594, type locality “Rio

Cunucunumá (altitude 700 feet), base of Mount Duida,

[Amazonas,] Venezuela.”

Sciurus igniventris zamorae J. A. Allen, 1914d:594, type

locality “Zamora (altitude 2,000 feet), [Zamora–

Chinchipe,] Ec ua dor.”

Urosciurus duida: J. A. Allen, 1915a:270; name

combination.

Urosciurus igniventris igniventris: J. A. Allen, 1915a:271;

name combination.

Urosciurus igniventris taedifer: J. A. Allen, 1915a:272;

name combination.

Urosciurus igniventris zamorae: J. A. Allen, 1915a:274;

name combination.

S[ciurus]. (Hadrosciurus) igniventris: Thomas, 1928c:290;

name combination.

Hadrosciurus igniventris igniventris: Tate, 1939:176; name

combination.

Hadrosciurus igniventris duida: Tate, 1939:176; name

combination.

Hadrosciurus igniventris: Cabrera and Yepes, 1940:195;

name combination.

Sciurus [(Hadrosciurus)] duida: Ellerman, 1940:344; name

combination.

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Suborder Sciuromorpha: Family Sciuridae 19

dorsum black to dark brown heavily streaked with or-

ange or red; venter red, with same color extending to

undersides of legs; and tail dark, mostly black over the

proximal third, with some orange distally. J. A. Allen

(1915a:267– 268) placed Sciurus igniventris Wagner in his

new genus Urosciurus.

Hadrosciurus igniventris ﬂ ammifer is similar to H. i.

igniventris, with orange to red limbs and a black dorsum

heavily streaked with pale yellow to orange, but the former

differs from the latter by having conspicuous pale orange

postauricular patches and a white venter. The genus Had-

rosciurus was created by J. A. Allen (1915a), with Sciurus

ﬂ ammifer as its type and only species.

Hadrosciurus pyrrhinus (Thomas, 1898)

Junín Red Squirrel, Quatipuru- açú

synonyms: See under subspecies.

description: Equivalent in size to H. igniventris,

with head and body length 220– 290 mm (N = 44 adults),

tail length 252– 305 mm (N = 42), hindfoot length 60–

70 mm (N = 40), and ear length 31– 35 mm (N = 19). Color

of dorsum, forelimbs, and hindlegs uniform dark red to

brownish red, without streaking by hairs with light api-

cal or subapical bands. Conspicuous postauricular patches

present, generally pale yellow or light orange throughout

most of range but inconspicuous in specimens from up-

per Rio Negro area of Brazil at Colombian border. Venter

white to pale red, mixture of both colors in some individu-

als; when white, transition from venter to torso, limbs,

and head demarcated by a red strip.

distribution: This species occurs from the eastern

Andean slopes of central Peru to western Brazilian Amazo-

nia, probably limited by the Purus river basin on the east

and Negro basin on the north.

selected localities (Map 6): BRAZIL: Amazo-

nas, between Cajutuba and Airão, Rio Jaú, Rio Negro

(syntype of Sciurus igniventris Wagner), Codajás, left

bank Rio Solimões (MZUSP 4202), Dejedá, Rio Juruá

(type locality of Urosciurus nigratus O. M. O. Pinto), Es-

tirão do Equador, right bank Rio Javari (MPEG 1807),

Manaus (syntype of Sciurus igniventris Wagner), Santo

Antônio do Içá, left bank Rio Içá (MZUSP 20384), Ta-

huapunto (Tauá), Rio Vaupés, left bank (AMNH 78620).

PERU: Cusco, San Juan Grande, Quincemil, Quispi-

canchis (FMNH 75208); Huánuco, Tingo Maria, Río

Huallaga (FMNH 24112); Junín, Tarma, 2 mi NW of

San Ramón (AMNH 231771); Pasco, Puerto Bermudez

(FMNH 24110).

subspecies: We tentatively recognize two subspe-

cies of H. pyrrhinus, which we compare in the Remarks

section.

solitary, although several may feed together in a single

fruiting tree. Primary foods include large palm nuts and

other tree nuts and fruits. Silvius (2002) reported indi-

viduals feeding on beetle larvae as well as the endocarp

of the palm Attalea maripa. The species is largely limited

to closed- canopy rainforest, either mature or disturbed

(Emmons and Feer 1997). Youlatos (1999) reported that

individuals used all forest heights, preferring tree crowns

and lianas, with an ability to walk, leap, and bound

along small horizontal supports. Jessen, Palmer, and

Koprowski (2013) reported a nest for an unidentiﬁ able

melanistic female of either H. igniventris or H. spadiceus

from Loreto, Peru, built at 3.6 m above ground with two

offspring.

remarks: Hadrosciurus igniventris cocalis is a large

squirrel with orange forelimbs, which differs from the

nominotypical subspecies by a dark to almost black dor-

sal color streaked with orange or dark red, the presence of

conspicuous orange postauricular patches, and a yellowish

orange venter.

Hadrosciurus igniventris igniventris is also a large

squirrel with orange to red forelimbs, with hindlimbs col-

ored as the forelimbs or dorsum; head blackish above,

with an orange muzzle diluted with black and with no

conspicuous eye- ring; ears brown to almost black on

both sides; postauricular patches, if present, inconspicuous;

Map 5 Selected localities for Hadrosciurus igniventris (●). Contour line = 2,000 m.

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20 Mammals of South America

comes clear when it is considered together with its sister

subspecies, as we shall do.

Hadrosciurus pyrrhinus fulminatus is similar to H. p.

pyrrhinus except that dorsal tones are paler, duller red;

pale orange postauricular patches are conspicuous; the

venter is uniformly yellow to orange. This taxon occurs

sympatrically with two other forms in the genus (namely

H. i. igniventris and H. spadiceus pyrrhonotus) in the region

of the lower Rio Solimões and Rio Purus in the Brazilian

state of Amazonas. In this area of sympatry, H. pyrrhinus

can be distinguished from H. i. igniventris by its uniform

dull dark red dorsal color with very little streaking (which

results from a pale band near the tip of the hairs) and the

possession of conspicuous postauricular patches. Hadro-

sciurus pyrrhinus is distinguished from H. spadiceus pyr-

rhonotus by presence of distinct postauricular patches, a

less vivid red dorsum with less streaking, and yellowish

orange as opposed to white venter. Since its description,

fulminatus has been allied to Sciurus igniventris, and

since Thomas (1927a) transferred pyrrhinus to the igni-

ventris complex, both have been usually treated as dis-

tinct subspecies under igniventris. The similarity in coat

color between pyrrhinus and fulminatus made us con-

sider a probable afﬁ nity, here inferred at the subspeciﬁ c

level.

H. p. fulminatus (Thomas, 1926)

synonyms:

Sciurus [(Urosciurus)] igniventris fulminatus Thomas,

1926f:637; type locality “Manacapuru,” Rio Solimões,

Amazonas, Brazil.

S[ciurus. (Urosciurus)] i[gniventris]. fulminatus: Thomas,

1928c:290; name combination.

Urosciurus nigratus O. M. O. Pinto, 1931:309; type local-

ity “Rio Juruá (Amazonas),” Brazil.

Sciurus [(Hadrosciurus)] nigratus: Ellerman, 1940:344;

name combination.

Sciurus [(Hadrosciurus)] igniventris fulminatus: Ellerman,

1940:345; name combination.

Hadrosciurus igniventris fulminatus: Moojen, 1942:22;

name combination.

Sciurus igniventris fulminatus: Moojen, 1952b:29; name

combination.

Sciurus (Hadrosciurus) pyrrhonotus fulminatus: Avila-

Pires, 1964:15; name combination.

This subspecies occurs primarily in Amazonas state,

Brazil; it is also known from a single locality in Cusco de-

partment, Peru.

H. p. pyrrhinus (Thomas, 1898)

synonyms:

Sciurus pyrrhinus Thomas, 1898f:265; type locality “Gari-

tal del Sol, Vitoc, [Junín,] Peru.”

Mesosciurus [(Histriosciurus)] pyrrhinus: J. A. Allen,

1915a:252; name combination.

Sciurus [(Urosciurus)] pyrrhinus: Thomas, 1927a:369;

implicit allocation to subgenus due to accepted resem-

blance to S. igniventris.

Sciurus [(Guerlinguetus)] pyrrhinus: Ellerman, 1940:342;

name combination.

Sciurus (Hadrosciurus) pyrrhinus: Cabrera, 1961:378;

name combination.

The nominotypical subspecies occurs on the eastern

slopes of the Andes in the Peruvian departments of Huá-

nuco, Junín, Pasco, and Ucayali.

natural history: Emmons and Feer (1997) stated

that this species inhabits montane forests across an eleva-

tional range from 600 to 2,000 m, but it occurs at less than

300 m elevation at localities in the central Amazon.

remarks: The taxonomic history of Hadrosciurus

pyrrhinus (Thomas) is marked by its erroneous allocation

to a group of Andean and peri- Andean taxa allied to No-

tosciurus granatensis by J. A. Allen (1915a). Although it is

superﬁ cially similar to these squirrels, its distribution on

the eastern slopes of the central Peruvian Andes only added

to the confusion. The alliance of pyrrhinus to other large

South American squirrels we include in Hadrosciurus be- Map 6 Selected localities for Hadrosciurus pyrrhinus (●). Contour line = 2,000 m.

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Suborder Sciuromorpha: Family Sciuridae 21

EC UA DOR: Morona– Santiago, Río San José (AMNH

68264); Napo, mouth of Río Coca, upper Río Napo (BM

0.6.3.2), boca del Río Lagarto Cocha (AMNH 72231);

Pastaza, Mera (AMNH 67674). PARAGUAY: Alto Para-

guay, Estancia Kambá Aka, Parque Nacional Río Negro

(D’Elía, Mora et al. 2008). PERU: Cajamarca, Huarandosa

(AMNH 67095); Huánuco, Tingo Maria, Río Huallaga

(FMNH 24113); Loreto, boca del Río Curaray (AMNH

72177), Río Apayacu (AMNH 74389), San Fernando, Río

Yavari (FMNH 88967); Puno, Pauquiplaya, below San Ig-

nacio, Río Tambopata (FMNH 79903).

subspecies: We tentatively recognize four subspecies

of H. spadiceus, as follows.

H. s. castus (Thomas, 1903)

synonyms:

Sciurus castus Thomas, 1903b:488, type locality “Chi-

mate, [La Paz,] Bolivia, . . . on the Upper Rio Beni.”

Urosciurus pyrrhonotus castus: J. A. Allen, 1915a:276;

name combination.

Hadrosciurus pyrrhonotus castus: Cabrera and Yepes, 1940:

195; name combination.

Sciurus [(Hadrosciurus)] pyrrhonotus castus: Ellerman,

1940:345; name combination.

Hadrosciurus langsdorfﬁ i rondoniae Moojen, 1942:39;

type locality “Cabeceiras do Rio Aripuanã, Rondonia,

Mato- Grosso,” Brazil.

Sciurus [(Hadrosciurus)] langsdorfﬁ i rondoniae: W. P. Har-

ris, 1944:12; name combination.

Sciurus langsdorfﬁ i rondoniae: Moojen, 1952b:32; name

combination.

Sciurus pyrrhonotus castus: Soukup, 1961:244; name

combination.

This subspecies is known from Rondônia state, Brazil,

and an isolated locality in La Paz department, Bolivia. It is

quite similar morphologically to the nominotypical form,

differing primarily by more extensive white on the venter

and the lack of a conspicuous eye- ring. The head is black,

mixed with orange on the crown; the rostrum also black

and orange; the ears brown on both sides; postauricular

patches are absent; the dorsum black, ﬁ nely streaked with

vivid yellow; forefeet and limbs the same color as the dor-

sum, only slightly lighter distally; hindfeet orange streaked

with black; bicolored legs— their anterior portion the same

color as the sides but darker orange posteriorly; venter

entirely white to white with some yellow from throat to

chest, white extending to the undersides of fore and hind

limbs almost to the feet, with the color near the feet the

same as dorsal parts of the legs; and the proximal quarter

to third of the tail is dark, and also turning black with an

orange tip. Hadrosciurus langsdorfﬁ i rondoniae Moojen,

Hadrosciurus spadiceus (Olfers, 1818)

Southern Amazon Red Squirrel, Quatipuru- açú

synonyms: See under subspecies.

description: Slightly larger than other species in

the genus, with head and body length 215– 316 mm (N = 78

adults), tail length 225– 325 mm (N = 78), hindfoot 50–

75 mm (N = 76), ear length 20– 35 mm (N = 36), and mass

650– 750 g (N = 3). Highly variable species, as apparent

from several subspecies we recognize (see Remarks for

distinctive characters of each subspecies). Dorsum dark,

with black background color, strongly streaked with pale

yellow, yellow, orange, or red; specimens from Peru have

darker mid- dorsal strip. Alternatively, the dorsum may be

mostly or entirely red, or divided into relatively distinct an-

terior and posterior regions, with posterior part uniformly

colored orange but anterior portion dark and streaked.

Dorsal sides of forelimbs and hindlimbs colored same as

adjacent region of upper body; in southeastern portion

of distribution, hindlegs divided into paler anterior and

darker posterior regions. Differently colored postauricular

patches absent. Ventral color variable: white in specimens

from Brazilian Amazon and parts of Bolivia, white to yel-

low in those from Peru, and red in west- central Brazil, cen-

tral Bolivia, and northern Paraguay.

distribution: The species occurs from southern Co-

lombia, Ec ua dor, and Peru east of the Andes, eastward into

Amazonian Brazil, where it is limited to the region south of

the Rio Solimões- Amazonas system, from the Rio Tapajós

west to the Peruvian border, and south to west- central Bra-

zil in the states of Mato Grosso and Mato Grosso do Sul as

well as across the lowlands of Bolivia to northern Paraguay.

selected localities (Map 7): BOLIVIA: La Paz, Chi-

mate (BM 1.2.1.7); Santa Cruz, Santa Cruz de la Sierra

(FMNH 21420). BRAZIL: Acre, Oriente (= Seringal Ori-

ente), near Taumaturgo, Rio Juruá (MPEG 725); Amazonas,

Eirunepé, left bank Rio Juruá (MZUSP 4634), Lago do Ba-

tista, Ilha de Tupinambarama, right bank Rio Amazonas

(MZUSP 4567), Lago do Mapixi, east of Rio Purus (FMNH

38881), Manaqueri, Rio Solimões (NMW B 1533), mouth

Rio Guariba, left bank Rio Aripuanã (MPEG 8785); Mato

Grosso, Mato Grosso (= Vila Bela da Santíssima Trindade)

(NMW B 1520), Palmeiras (MZUSP 6338), Villa Maria,

NNW of Cáceres (NMW B 1518); Mato Grosso do Sul,

Urucum (type locality of Sciurus langsdorfﬁ i urucumus J. A.

Allen); Pará, Flexal, Itaituba– Jacareacanga road, km 212,

left bank Rio Tapajós (MPEG 10165), Rio Arapiuns, left

bank Rio Tapajós (MPEG 424), Sumaúma, left bank Rio

Tapajós (MZUSP 20365); Rondônia, Ouro Preto d’Oeste,

Rio Paraíso (MPEG 12411). COLOMBIA: Meta, Cabaña

Duda, junction of Río Duda and Río Guayabero (Lemke

et al. 1982), Fundo “Guami,” Piñalto (Lemke et al. 1982).

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22 Mammals of South America

Sciurus Langsdorﬁ : Thomas, 1900e:137; name combina-

tion and incorrect subsequent spelling of Sciurus langs-

dorfﬁ i Brandt.

Sciurus langsdorfﬁ i urucumus J. A. Allen, 1914d:595;

type locality “Urucum, (altitude 400 feet), Rio Para-

guay (at mouth of Rio Tacuari), [Mato Grosso do Sul,]

Brazil.”

Sciurus langsdorfﬁ i steinbachi J. A. Allen, 1914d:596; type

locality “Santa Cruz de La Sierra, [Santa Cruz,] Bolivia.”

Guerlinguetus langsdorfﬁ : Miranda- Ribeiro, 1914:36;

name combination and incorrect subsequent spelling of

Sciurus langsdorfﬁ i Brandt.

Urosciurus langsdorfﬁ i langsdorfﬁ i: J. A. Allen, 1915a:276;

name combination.

Urosciurus langsdorfﬁ i urucumus: J. A. Allen, 1915a:278;

name combination.

Urosciurus langsdorfﬁ i steinbachi: J. A. Allen, 1915a:279;

name combination.

Urosciurus langsdorfﬁ urucumus: J. A. Allen, 1916d:572;

name combination.

Hadrosciurus langsdorfﬁ i urucumus: Cabrera and Yepes,

1940:196; name combination.

Hadrosciurus langsdorfﬁ i steinbachi: Cabrera and Yepes,

1940:196; name combination.

Sciurus [(Hadrosciurus)] langsdorfﬁ langsdorfﬁ : Ellerman,

1940:345; name combination.

Sciurus [(Hadrosciurus)] langsdorfﬁ urucumus: Ellerman,

1940:345; name combination.

Sciurus [(Hadrosciurus)] langsdorfﬁ steinbachi: Ellerman,

1940:345; name combination.

Hadrosciurus langsdorfﬁ i langsdorfﬁ i: Moojen, 1942:36;

name combination.

Sciurus langsdorfﬁ i langsdorfﬁ i: Moojen, 1952b:32; name

combination.

Guerlinguetus (Hadrosciurus) urucumus: Moore, 1959:

203; name combination.

Guerlinguetus (Hadrosciurus) spadiceus: C. T. Carvalho,

1965:52; name combination.

Sciurus [(Urosciurus)] spadiceus: Patton, 1984:53; name

combination.

Sciurus (Hadrosciurus) spadiceus: Eisenberg, 1989:335;

name combination.

Sciurus [(Urosciurus)] spadiceus [steinbachi]: Thorington

and Hoffmann, 2005:763; name combination.

Urosciurus spadiceus: J. A. Oliveira and Bonvicino, 2006:

350; name combination.

The nominotypical subspecies occurs in the southern

part of Mato Grosso and northwestern Mato Grosso do

Sul states, Brazil, and extends west to Santa Cruz depart-

ment, Bolivia. Specimens are characterized by a black

head mixed with orange on the crown, a black and or-

ange muzzle, a conspicuous eye- ring, ears brown on both

from west- central Brazil, is identical to castus and thus is

placed as a synonym.

H. s. pyrrhonotus (Wagner, 1842)

synonyms:

Sciurus pyrrhonotus Wagner, 1842c:360, type locality

“Borba,” Amazonas, Brazil.

Urosciurus pyrrhonotus pyrrhonotus: J. A. Allen, 1915a:

275; name combination.

Urosciurus pyrrhonotus: O. M. O. Pinto, 1931:303; name

combination.

Sciurus [(Hadrosciurus)] pyrrhonotus pyrrhonotus: Eller-

man, 1940:345; name combination.

Hadrosciurus pyrrhonotus pyrrhonotus: Moojen, 1942:26;

name combination.

Sciurus pyrrhonotus pyrrhonotus: Moojen, 1952b:30; name

combination.

Sciurus pirrhonotus Alho, 1982:144; incorrect subsequent

spelling of Sciurus pyrrhonotus Wagner.

The range of H. s. pyrrhonotus is limited to Amazonas

state, Brazil, south of the Rio Amazonas– Solimões along

the lower and middle courses of the Rio Purus and Rio Ma-

deira. Specimens typically have a black head but orange ros-

trum, and lack a conspicuous eye- ring. The ears are red on

both sides. Postauricular patches are absent. The dorsum is

red, slightly darker on the midline. The forelimbs and feet

are the same color as the dorsum, only slightly lighter dis-

tally. The hindlegs and feet are reddish. The venter is white,

with the white extending to the undersides of arms and legs

almost to the feet. The color near forefeet and hindfeet is

the same as dorsal parts of the legs. The tail is dark over the

proximal third of its length, with red predominating in the

middle section and black predominating distally. Some spec-

imens of this subspecies have the dorsum partially divided

into anterior and posterior sections with slightly distinct

hues of red and distinct amounts of banding of the hairs, but

the dorsal color is always red, and the venter is white.

H. s. spadiceus (Olfers, 1818)

Sc[iurus]. spadiceus Olfers, 1818:208; type locality re-

stricted by Hershkovitz (1959b:346) to Cuiabá, Mato

Grosso, Brasil.

Sciurus langsdorfﬁ i Brandt, 1835:89, type locality “Bra-

zil,” restricted to Cuiabá, Mato Grosso, Brazil by Wag-

ner (1850:275).

Sciurus Langsdorﬁ i Gray, 1843b:137; incorrect subsequent

spelling of Sciurus langsdorfﬁ i Brandt.

Sciurus langsdorfﬁ i Var. rufa Wagner, 1844:plate CCXV;

preoccupied by Sciurus rufus Kerr, 1792.

Sciurus Langsdorfﬁ : E.- L. Trouessart, 1897:429; name

combination and incorrect subsequent spelling of Sci-

urus langsdorfﬁ i Brandt.

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Suborder Sciuromorpha: Family Sciuridae 23

a tributary on the right bank of the middle Rio Madeira,

Amazonas, Brazil, and from there to Pará state west of the

Rio Tapajós.

Melanistic individuals of H. s. tricolor are common, with

some individuals entirely black, others only partially so. The

color pattern for non- melanistic specimens includes a black

head with orange rostrum, no conspicuous eye- ring, reddish

brown to brownish black ears on both sides; no postauricu-

lar patches; dorsum generally black, sometimes darker red

posteriorly, but heavily streaked with orange; forefeet and

limbs same color as the dorsum, only slightly paler distally;

hindfeet and limbs reddish orange, with or without black

streaking; venter white, extending to undersides of forelegs

and hindlegs almost to the paws, color near hands and feet

the same as dorsal parts of the legs; proximal third of the

tail dark, with red predominating in the middle section and

orange and black predominating distally.

natural history: Gwinn et al. (2012) compiled

information on the natural history of this species. Peres,

Barlow, and Haugaasen (2003) reported an estimated den-

sity of 0.08 individuals/ha at a site in the headwaters of

Rio Arapiuns, Pará state, Brazil; they observed solitary in-

dividuals twice. H. Gómez et al. (2003) reported a larger

average density for a site in the department of La Paz,

Bolivia, with 0.122 individuals/ha. Patton et al. (2000) re-

corded scrotal males in the headwaters of the Rio Juruá,

Amazonas state, Brazil, in September and February. They

also collected a lactating female with three placental scars

in February. Anderson (1997) reported litter sizes ranging

from two to four in August from Bolivia. These very lim-

ited data suggest breeding in both wet and dry seasons. Ac-

cording to sources cited in Gwinn et al. (2012), the species

is mostly seen at ground level, using the canopy when the

forest is ﬂ ooded. Nuts of four genera of trees (the palms

Astrocaryum, Attalea, and Scheelea, and the legume Dip-

teryx) comprise the majority of its diet. Della- Flora et al.

(2013) reported the association of this species with mixed-

species bird ﬂ ocks in western Brazil.

remarks: The distributions of three subspecies (H. s.

castus, H. s. tricolor, and H. s. spadiceus) approach one

another in Bolivia. Here, those specimens we assign to tri-

color differ from both spadiceus and castus in dorsal color,

with the dorsum black streaked with red or orange, not

the pale yellow characteristic of the latter two subspecies.

However, Bolivian tricolor also differ from their counter-

parts east of the Andes in Ec ua dor and Peru, where speci-

mens typically exhibit a mid- dorsum darker than the sides,

a characteristic not present in Bolivian tricolor. Samples

from the western Brazilian Amazon, in the states of Acre

and Amazonas, also differ from typical tricolor, with the

mid- dorsum not markedly darker than the sides and the

venter tending to white, with some orange. The dorsum of

sides; no postauricular patches; dorsum black ﬁ nely lined

with yellow and slightly darker on the midline; forefeet

and limbs the same color as dorsum, only slightly lighter

distally; black hindfeet streaked with orange; bicolored

hindlegs, anteriorly the same pale orange as the sides but

posteriorly darker reddish; venter yellowish orange, with

some white on the throat and with color extending to the

undersides of forelegs and hindlegs, almost to the feet;

color near feet same as dorsal parts of legs; and proximal

quarter of tail dark, becoming black before terminating

in an orange tip.

H. s. tricolor (Tschudi, 1845)

synonyms:

Sc[iurus]. tricolor Tschudi, 1845:156, plate XI; type local-

ity “Peru”; restricted to “Maynas, in the angle between

the lower Huallaga and the Marañon [Loreto]” (Thomas

1900e:137).

Urosciurus tricolor: J. A. Allen, 1915a:269; name combi-

nation.

Sciurus (Urosciurus) pyrrhonotus taparius Thomas, 1926f:

635; type locality “Uricurituba, Santarem [= San-

tarém],” Pará, Brazil.

Sciurus [(Urosciurus)] pyrrhonotus juralis Thomas,

1926f:636, type locality “Juruá River,” Amazonas,

Brazil.

Sciurus (Urosciurus) tricolor: Thomas, 1927f:599; name

combination.

Sciurus (Hadrosciurus) tricolor: Thomas, 1927f:599; name

combination.

Hadrosciurus tricolor: Cabrera and Yepes, 1940:195;

name combination.

Sciurus [(Hadrosciurus)] pyrrhonotus taparius: Ellerman,

1940:345; name combination.

Sciurus [(Hadrosciurus)] pyrrhonotus juralis: Ellerman,

1940:345; name combination.

Hadrosciurus pyrrhonotus taparius: Moojen, 1942:30;

name combination.

Hadrosciurus pyrrhonotus juralis: Moojen, 1942:34; name

combination.

Sciurus pyrrhonotus taparius: Moojen, 1952b:30; name

combination.

Sciurus pyrrhonotus juralis: Moojen, 1952b:31; name

combination.

Sciurus [(Urosciurus)] spadiceus [tricolor]: Thorington and

Hoffmann, 2005:763; name combination.

This subspecies occurs east of the Andes in Colombia

south through Ec ua dor and Peru to central Bolivia, extend-

ing to the east to Acre and Amazonas states in Brazil, and

south of the Rio Solimões along the upper course of the Rio

Juruá and Rio Purus. There is an apparent gap in the distri-

bution, as the taxon is also known from the Rio Aripuanã,

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24 Mammals of South America

ceus in color but average distinctly smaller in size. J. A. Al-

len (1914d) described these specimens as Sciurus langsdorfﬁ i

urucumus; we have placed urucumus as a ju nior synonym of

S. spadiceus because there is an overlap in size between the

largest specimens of urucumus and the smallest of spadiceus.

Genus Microsciurus J. A. Allen, 1895

Microsciurus is a genus of small tree squirrels distributed

in Central and South America. Among neotropical squir-

rels, Microsciurus species are intermediate in size between

the diminutive Sciurillus and the slightly larger Syntheosci-

urus, with mass ranging from 80– 120 g and external mea-

sure ments of head and body length of 112– 170 mm, tail

length 82– 165 mm, hindfoot length 28– 45 mm, and ear

length 8– 19 mm. The ears are small and rounded. Females

possess three pairs of evenly spaced mammae. Except for

a few taxa, the color pattern is very conservative, usually

some shade of brown ﬁ nely streaked with orange or red on

the dorsum, and with a paler ventrum. Several species have

colored postauricular patches. High- elevation populations

typically have longer and denser fur than lowland ones, and

the tail is never very bushy.

The skull has a strongly curved dorsal proﬁ le with an in-

ﬂ ection point at the middle of the frontals. From the inﬂ ec-

tion point to the anteriormost portion of skull, the rostrum

maintains a straight or even slightly concave proﬁ le. The

rostrum is short, at most one- third the length of the skull.

The frontals are longer than they are wide, and the brain-

case is globose. The temporal portion of the orbitotemporal

fossa is greatly reduced but not obliterated; the postorbital

pro cess extends posteriorly almost to a vertical plane pass-

ing through the internal (anterior) portion of the squamosal

zygomatic pro cess in the fossa. The maxillary pro cess of the

zygomatic arch is oblique in lateral view, originating ante-

rior to the cheek teeth and above the infraorbital foramen.

The sphenopalatine foramen is situated above the second

molar or the contact between the second and third molars.

There are two complete bullar septa, and rarely a third in-

complete septum. The mandible has a very short coronoid

pro cess.

The dental formula is I 1/1, C 0/0, PM 2– 1/1, M 3/3 = 20–

22; PM3 is absent in some populations, particularly those

of M. ﬂ aviventer. When present, PM3 is simple, small, uni-

cuspidate, and has one root; all remaining permanent cheek

teeth possess three roots. PM4 is molariform but varies in

size, from one- third smaller to only slightly smaller than

M1. M1 and M2 are subequal, but M3 is distinctly smaller

in size. Paraconule, metaconule, and ectostyle are absent

or vestigial; the trigon is shallow. Only pm4 has a vestigial

paraconid, a structure absent in all remaining lower cheek

teeth. The trigonid is vestigial.

these Brazilian samples also varies from being uniformly

colored from front to back or divided between a more or-

ange posterior section and a black grizzled with orange

anterior section; this phenotype has an available name, Sci-

urus pyrrhonotus juralis Thomas, with type locality on the

middle Rio Juruá, Amazonas, Brazil.

There are few samples from the east and northeast

along the Rio Amazonas, but those from the Rio Aripuanã,

in southeastern Amazonas state, as well as those from the

left bank of the Rio Tapajós, Pará state, are similar to those

from the Juruá region, except that the top of the head is

paler. As is usual for most color variants of South Ameri-

can squirrels, there is a name available for these squir-

rels, namely, Sciurus (Urosciurus) pyrrhonotus taparius

Thomas. If taken in isolation, specimens from central Peru

and from the Rio Tapajós are quite distinct from each other

and could be distinguished as valid taxa. We do not do so,

however, because we cannot presently establish clear geo-

graph i cal limits to this variation. We believe that the taxon

we are recognizing here is a complex and will probably be

split into a number of distinct, valid taxa with a deeper

study of geo graph i cal variation.

Sciurus langsdorfﬁ steinbachi J. A. Allen, described from

Santa Cruz de la Sierra, Bolivia, is identical to our concept

of H. s. spadiceus, and we treat it as a ju nior synonym. In

the southern portion of the geo graph i cal distribution of spa-

diceus, specimens from the Serra do Urucum and Cáceres,

Mato Grosso do Sul, Brazil, are identical to typical spadi-

Map 7 Selected localities for Hadrosciurus spadiceus (●). Contour line = 2,000 m.

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Suborder Sciuromorpha: Family Sciuridae 25

6. Venter with predominantly red tones . . . . . . . . . . . . . 7

6. Ventral color variable, dark gray, pale gray, or pale yel-

low on throat, and dark gray to dark brown and yellow

on chest and belly. . . . . . . . . . . . Microsciurus simonsi

7. Venter red to mixed red and gray on throat and chest,

dark gray to dark brown punctuated with red on belly

. . . . . . . . . . . . . . . . . . . . . . . . . . . Microsciurus similis

7′. Venter brown washed with red on throat and chest,

dark gray and red on belly . . . . Microsciurus sabanillae

Microsciurus ﬂ aviventer (Gray, 1867)

Amazon Dwarf Squirrel

synonyms:

Macroxus ﬂ aviventer Gray, 1867:432; type locality

“Brazil.”

Sciurus aestuans: Alston, 1878:668 part (inclusion of ﬂ avi-

ventris [= ﬂ aviventer]); not aestuans Linnaeus.

Macroxus ﬂ aviventris Alston, 1878:668; incorrect subse-

quent spelling of Macroxus ﬂ aviventer Gray.

Sciurus chrysurus: Thomas, 1893a:337; part; not chrys-

urus Pucheran.

Sciurus (Microsciurus) peruanus J. A. Allen, 1897c:115;

type locality “Guayabamba, alt. 4000 feet,” = Santa

Rosa de Huayabamba, Amazonas, Peru.

Sciurus (Microsciurus) peruanus napi Thomas, 1900f:295;

type locality “Mouth of Coca River, Upper Rio Napo,”

Orellana, Ec ua dor.

Sciurus peruanus: Ihering, 1904:420; name combination.

Microsciurus peruanus: E.- L. Trouessart, 1904:329; name

combination.

Microsciurus peruanus napi: E.- L. Trouessart, 1904:329;

name combination.

Microsciurus brevirostris J. A. Allen, 1914a:147; nomen

nudum; indicated as a new species in a list but with no

further description or reference to a description or il-

lustration, likely a lapsus calami for M. rubrirostris, de-

scribed in the same paper.

Microsciurus napi: J. A. Allen, 1914a:163; name combi-

nation.

Microsciurus rubrirostris J. A. Allen, 1914a:163; type lo-

cality “Chanchamayo, [Junín,] central Peru; altitude

2000 m.”

Microsciurus ﬂ orenciae J. A. Allen, 1914a:164; type lo-

cality “Florencia (altitude 1000 feet), Caquetá district,

Colombia.”

Microsciurus avunculus Thomas, 1914b:574; type locality

“Gualaquiza; alt. 2500′,” Morona– Santiago, Ec ua dor.

Microsciurus rubricollis Thomas, 1914b:574; incorrect sub-

sequent spelling of Microsciurus rubrirostris J. A. Allen.

Microsciurus manarius Thomas, 1920f:275; type locality

“Acajutuba, Rio Negro, near its mouth,” Amazonas,

Brazil.

synonyms:

Macroxus: Gray, 1867:432; part (description of ﬂ aviven-

ter); not Macroxus F. Cuvier.

Sciurus: J. A. Allen, 1877:750; part (inclusion of rufoni-

ger); not Sciurus Linnaeus.

[Sciurus] (Microsciurus) J. A. Allen, 1895b:332; type species

Sciurus (Microsciurus) alfari, by original designation.

Microsciurus: E.- L. Trouessart, 1897:429; elevation to ge-

neric status.

Microsciurus (Microsciurus): Moore, 1959:179; name

combination.

remarks: J. A. Allen (1895b) described Microsciurus

as a subgenus, designating Sciurus (Microsciurus) alfari

J. A. Allen as the type species. Being quite easily diagnos-

able in terms of size and a number of cranial characters

from other squirrels, the taxonomic history of Microsci-

urus is relatively unproblematic. The only major taxo-

nomic changes subsequent to Allen’s description were E.- L.

Trouessart’s (1897) elevation to full genus and Thomas’s

(1914b,f) placement of Microsciurus pusillus (sensu J. A.

Allen 1914a) in its own genus, Sciurillus. Moore (1959)

used the name in a broader sense, including several genera

and subgenera in it, but his subgenus Microsciurus is equal

to the genus Microsciurus of other authors and ourselves.

We recognize 11 species of Microsciurus, three in Cen-

tral America (M. alfari, M. boquetensis, and M. venustulus)

and eight in South America (M. ﬂ aviventer, M. isthmius,

M. mimulus, M. otinus, M. sabanillae, M. santanderensis,

M. similis, and M. simonsi). Only the South American spe-

cies are detailed herein. Both Hall (1981) and Thorington

et al. (2012) extended the range of boquetensis into extreme

northwestern Colombia (as a subspecies of M. mimulus).

KEY TO THE SOUTH AMERICAN SPECIES OF

MICROSCIURUS:

1. Postauricular patch present . . . . . . . . . . . . . . . . . . . . 2

1′. Postauricular patch absent . . . . . . . . . . . . . . . . . . . . . 3

2. Dorsum brown, streaked with red . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . Microsciurus ﬂ aviventer

2′. Dorsum dark brown, with mid- dorsal dark stripe . . . .

. . . . . . . . . . . . . . . . . . . . . Microsciurus santanderensis

3. Dorsum dark brown streaked with red to reddish or-

ange tones . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4

3′. Dorsum dark brown streaked with yellow to pale yel-

low tones . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .5

4. Venter orange, mixed with brown on belly . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . Microsciurus isthmius

4′. Venter washed with red on throat and chest; mixed brown,

dark gray, and red on belly . . . . . . . .Microsciurus otinus

5. Basal third of hairs of venter gray. . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . .Microsciurus mimulus

5′. Hairs of ventrum without gray basal third . . . . . . . . . 6

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26 Mammals of South America

71606), Hacienda Santa Elena, ca. 35 km NE of Tingo Ma-

ria (LSUMZ 17720), left bank at mouth of Río Yaquerana,

alto Río Yavarí (FMNH 88991), Pebas. Río Amazonas (BM

28.7.21.74– 78), Quebrada Pushaga, left bank Río Mo-

rona, alto Río Amazonas (FMNH 88984), San Fernando,

left bank Río Yavari (FMNH 88990), Santa Rita, Iquitos

(FMNH 87178), Sarayacu, Río Ucayali (AMNH 75279);

Madre de Dios, Altamira, Manu (FMNH 98067); Ucayali,

Lagarto, upper Río Ucayali (AMNH 76538).

subspecies: We regard M. ﬂ aviventer as monotypic

(but see Remarks). Thorington and Hoffmann (2005) rec-

ognized seven subspecies, and Thorington et al. (2012)

listed eight, three and four of which, respectively, we re-

gard herein as full species.

natural history: This species is found only in ev-

ergreen rainforest (Emmons and Feer 1997). Surprisingly,

few ecological, reproductive, behavioral, or other natural

history data are available for this wide- ranging squirrel.

Emmons and Feer (1997) stated that individuals search ac-

tively over the trunks of large trees, on vines, and in treef-

alls, apparently searching for insects, but that they also feed

on substances scraped from tree bark such as gum (as was

suggested by A. C. Smith [1999], who observed two soli-

tary individuals moving through the subcanopy). Youlatos

(1999) noted that individuals primarily used the subcanopy

or canopy at heights above 5 m. Emmons and Feer (1997)

recorded ball nests of leaves lined with ﬁ bers in the top of

palms. Eisenberg (1989) reported a litter size of two off-

spring. Buitron- Jurado and Tobar (2007) reported on asso-

ciations between individuals of this species and bird ﬂ ocks,

with squirrels actively searching for insects at heights of 2–

10 m in the company of birds.

remarks: Microsciurus ﬂ aviventer has a wide dis-

tribution, from the eastern slopes of the Andes to central

Amazonia. In this vast area, authors have distinguished sev-

eral taxa, all listed in the previous synonymy. Our study

of material deposited in many collections revealed that this

variation is real, but we have not been able to decide how

much of it is of taxonomic value. There are important dif-

ferences in size (Andean forms such as napi are large, for

example). And the upper third premolar may be present

or absent in different samples. The fur is also longer and

denser in Andean samples, and the color, especially that of

ventral surfaces, varies regionally. We believe that recogniz-

ing a single species for the whole of the samples examined

is, in this case, a decision of con ve nience. It is quite possible

that further study of this material will reveal the existence

of additional taxa, either to be recognized as valid subspe-

cies or as full species.

Microsciurus ﬂ aviventer is one of the two Colombian spe-

cies of the genus occurring east of the Andes, the other being

M. santanderensis. Their known distributions are interrupted

Microsciurus ﬂ aviventer: Thomas, 1920f:275; ﬁ rst use of

current name combination.

Microsciurus mannarius: O. M. O. Pinto, 1931:284; inad-

vertent subsequent spelling of Microsciurus manarius

Thomas.

Microsciurus [(Microsciurus)] napi: Moore, 1959:203;

name combination.

Microsciurus ﬂ aviventer ﬂ aviventer: Cabrera, 1961:355;

name combination.

Microsciurus ﬂ aviventer napi: Cabrera, 1961:356; name

combination.

Microsciurus ﬂ aviventer peruanus: Cabrera, 1961:357;

name combination.

Microsciurus ﬂ aviventer rubrirostris: Cabrera, 1961:357;

name combination.

description: Adult mass 80– 111 g (N = 16); head

and body length 124– 170 mm (N = 76), tail length 115–

165 mm (N = 79), hindfoot length 30– 44 mm (N = 76),

and ear length 13– 19 mm (N = 65). Dorsal color brown

streaked with red; most samples have yellowish to white

postauricular patches; venter of mixed colors, with throat

and chest yellowish orange and belly slightly darker.

distribution: Microsciurus ﬂ aviventer occurs widely

in western Amazonia, from southern Colombia through east-

ern Ec ua

dor, Peru, and northeastern Bolivia, then eastward

to the Brazilian states of Amazonas, Acre, and Rondônia. It

ranges in elevation from 100 to 1,500 m.

selected localities (Map 8): BOLIVIA: Pando,

left bank Río Madre de Dios near Santa Elena (Salazar-

Bravo, Yensen et al. 2002). BRAZIL: Acre, Oriente (=

Seringal Oriente), near Taumaturgo, Rio Juruá (MZUSP

20351); Amazonas, Acajutuba, Rio Negro (type locality

of M. manarius Thomas), Codajás, left bank Rio Solimões

(MZUSP 4207), Eirunepé, left bank Rio Juruá (MZUSP

4621), Estirão do Equador, right bank Rio Javari (MZUSP

20350), Lago do Mapixi, east of Rio Purus (FMNH 38880),

Manacapuru, Rio Solimões (BM 26.5.5.36), Paraná do

Aiapuá, Rio Purus (BM 27.8.11.61); Rondônia, Cachoeira

Nazaré, left bank Rio Ji- Paraná (MPEG 20826). COLOM-

BIA: Caquetá, Florencia (type locality of M. ﬂ orenciae J.

A. Allen); Putumayo, La Tagua, Tres Troncos, Río Caquetá

(FMNH 71118); Vaupés, Río Vaupés, in front of Tahua-

punto (AMNH 78625). EC UA DOR: Loja, El Porotillo,

San José (MNHN 1934/1210); Morona– Santiago, Men-

dez Sur, Oriente (MNHN 1987/338); Napo, Archidona

(MNHN 1932/2876), San José abajo (AMNH 66792).

PERU: Amazonas, Guayabamba (AMNH type locality

of Sciurus (Microsciurus) peruanus J. A. Allen), near

Huampani, Río Cenepa (MVZ 154929), 12 km E of La

Peca Nueva (LSUMZ 21876); Cusco, Quincemil (FMNH

75210); Junin, Chanchamayo (type locality of Microsciurus

rubrirostris J. A. Allen); Loreto, boca Río Curaray (AMNH

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Suborder Sciuromorpha: Family Sciuridae 27

Microsciurus [(Microsciurus)] isthmius: Moore, 1959:203;

name combination.

Microsciurus ﬂ aviventer isthmius: Cabrera, 1961:355;

part; name combination.

Microsciurus mimulus isthmius: Handley, 1966:778; part;

name combination.

description: Adult mass 120 g (one female); head

and body length 120– 165 mm (N = 41), tail length 98–

150 mm (N = 36), hindfoot length 32– 45 mm (N = 39), and

ear length 10– 19 mm (N = 20). Dorsum dark brown to black,

heavily speckled with yellow; postauricular patches absent;

venter not sharply distinct from dorsum, generally orange in

color but with mixture of orange and brown on belly.

distribution: Microsciurus isthmius occurs in rain-

forests west of the Andes in northwestern Colombia and ad-

jacent Panama; in Colombia, it ranges in elevation from sea

level to 800 m.

selected localities (Map 9; South America

only): COLOMBIA: Antioquia, Alto Bonito, upper Río

Sucio (AMNH 37645), Murindó (FMNH 44855); Cauca,

Río Saija (FMNH 90210); Chocó, Bagadó (AMNH

34132), Baudó (AMNH 33812), Nóvita, Río Tamaná

(BM 9.7.17.26), Río Nuqui, base of Baudó Mts. (USNM

292132), Río Truandó (type locality of Sciurus [Microsci-

urus] isthmius E. W. Nelson), Sipí, Río Sipí, Río San Juan

(type locality of Sciurus [Microsciurus] palmeri Thomas);

Nariño, Buenavista (AMNH 34158), La Guayacana

(FMNH 89518); Valle Del Cauca, Mechenquito (ROM

63217), Río Mechengue (FMNH 90201), San José (AMNH

31680).

subspecies: We treat M. isthmius as monotypic.

natural history: Goldman (1920: 144) reported

specimens from about 1,150 m on Cerro Pirre, Darién,

Panama (his M. isthmius vivatus), collected from lower

branches or the trunks of trees in cloud forest.

remarks: Variation in the coat color is limited to

the amount of black on the crown and rest of dorsum,

which is slightly more pronounced in some specimens. J. A.

Allen (1877:756– 763) recognized ﬁ ve species of squirrels

in South America, among which was Sciurus aestuans

and in which he listed “aestuans” and “rufoniger” as two

“varieties.” These together encompassed all of the then-

known smaller South American squirrels. This was a very

broad concept, and even Allen himself (1914a, 1915) did

not succeed in settling all the synonymic implications of

his earlier lumping effort. E. W. Nelson (1899a:77– 78)

subsequently described Sciurus (Microsciurus) isthmius

based on the material collected at the Río Truandó, Chocó,

Colombia.

Cabrera (1961) proposed a new arrangement for the

species level taxa in the genus Microsciurus, one in which

isthmius E. W. Nelson and all other South American taxa

by a large gap in the eastern Andes, a region well sampled for

other squirrels, and this gap likely represents a true disjunc-

tion. Salazar- Bravo, Yensen et al. (2002) extended the range

of this species into northern Bolivia, reidentifying a specimen

that S. Anderson (1997:397) originally assigned to Sciurus

ignitus (= Notosciurus pucheranii herein).

Microsciurus isthmius (E. W. Nelson, 1899)

Isthmian Dwarf Squirrel

synonyms:

(?) Sciurus aestuans var. rufoniger: J. A. Allen, 1877:757;

part.

Sciurus (Microsciurus) isthmius E. W. Nelson, 1899a:77;

type locality “Truando River [= Río Truandó], [Chocó,]

Colombia.”

Microsciurus isthmius: E.- L. Trouessart, 1904:329; name

combination.

Sciurus (Microsciurus) palmeri Thomas, 1909:234; type

locality “Sipi, Rio Sipi, Rio San Juan, Chocó, western

Colombia.”

Microsciurus isthmius vivatus Goldman, 1912b:4; type

locality “near Cana, (altitude 3,500 feet), in the Pirri

range of mountains, [Darien,] eastern Panama.”

Microsciurus isthmius isthmius: J. A. Allen, 1914a:57;

name combination.

Microsciurus palmeri: J. A. Allen, 1914a:160; name

combination.

Map 8 Selected localities for Microsciurus ﬂ aviventer (●). Contour line = 2,000 m.

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28 Mammals of South America

patches absent; venter not sharply distinct from dorsum,

with color ranging from orange to pale yellow with basal

third of hairs gray, with belly orange to pale yellow with

some brown.

distribution: Microsciurus mimulus occurs in

rainforests west of the Andes in northwestern Ec ua dor

and southwestern Colombia, at elevations from sea level

to 1,000 m.

selected localities (Map 10): COLOMBIA:

Nariño, Barbacoas (AMNH 34159), La Guayacana

(USNM 309028). EC UA DOR: Esmeraldas, Cachabí (type

locality of Sciurus [Microsciurus] mimulus Thomas),

Carondelet (USNM 113310); Imbabura, Lita (BM

1.1.6.3).

subspecies: Microsciurus mimulus is monotypic.

natural history: We have found no information

about the natural history of this species.

remarks: Thomas (1898f:266– 267) described Sci-

urus (Microsciurus) mimulus based on three specimens

from northwestern Ec ua dor. This was the ﬁ rst name ap-

plied to dwarf squirrels from South America west of the

Andes. Cabrera (1961:356) considered M. ﬂ aviventer to

be the only South American species in the genus, using

all other names either for subspecies (including mimulus

Thomas) or as synonyms. As in many occasions through-

out his cata log of South American mammals, this decision

was based on the literature and on his own (limited, in this

case) experience with the group. Handley (1966:777) con-

sidered M. mimulus to be a good species, and although he

did not provide a reason for this decision, his arrangement

were placed as subspecies, or synonyms, of a single species,

namely, M. ﬂ aviventer (Gray). Cabrera’s arrangement, how-

ever, was not uniformly followed (e.g., see Handley 1966;

Hall 1981; and Honacki et al. 1982). Thorington et al.

(2012) retained isthmius as a subspecies in their concept of

M. mimulus.

Microsciurus mimulus (Thomas, 1898)

Western Dwarf Squirrel

synonyms:

Sciurus (Microsciurus) mimulus Thomas, 1898f:266; type

locality “Cachavi [= Cachabí], [Esmeraldas,] N. Ec ua dor,

alt. 167 m.”

Microsciurus mimulus: E.- L. Trouessart, 1904:329; ﬁ rst

use of current name combination.

Microsciurus [(Microsciurus)] mimulus: Moore, 1959:203;

name combination.

Microsciurus ﬂ aviventer mimulus: Cabrera, 1961:356;

name combination.

description: Head and body length 114– 159 mm

(N = 17), tail length 90– 125 mm (N = 17), hindfoot length

30– 38 mm (N = 16), and ear length 13– 16 mm (N = 8).

Dorsum dark brown to black, speckled with red or or-

ange, sometimes with mid- dorsal dark stripe; postauricular

Map 9 Selected localities for Microsciurus isthmius (●). Range in Panama from

Hall (1981). Contour line = 2,000 m.

Map 10 Selected localities for Microsciurus mimulus (●). Contour line = 2,000 m.

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Suborder Sciuromorpha: Family Sciuridae 29

was on the Río Porce in the Cordillera Central of Antio-

quia department, Colombia, the taxon they studied was

likely M. otinus, not M. mimulus as they originally re-

ported. Otherwise, little information is available on this

species.

remarks: Thomas (1901c) contrasted M. otinus with

the geo graph i cally adjacent M. isthmius, noting that the

two were similar in all external aspects except that otinus

had white pinnae. Although we recognize M. otinus here

as a valid species, this scheme needs to be tested by ad-

ditional collecting and comparisons of existing specimens.

Thorington et al. (2012) included otinus as a subspecies of

M. ﬂ aviventer.

Microsciurus sabanillae Anthony, 1922

Sabanilla Dwarf Squirrel

synonyms:

Microsciurus sabanillae Anthony, 1922: 2, type local-

ity “Sabanilla, Prov. de Loja, Ec ua dor; altitude 5700

feet.”

Microsciurus ﬂ aviventer sabanillae: Cabrera, 1961:357;

name combination.

description: Head and body length 134– 151 mm

(N = 3), tail length 140– 148 mm (N = 3), and hindfoot

met with wide ac cep tance and has been adopted by most

subsequent authors (e.g., Emmons and Feer 1997; Thor-

ington and Hoffmann 2005). Our concept of M. mimu-

lus is more restricted geo graph i cally than that espoused by

these authors, generally equivalent to that given by Thor-

ington et al (2012:35) for the nominotypical form of their

polytypic M. mimulus.

Some specimens from Nariño, Colombia, show indi-

vidual variation in belly color, from almost black punctu-

ated with orange to brown mixed with orange. The type

series of M. mimulus and other specimens from north-

western Ec ua dor have a black mid- dorsal stripe continu-

ing to the crown (Thomas 1898f; J. A. Allen 1914a). This

character, however, while present in Ec ua dorean mate-

rial, is variable in specimens from farther to the north in

Nariño, Colombia, with only one specimen in ﬁ ve pre-

senting it fully.

Microsciurus otinus (Thomas, 1901)

White- eared Dwarf Squirrel

synonyms:

Sciurus (Microsciurus) otinus Thomas, 1901c:193; type lo-

cality “Medellin, [Antioquia,] Colombia.”

Microsciurus otinus: E.- L. Trouessart, 1904:329; ﬁ rst use

of current name combination.

Microsciurus ﬂ aviventer otinus: Cabrera, 1961:357; name

combination.

description: Head and body length 120– 141 mm

(N = 15), tail length 95– 132 mm (N = 12), hindfoot length

35– 39 mm (N = 13), and ear length 11– 16 mm (N = 6). Dor-

sum dark brown streaked with pale yellow; postauricular

patches absent but ears tipped with white; throat and chest

washed with reddish orange, belly mixed brown, dark

gray, and red; tail frosted whitish.

distribution: Microsciurus otinus occurs west of

the Andes, in rainforests in the Colombian departments of

Antioquia, Bolívar, and Córdoba, at elevations from 200

to 1,500 m.

selected localities (Map 11): COLOMBIA:

Antioquia, La Tirana, 25 km S and 22 km W of Zaragoza

(USNM 499515), Medellín (type locality of Sciurus [Mi-

crosciurus] otinus Thomas), Purí, above Caceres (FMNH

70046), Valdívia, right bank of lower Río Cauca (AMNH

37640); Bolívar, 15 mi W of Simiti, 6 mi above Santa

Rosa (USNM 282755); Córdoba, Socorré, upper Río Sinú

(FMNH 69034).

subspecies: Microsciurus otinus is monotypic.

natural history: Rojas- Robles et al. (2008) re-

ported on the importance of a species of Microsciurus in

the short- distance dispersal (typically <50 m) of the large

seeds of Oenocarpus bataua, a subcanopy and canopy

palm in premontane humid forest. Because their study site

Map 11 Selected localities for Microsciurus otinus (●). Contour line = 2,000 m.

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30 Mammals of South America

Microsciurus santanderensis

(Hernández- Camacho, 1957)

Santander Dwarf Squirrel

synonyms:

Sciurus pucheranii santanderensis Hernández- Camacho,

1957: 219, type locality “Meseta de los Caballeros,

5 km NW of La Albania, San Vicente de Chururí [= San

Vicente de Chucurí], Santander, Colombia.”

Microsciurus santanderensis: Hernández- Camacho, 1960:

360; ﬁ rst use of current name combination.

description: (The following summarized from the

original description, which was based on ﬁ ve specimens;

no specimens seen by us.) Head and body length 133–

160 mm, tail length 136– 152 mm, hindfoot length 42–

45 mm, and ear length 15– 18 mm. Dorsum dark brown,

with mid- dorsal black stripe; pale ochraceous postauricu-

lar patches present; venter with mixed colors, but predomi-

nantly light ochraceous salmon.

distribution: This species occurs in the rainforests

on the western slopes of the eastern Cordillera, and from

the right bank of the middle Río Magdalena, in the vicinity

of Barrancabermeja and San Vicente de Chucurí, Santander,

Colombia. Thorington et al. (2012) extended the range west

and northwest to include the upper portion of the Cordillera

Occidental, but did not cite speciﬁ c localities.

selected localities (Map 13): COLOMBIA:

Santander, Barancabermeja (Hernández- Camacho 1960),

Meseta de Los Caballeros, 5 km NW of La Albania, San

Vicente de Chucurí (type locality of Sciurus pucheranii

santanderensis Hernández- Camacho).

subspecies: Microsciurus santanderensis is

monotypic.

natural history: Little is known about the ecol ogy

of this species, other than its distribution in humid forests in

the low to mid elevations in the Cordillera de La Paz, Colom-

bia (Hernández- Camacho 1960). Thorington et al. (2012)

included marshlands and montane forests (at an elevational

range of 2,700– 3,800 m) among habitats this species occupies.

remarks: Variation in this species has not been

studied beyond what Hernández- Camacho (1957, 1960)

presented in his original description and subsequent rede-

scription. It appears to be a valid species because it dif-

fers from similarly colored squirrels occurring east of the

Andes. For example, M. santanderensis differs from M.

ﬂ aviventer by the possession of a mid- dorsal dark stripe

and from M. sabanillae by its possession of postauricular

patches. Although originally described as a subspecies of

Notosciurus pucheranii (as Sciurus pucheranii santand-

erensis) by Hernández- Camacho, he (1960) subsequently

transferred santanderensis to Microsciurus, as more speci-

mens became available.

length 40– 42 mm (N = 3). Dorsum dark brown to black

streaked with red; postauricular patches absent; throat

and chest washed with red, belly mixed dark gray and

red.

distribution: Microsciurus sabanillae is known

from Ec ua dor and adjacent northwestern Peru, in

midelevation forests on the eastern Andean slope, at an el-

evational range from 600 to 1,750 m.

selected localities (Map 12): EC UA DOR:

Loja, Sabanilla, near Río Destrozo, a small tributary of Río

Zamora (type locality of Microsciurus sabanillae Anthony);

Napo, Río Napo (BM 54.406). PERU: Amazonas, Santa

Rosa (AMNH 69237).

subspecies: Microsciurus sabanillae is monotypic.

natural history: Nothing has been published on

the ecol ogy of this species.

remarks: Contra Thorington et al. (2012), we recog-

nize M. sabanillae as a species distinct from M. ﬂ aviventer

on the basis of details of ventral coloration, the lack of the

postauricular patch, and partial sympatry. However, as in

the case of M. otinus, this kind of variation has to be con-

sidered in the context of the highly variable M. ﬂ aviventer.

The validity of supposed species- level taxa such as M. sa-

banillae, M. otinus, and several other forms here treated as

synonyms of M. ﬂ aviventer requires additional study and

further collecting efforts.

Map 12 Selected localities for Microsciurus sabanillae (●). Contour

line = 2,000 m.

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Suborder Sciuromorpha: Family Sciuridae 31

Microsciurus [(Microsciurus)] similis: Moore, 1959:203;

name combination.

Microsciurus ﬂ aviventer similis: Cabrera, 1961:357; name

combination.

Microsciurus alfari fusculus: Handley, 1966:777; name

combination.

description: Head and body length 112– 147 mm

(N = 23), tail length 82– 133 mm (N = 17), hindfoot length

28– 38 mm (N = 21), and ear length 8– 16 mm (N = 17).

Dorsum dark brown to black punctuated with red; postau-

ricular patches absent; venter not sharply distinct from the

dorsum, colored red to mixed red and dark gray on throat

and chest but dark gray to dark brown heavily streaked

with red on belly.

distribution: Microsciurus similis occurs in rain-

forests west of the Andes in western Colombia, across an

elevational range from 130 to 2,200 m.

selected localities (Map 14): COLOMBIA:

Antioquia, Alto Bonito, upper Río Sucio (AMNH 37643);

Cauca, Munchique (FMNH 86788), Rio Munchique

(FMNH 90200); Chocó, Baudó (AMNH 33184, 33186),

Juntas de Tamaná (type locality of Sciurus [Microsci-

urus] similis fusculus Thomas), Río Jurubidá, Baudó Mts.

The remarkable distribution gap between this species and

M. ﬂ aviventer, which happens to be in a region of Colom-

bia well sampled for squirrels in general, and this species’ re-

semblance to Notosciurus pucheranii, may indicate that M.

santanderensis does not belong in the genus Microsciurus.

Because we have not seen specimens (and, so far as we know,

neither has anyone but the author of the name), we keep it

among the dwarf squirrels, but with this cautionary note.

Microsciurus similis (E. W. Nelson, 1899)

Chocó Dwarf Squirrel

synonyms:

Sciurus (Microsciurus) similis E. W. Nelson, 1899a:78;

type locality “near Cali, Cauca Valley, [Valle de Cauca,]

Colombia (alt. 6000 ft.)”.

Microsciurus similis: E.- L. Trouessart, 1904:329; ﬁ rst use

of current name combination.

Sciurus ( Microsciurus) similis fusculus Thomas, 1910d:503;

type locality “Juntas, Rio San Juan. Alt. 400′, Chocó,

W. Colombia.”

Microsciurus similis similis: J. A. Allen, 1914a:153; name

combination.

Microsciurus similis fusculus: J. A. Allen, 1914a:154; name

combination.

Map 13 Selected localities for Microsciurus santanderensis (●). Contour

line = 2,000 m.

Map 14 Selected localities for Microsciurus similis in South America (●).

Contour line = 2,000 m.

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32 Mammals of South America

cluded simonsi as a subspecies of a highly polytypic M.

ﬂ aviventer.

Genus Notosciurus J. A. Allen, 1914

Notosciurus is a genus of medium- sized Central and South

American squirrels occurring from eastern Costa Rica

through Colombia, Venezuela, Ec ua dor, Peru, western Am-

azonian Brazil, and Bolivia, to northernmost Argentina. In

South America, its distribution is mostly associated with

the Andean Cordillera, but in western Colombia, eastern

Peru, and neighboring areas of Brazil, it is also found in

lowland evergreen forests. The distribution of the genus is

largely parapatric to that of Guerlinguetus, but both gen-

era may be found living together at the margins of their

ranges, particularly in the Marañón, Solimões, and Ma-

deira river basins of Amazonia and possibly also in north-

western Mato Grosso state in Brazil.

Species of Notosciurus can be distinguished from all

other South American squirrels except for Guerlinguetus

by their size (larger than species of Microsciurus, Sciurillus,

and Syntheosciurus, but smaller than those of Hadrosci-

urus and Simosciurus). Compared with the similarly sized

Guerlinguetus, Notosciurus possess three pairs of mammae

instead of four and typically have orange postauricular

patches, which are generally lacking in Guerlinguetus. No-

tosciurus is highly variable in size and mass, varying from

136– 418.5 g (N = 31 adults). The head and body length range

from 151– 262 mm (N = 469), tail length from 134.5– 265 mm

(N = 459), hindfoot length from 34– 65 mm (N = 455), and

(USNM 292134), Río San Juan (BM 10.7.16.1); Nariño,

Ricaurte (USNM 309027); Risaralda, Siató, Rio Siató, San

Juan (FMNH 18877); Valle del Cauca, Albán (= Salencio)

(AMNH 32852), Cali (BM 99.9.6.56).

subspecies: We treat M. similis as monotypic, al-

though Thomas (1910d) differentiated specimens from

Chocó in western Colombia as M. s. fusculus.

natural history: We found no information about

the natural history of this species.

remarks: Microsciurus similis, on the scale of Maps 9

and 14, appears to broadly overlap the range of M. isthmius.

The two species, however, occupy separate elevational ranges

in the Colombian Andes, with limited overlap. The lowest

elevation recorded for M. similis is 150 m, and the highest for

M. isthmius is 800 m. This species is also sympatric with M.

mimulus in Nariño department in southwestern Colombia.

Microsciurus simonsi (Thomas, 1900)

Simons’s Dwarf Squirrel

synonyms:

Sciurus (Microsciurus) simonsi Thomas, 1900f:294; type

locality “Porvenir, near Zaparal, Province of Bolivar,

Ec ua dor. Altitude 1500 m.”

Microsciurus simonsi: E.- L. Trouessart, 1904:329; ﬁ rst use

of current name combination.

Microsciurus similis fusculus: Lönnberg, 1913:26; not fus-

culus Thomas.

Microsciurus ﬂ aviventer simonsi: Cabrera, 1961:358; name

combination.

description: Head and body length 126– 154 mm

(N = 10), tail length 104– 124 mm (N = 10), hindfoot length

32– 37 mm (N = 11), and ear length 14– 16 mm (N = 11).

Dorsum dark brown streaked with orange; postauricular

patches absent; ventrum mixture of dark gray, pale gray,

pale yellow on throat, dark gray to dark brown and yellow

on chest and belly.

distribution: Microsciurus simonsi occurs west of

the Andes in the Ec ua dorean provinces of Bolívar and Pi-

chincha, at elevations from 250 to 1,500 m.

selected localities (Map 15): EC UA DOR:

Bolívar, Porvenir (type locality of Sciurus [Microsciurus] si-

monsi Thomas); Pichincha, Ila, 36 mi SW of Santo Domingo

(BM 15.1.1.25), Mindo, Río Blanco (BM 13.10.24.36),

Santo Domingo (BM 15.1.1.23– 24).

subspecies: Microsciurus simonsi is monotypic.

natural history: Nothing has been published on

the ecol ogy or behavior of this species.

remarks: Microsciurus simonsi is the southernmost

species of the genus west of the Andes. Each of the four

trans- Andean species can be distinguished mainly by

its ventral coloration and the color of the subterminal

band of dorsal guard hairs. Thorington et al. (2012) in-

Map 15 Selected localities for Microsciurus simonsi (●). Contour line = 2,000 m.

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Suborder Sciuromorpha: Family Sciuridae 33

South American species until J. A. Allen (1915a) created

new genera to accommodate the several groups of appar-

ently related species he was able to recognize. Among the

new genera were Leptosciurus (type species Sciurus rufo-

niger Pucheran, 1845) and Mesosciurus (type species Sci-

urus aestuan var. hoffmanni Peters, 1864), the latter with

two subgenera: the nominotypical one and Histriosciurus

(type species Sciurus gerrardi Gray). Allen did not com-

pare Leptosciurus and Mesosciurus with each other, ex-

cept in his key for the genera; in his characterizations, he

chose to compare both only to Guerlinguetus, and quite

superﬁ cially. Furthermore, he distinguished between Lep-

tosciurus and Mesosciurus only on the basis of size, which

overlapped slightly in his own key (J. A. Allen 1915a:187).

However, a year earlier, J. A. Allen (1914d) had de-

scribed a new genus and species based on a single specimen

from Ec ua dor, which he named Notosciurus rhoadsi. We

have not seen the holotype of this taxon, but Hershkovitz

(1947:8) identiﬁ ed it as a young specimen of Sciurus hoff-

manni soederstroemi Stone. Thorington and Hoffmann

(2005:761) also listed Allen’s rhoadsi as a synonym of so-

ederstroemi Stone, which they regarded as a subspecies of

a polytypic Sciurus granatensis Humboldt. These authors

also included chrysuros Pucheran and hoffmanni Peters as

additional subspecies of S. granatensis. Because we con-

sider soederstroemi Stone to be a synonym of chrysuros

Pucheran and we treat granatensis Humboldt as a species

in the genus we discuss here, Notosciurus J. A. Allen takes

priority over Leptosciurus J. A. Allen as the generic name

for these squirrels.

We do not recognize subgenera in Notosciurus, as all

species share the presence of three pairs of mammae; addi-

tionally, most of the included species and subspecies possess

postauricular patches, both characteristics that distinguish

our concept of Notosciurus from Guerlinguetus. These

two genera, as here recognized, are largely parapatric, with

marginal sympatry in parts of western Brazil and eastern

Peru. See the discussion on the validity of the presently rec-

ognized supraspeciﬁ c groups in the family account.

KEY TO THE SPECIES OF NOTOSCIURUS:

1. Size smaller, head and body length approximately 150–

210 mm; ventral hairs of various colors but usually with

a gray base (except in Bolivia and Argentina) . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . .Notosciurus pucheranii

1′. Size larger (but overlapping; in eastern Andean Colom-

bia, where both species occur in sympatry [N. g. chrys-

uros and N. p. pucheranii], they diverge greatly in size),

head and body length approximately 170– 265 mm; ven-

tral hairs of various colors but usually of a single color

throughout (species does not occur in Bolivia or Argen-

tina) . . . . . . . . . . . . . . . . . . . . Notosciurus granatensis

ear length from 15– 32 mm (N = 359). The skull has the orbi-

totemporal fossa equally divided by a well- developed postor-

bital pro cess, and an upper second molar with two parallel,

buccolingual crests. These crests are obliquely oriented rela-

tive to the anteroposterior axis of the skull.

We recognize two polytypic species: Notosciurus grana-

tensis with six subspecies (ﬁ ve in South America), and N.

pucheranii with four subspecies. Our concept of both spe-

cies is considerably different from that of previous authors,

most recently Thorington et al. (2012). These authorities,

for example, recognized 32 subspecies for granatensis, 29 in

South America. They also treated the trans- and cis- Andean

populations of our N. pucheranii as separate species.

synonyms:

Sciurus: Pucheran, 1845:336; part (description of rufoni-

ger and chrysurus); not Sciurus Linnaeus, 1758.

Funambulus: Fitzinger, 1867a:487; part (listing of pucher-

anii and chrysurus); not Funambulus Lesson, 1835.

Macroxus: Gray, 1867:429; part (description of ignitus);

not Macroxus F. Cuvier, 1823.

Sciurus (Guerlinguetus): J. A. Allen, 1899b:217; part (de-

scription of quebradensis); not Guerlinguetus Gray, 1821.

Notosciurus J. A. Allen, 1914d:585; type species Notosci-

urus rhoadsi J. A. Allen, 1914, by original designation.

Guerlinguetus: J. A. Allen, 1914d:587; part (description

of pucheranii salentensis, hoffmanni quindianus, hoff-

manni manavi, and griseimembra); not Guerlinguetus

Gray, 1821.

Leptosciurus J. A. Allen, 1915a:199; type species Sciurus

rufoniger Pucheran, 1845 (preoccupied by Sciurus rufo-

nigra Gray, 1842:263, replaced by Funambulus pucher-

anii Fitzinger, 1867).

Mesosciurus J. A. Allen, 1915a:212; type species Sci-

urus aestuans var. hoffmanni Peters, 1864, by original

designation.

Histriosciurus J. A. Allen, 1915a:213, 236; as subgenus;

type species Sciurus gerrardi Gray, 1861, by original

designation (p. 236).

Mesosciurus (Mesosciurus): J. A. Allen; 1915a:216; name

combination.

Guerlinguetus (Mesosciurus): J. A. Allen; 1915a:228; name

combination, but lapsus calami.

Mesosciurus (Histriosciurus): J. A. Allen; 1915a:236; name

combination.

Sciurus (Mesosciurus): Thomas, 1921k:609; name

combination.

Sciurus (Leptosciurus): Anthony, 1923a:5; name

combination.

Sciurus [(Notosciurus)]: Ellerman, 1940:344; name

combination.

remarks: The midsized South American squirrels

with three pairs of mammae were or ga nized with all other

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34 Mammals of South America

and Boher); Carabobo, Urama, E of Yaracuy (AMNH

130289); Guárico, Guárico (AMNH 32877); Mérida, Es-

corial, Sierra de Mérida (type locality of Sciurus griseogena

meridensis Thomas); Monagas, near San Agustin, 2 km N

and 4 km E of Caripe (USNM 409776); Nueva Esparta, El

Valle, Isla de Margarita (type locality of Sciurus nesaeus

G. M. Allen); Sucre, Cristobal Colón (AMNH 36156), Ne-

vera (AMNH 69911); Vargas, Galiparé, Cerro del Ávila,

near Caracas (type locality of Sciurus griseogena klagesi

Thomas); Zulia, El Panorama, Río Aurare (type locality of

Sciurus granatensis maracaibensis Hershkovitz), Hacienda

Platanal, 18 km N and 49 km W of Maracaibo (USNM

442017), Novita, 3.1 km S and 19 km W of Machiquos

(USNM 442013).

subspecies: We recognize six subspecies of N. grana-

tensis, with ﬁ ve in South America. A sixth subspecies, N. g.

hoffmanni, occurs only in Costa Rica and western Panama.

For comparison with South American taxa, specimens of

hoffmanni possess postauricular patches, a reddish dorsum

streaked with orange or yellow but without a dark mid-

dorsal area, an orange venter with gray- based hairs, and a

homogeneously colored tail fringed and/or streaked with

orange.

N. g. chapmani (J. A. Allen, 1899)

synonyms:

Sciurus chapmani J. A. Allen, 1899a:16, type locality

“Caparo, Trinidad,” Trinidad and Tobago.

Sciurus (Guerlinguetus) aestuans quebradensis J. A. Allen,

1899b:217, type locality “Quebrada Secca [= Quebrada

Seca, = Villarroel], [Sucre,] Venezuela.”

Sciurus (Guerlinguetus) chapmanni E.- L. Trouessart, 1904:

327; name combination, and incorrect subsequent spell-

ing of Sciurus chapmani J. A. Allen.

Sciurus tobagensis Osgood, 1910:27, type locality “To-

bago Island, Ca rib be an Sea,” Trinidad and Tobago.

Mesosciurus (Mesosciurus) chapmani: J. A. Allen, 1915a:

230; name combination.

Mesosciurus (Mesosciurus) chapmani tobagensis: J. A. Allen,

1915a:232; name combination.

Sciurus [(Guerlinguetus)] chapmani chapmani: Ellerman,

1940:341; name combination.

Sciurus [(Guerlinguetus)] chapmani tobagensis: Ellerman,

1940:341; name combination.

Sciurus granatensis chapmani: Hershkovitz, 1947:37; name

combination.

Sciurus granatensis tobagensis: Hershkovitz, 1947:37; name

combination.

Sciurus (Guerlinguetus) granatensis chapmani: Goodwin,

1953:277; name combination.

Sciurus (Guerlinguetus) granatensis: Eisenberg, 1989:331;

part; name combination.

Notosciurus granatensis (Humboldt, 1811)

Red- tailed Squirrel

synonyms: See under subspecies.

description: Highly variable species, with head and

body length 169– 262 mm (N = 348 adult specimens), tail

length 145– 265 mm (N = 342), hindfoot length 40– 65 mm

(N = 330), ear length 17– 32 mm (N = 250), and mass 190–

420 g (N = 25). We brieﬂ y describe each subspecies follow-

ing its synonymy.

distribution: This species ranges from Costa Rica

to Colombia, Ec ua dor, and Venezuela. In South America, N.

granatensis is essentially linked to the Andes, but may also

occur in adjacent lowlands and coastal areas at sea level.

selected localities (Map 16): COLOMBIA:

Antioquia, Puerto Valdivia (type locality of Mesosciurus

(Histriosciurus) gerrardi valdiviae J. A. Allen), Sonsón

(USNM 239942); Bolívar, Cartagena (type locality of Sci-

urus granatensis Humboldt); Cauca, Cocal (type locality

of Sciurus milleri J. A. Allen); Cesar, Colonia Agricola de

Caracolicito (type locality of Sciurus granatensis agricolae

Hershkovitz); Chocó, Baudó (type locality of Mesosciurus

(Histriosciurus) gerrardi baudensis J. A. Allen), Río Salaqui

(type locality of Sciurus gerrardi salaquensis J. A. Allen);

Córdoba, Catival, upper Río San Jorge (FMNH 69008),

Socorré, upper Río Sinú (FMNH 68987); Cundinamarca,

Bogotá (type locality of Sciurus chrysuros Pucheran and

Sciurus hyporrhodus Gray); Huila, La Palma (AMNH

33669); La Guajira, Laguna de Junco (USNM 271318),

Puerto Estrela (USNM 216696); Magdalena, Don Diego

(AMNH 23317); Meta, Villavicencio (USNM 241350);

Nariño, Guayacana (USNM 309034); Norte de Santander,

El Guayabal, 10 mi from San José de Cucuta (type locality

of Sciurus gerrardi cucutae J. A. Allen), San Calixto, Río

Tarra (type locality of Sciurus granatensis tarrae Hershko-

vitz); Valle del Cauca, Río Frio, Río Cauca (type locality of

Guerlinguetus hoffmanni quindianus J. A. Allen), Río Ra-

poso, (USNM 334696). EC UA DOR: Carchi, Atal, 5 mi SE

of San Gabriel (type locality of Sciurus candelensis carch-

ensis W. P. Harris and Hershkovitz); Chimborazo, Chunchi

(type locality of Notosciurus rhoadsi J. A. Allen); Esmer-

aldas, Panbilar (USNM 113314); Manabí, Manaví, Río

de Oro (type locality of Guerlinguetus hoffmanni manavi

J. A. Allen); Napo, Sumaco abajo (AMNH 68159); Pich-

incha, Mt. Pichincha (type locality of Sciurus hoffmanni

soederstroemi Stone). TRINIDAD AND TOBAGO: To-

bago, Runnemede (USNM 461886); Trinidad, Carenage

(AMNH 14008), Diego Martin (AMNH 188365). VEN-

EZUELA: Anzoátegui, Cantaura (USNM 283879); Apure,

Nulita, 3 km N of Nula (USNM 442011), Tama (type

locality of Sciurus griseogena tamae Osgood); Barinas,

Hato Corozal, Caño Tutumito, Arismendi (type locality

of Sciurus [Guerlinguetus] granatensis llanensis Mondolﬁ

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Suborder Sciuromorpha: Family Sciuridae 35

Guerlinguetus griseimembra J. A. Allen, 1914d:589; type

locality “Buena Vista (altitude 4500 ft.), eastern slope

of Eastern Andes, about 50 miles southeast of Bogotá,”

Meta, Colombia.

Guerlinguetus candelensis J. A. Allen, 1914d:590; type lo-

cality “La Candela (altitude 6500 ft.), near San Agustin,

Huila, Colombia.”

Sciurus gerrardi cucutae J. A. Allen, 1914d:592; type lo-

cality “El Guayabal, 10 miles north of San José de Cu-

cuta, [Norte de Santander,] Colombia (near Venezuela

boundary).”

Mesosciurus hoffmanni hoffmanni: J. A. Allen, 1915a:216;

part; name combination.

Mesosciurus (Mesosciurus) hoffmanni hyporrhodus: J. A.

Allen, 1915a:223; name combination.

Mesosciurus (Mesosciurus) griseogena griseogena: J. A. Al-

len, 1915a:226; name combination.

Guerlinguetus (Mesosciurus) griseogena meridensis: J. A.

Allen, 1915a:228; name combination.

Mesosciurus (Mesosciurus) griseimembra: J. A. Allen, 1915a:

233; name combination.

Mesosciurus (Mesosciurus) candalensis J. A. Allen, 1915a:

235; name combination and incorrect subsequent spell-

ing of Guerlinguetus candelensis J. A. Allen.

Mesosciurus (Histriosciurus) gerrardi cucutae: J. A. Allen,

1915a:247; name combination.

Mesosciurus ferminae Cabrera, 1917:49; type locality

“Baeza (Ec ua dor oriental),” Napo, Ec ua dor.

Mesosciurus candelensis sumaco Cabrera, 1917:51; type

locality “San José, al pie del monte Sumaco, [Napo,]

Ec ua dor oriental.”

Sciurus gerrardi imbaburae W. P. Harris and Hershkovitz,

1938:1; type locality “Peñaherrera (Intag) western sub-

tropical part of Imbabura Province, Ec ua dor; altitude,

approximately 1500 meters.”

Sciurus candelensis carchensis W. P. Harris and Hershkov-

itz, 1938:3; type locality “Atal, about ﬁ ve miles south-

east of San Gabriel, Montúfar, Carchi Province, Ec ua-

dor, in the cold temperate rainforests of the western

slopes of the eastern cordillera of the Andes. Altitude

about 2900 meters.”

Mesosciurus griseogena meridensis: Cabrera and Yepes,

1940:194; name combination.

Mesosciurus caudeleusis carchensis: Cabrera and Yepes,

1940:194; name combination and incorrect subsequent

spelling of Guerlinguetus candelensis J. A. Allen.

Mesosciurus gerrardi imbaburae: Cabrera and Yepes, 1940:

195; name combination.

Mesosciurus gerrardi cucutae: Cabrera and Yepes, 1940:

195; name combination.

Sciurus [(Guerlinguetus)] hoffmanni hyporrhodus: Ellerman,

1940:340; name combination.

Notosciurus g. chapmani occurs on the islands of Trini-

dad and Tobago as well as neighboring coastal Venezuela,

including both lowlands and coastal ranges. Specimens

on the islands average slightly smaller than those on the

mainland but are otherwise indistinguishable. Top of head

and dorsum dark brown streaked with orange or pale or-

ange; postauricular patches absent (the only case of this,

along with N. g. morulus, in the genus); venter differently

colored than sides, being washed with orange to reddish

orange on the throat, chest, and belly (hairs with a short,

gray base and orange on the remainder); and tail colored

like the dorsum, but the distal tips of hairs are washed

with red.

N. g. chrysuros (Pucheran, 1845)

synonyms:

Sciurus chrysuros Pucheran, 1845:337, type locality “Co-

lombie (Santa- Fé de Bogota),” Cundinamarca, Colombia.

Sciurus chrysurus: Wagner, 1850:272; unnecessary Lat-

inzation of chrysuros Pucheran.

Sciurus hyporrhodus Gray, 1867:419; type locality “Santa

Fé de Bogotá,” Cundinamarca, Colombia.

Macroxus griseogena Gray, 1867:429; type locality given

as a number of localities from Mexico to Colombia and

Venezuela.

[Funambulus] chrysurus: Fitzinger, 1867a:487; name combi-

nation, and incorrect Latinization of chrysuros Pucheran.

Sciurus griseogenys Alston, 1878:667; unjustiﬁ ed emenda-

tion of griseogena Gray.

Sciurus rufo- niger: Alston, 1878:669; part.

Sciurus aestuans: Thomas, 1880:400; part; not aestuans

Linnaeus, 1766.

Macroxus chrysuros: E.- L. Trouessart, 1897:427; part;

name combination.

Sciurus griseogena: W. Robinson and Lyon, 1901:144;

name combination.

Sciurus griseogena meridensis Thomas, 1901c:192; type

locality “Escorial, Sierra de Merida. Alt. 2500 m,”

Mérida, Venezuela.

Sciurus griseogena tamae Osgood, 1912:48; type locality

“Paramo de Tama, [Norte de Santander,] Colombia.

Alt. 6000– 7000 ft.”

Sciurus hoffmanni: J. A. Allen, 1912:90; part; name

combination.

Notosciurus rhoadsi J. A. Allen, 1914d:585; type locality

“Pagma Forest, Chunchi (altitude 6300 feet), [Chimbo-

razo,] Ec ua dor.”

Sciurus hoffmanni söderströmi Stone, 1914:14; type local-

ity “Mt. Pichincha,” Pichincha, Ec ua dor.

Sciurus griseogena klagesi Thomas, 1914d:240; type local-

ity “Galifaré, Cerro del Avila [= Pico Ávila], near Cara-

cas, [Vargas,] Venezuela. Alt. 6500′.”

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36 Mammals of South America

Sciurus (Guerlinguetus) granatensis chrysurus: Cabrera,

1961:363; name combination, and incorrect Latiniza-

tion of chrysuros Pucheran.

Sciurus (Guerlinguetus) granatensis ferminae: Cabrera,

1961:363; name combination.

Sciurus (Guerlinguetus) granatensis griseogena: Cabrera,

1961:364; name combination.

Sciurus (Guerlinguetus) granatensis imbaburae: Cabrera,

1961:365; name combination.

Sciurus (Guerlinguetus) granatensis meridensis: Cabrera,

1961:366; name combination.

Sciurus (Guerlinguetus) granatensis söderströmi: Cabrera,

1961:368; name combination.

Sciurus (Guerlinguetus) granatensis sumaco: Cabrera, 1961:

368; name combination.

Sciurus (Guerlinguetus) granatensis tarrae: Cabrera, 1961:

369; name combination.

Sciurus (Guerlinguetus) granatensis llanensis Mondolﬁ and

Boher, 1984:312, type locality “Hato Corozal, Caño

Tutumito, Arismendi, Barinas, Venezuela.”

Sciurus (Guerlinguetus) granatensis: Eisenberg, 1989:331,

part; name combination.

Sciurus (Guerlinguetus) granatensis soederstroemi: Thor-

ington and Hoffmann, 2005:761; name combination;

required replacement of diacritical marks used in origi-

nal description (ICZN 1999:Art. 32.5.2.1).

Notosciurus g. chrysuros is the name for the taxon oc-

curring on the eastern slopes of the Andes, from Venezu-

ela to Ec ua dor. Crown of the head brown to dark brown

streaked with pale yellow to dark red; dorsum uniformly

colored brown to dark brown streaked with yellow, gray-

ish yellow to red, sometimes with black on midline; pale

orange but inconspicuous postauricular patches present;

venter reddish orange on the throat, chest, and belly, with

white spots on the chest in some specimens. Tail colored

like dorsum on its proximal quarter to third, and washed

with reddish orange succeeded by a black tip on some

specimens; in others, tail may have an intermediate section

washed with red and a black distal portion.

N. g. granatensis (Humboldt, 1811)

synonyms:

Sciurus granatensis Humboldt, 1811:8, 13, plate III; type

locality “Carthagène [= Cartagena],” Bolívar, Colombia.

Sciurus Grenatensis Illiger, 1815:69; incorrect subsequent

spelling of Sciurus granatensis Humboldt.

Sciurus variabilis I. Geoffroy St.- Hilaire, 1833:2, plate 4;

type locality “Amérique [in reference to both North and

South America].” [The volume is dated 1832 but was

published in “Mars 1833.”]

Sciurus splendidus Gray, 1842:262; no type locality given

(see Thomas 1928e:590).

Sciurus [(Guerlinguetus)] hoffmanni soderstromi: Ellerman,

1940:340; name combination.

Sciurus [(Guerlinguetus)] griseogena griseogena: Ellerman,

1940:340; name combination.

Sciurus [(Guerlinguetus)] griseogena meridensis: Ellerman,

1940:340; name combination.

Sciurus [(Guerlinguetus)] griseimembra: Ellerman, 1940:

341; name combination.

Sciurus [(Guerlinguetus)] candelensis candelensis: Eller-

man, 1940:341; name combination.

Sciurus [(Guerlinguetus)] candelensis sumaco: Ellerman,

1940:341; name combination.

Sciurus [(Guerlinguetus)] ferminae: Ellerman, 1940:341;

name combination.

Sciurus [(Guerlinguetus)] gerrardi cucutae: Ellerman, 1940:

342; name combination.

Sciurus [(Notosciurus)] rhoadsi: Ellerman, 1940:344;

name combination.

Sciurus granatensis tarrae Hershkovitz, 1947:26; type lo-

cality “Rio Tarra, a small tributary of the upper Cata-

tumbo, San Calixto, Department of Norte de Santander,

Colombia; altitude 200 meters.”

Sciurus granatensis griseogena: Hershkovitz, 1947:29;

name combination.

Sciurus granatensis meridensis: Hershkovitz, 1947:30;

name combination.

Sciurus granatensis salaquensis: Hershkovitz, 1947:35;

name combination.

Sciurus granatensis chrysurus: Hershkovitz, 1947:36;

name combination, and unnecessary Latinization of

chrysuros Pucheran.

Sciurus granatensis griseimembra: Hershkovitz, 1947:36;

name combination.

Sciurus granatensis candalensis: Hershkovitz, 1947:36;

name combination.

Sciurus granatensis carchensis: Hershkovitz, 1947:37;

name combination.

Sciurus granatensis söderströmi: Hershkovitz, 1947:37;

name combination, and incorrect use of diacritical

marks (ICZN 1999:Art. 32.5.2.1).

Sciurus granatensis imbaburae: Hershkovitz, 1947:37;

name combination.

Sciurus granatensis ferminae: Hershkovitz, 1947:37; name

combination.

Sciurus granatensis sumaco: Hershkovitz, 1947:37; name

combination.

Sciurus (Guerlinguetus) granatensis griseimembra: Good-

win, 1953:276; name combination.

Sciurus (Guerlinguetus) granatensis candelensis: Goodwin,

1953:277; name combination.

Sciurus (Guerlinguetus) granatensis carchensis: Cabrera,

1961:363; name combination.

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Suborder Sciuromorpha: Family Sciuridae 37

Sciurus [(Guerlinguetus)] gerrardi zuliae: Ellerman, 1940:

342; name combination.

Sciurus [(Guerlinguetus)] gerrardi valdiviae: Ellerman,

1940:342; name combination.

Sciurus [(Guerlinguetus)] splendidus splendidus: Ellerman,

1940:342; name combination.

Sciurus [(Guerlinguetus)] splendidus saltuensis: Ellerman,

1940:342; name combination.

Sciurus [(Guerlinguetus)] splendidus bondae: Ellerman,

1940:342; name combination.

Sciurus granatensis granatensis: Hershkovitz, 1947:12;

name combination.

Sciurus granatensis bondae: Hershkovitz, 1947:13; name

combination.

Sciurus granatensis saltuensis: Hershkovitz, 1947:15; name

combination.

Sciurus granatensis agricolae Hershkovitz, 1947:16; type

locality “Colonia Agrícola de Caracolicito, Río Ara-

guaní, on the southern slopes of the Sierra Nevada de

Santa Marta, Department of Magdalena, Colombia; al-

titude 335 meters.”

Sciurus granatensis splendidus: Hershkovitz, 1947:17;

name combination.

Sciurus granatensis variabilis: Hershkovitz, 1947:19; name

combination.

Sciurus granatensis norosiensis Hershkovitz, 1947:21, type

locality “Norosí, Mompós, Department of Bolívar, Co-

lombia; altitude 120 meters.”

Sciurus granatensis perijae Hershkovitz, 1947:23, type lo-

cality “Sierra Negra, above Villanueva, on the western

slope of the Sierra de Perijá, Valledupar, Department of

Magdalena, Colombia; altitude 1,265 meters.”

Sciurus granatensis maracaibensis Hershkovitz, 1947:25,

type locality “El Panorama, Río Aurare, a small river

emptying into lake Maracaibo, opposite the city of Ma-

racaibo, Zulia, Venezuela; altitude near sea level.”

Sciurus granatensis zuliae: Hershkovitz, 1947:28; name

combination.

Sciurus granatensis gerrardi: Hershkovitz, 1947:35; name

combination.

Sciurus granatensis valdiviae: Hershkovitz, 1947:35; name

combination.

Sciurus granatensis quindianus: Hershkovitz, 1947:36;

name combination.

Sciurus (Guerlinguetus) granatensis bondae: Goodwin,

1953: 275; name combination.

Sciurus (Guerlinguetus) granatensis splendidus: Goodwin,

1953:275; name combination.

Sciurus (Guerlinguetus) granatensis valdiviae: Goodwin,

1953:276; name combination.

Sciurus (Guerlinguetus) granatensis quindianus: Goodwin,

1953:276; name combination.

Macroxus variabilis: Lesson, 1842:112; name combination.

Sciurus gerrardi Gray, 1861:92, type locality “New

Granada.”

Sciurus gerrardii: Gerrard, 1862:214; name combination and

incorrect subsequent spelling of Sciurus gerrardi Gray.

Macroxus gerrardii: Gray, 1867:430; name combination and

incorrect subsequent spelling of Sciurus gerrardi Gray.

Sciurus variabilis variabilis: Bangs, 1898b:184; name

combination.

Sciurus variabilis saltuensis Bangs, 1898b:185, type local-

ity “Pueblo Viejo [= El Pueblito], [north slope Sierra Ne-

vada de Santa Marta, La Guajira,] Colombia (altitude

8000 ft [2,000 ft; see Paynter 1997:139]).”

Sciurus saltuensis bondae J. A. Allen, 1899b:213, type

locality “Bonda, Santa Marta District, [Magdalena,]

Colombia.”

Sciurus saltuensis: J. A. Allen, 1904c:431; name combination.

Sciurus variabilis gerrardi: E.- L. Trouessart, 1904:326;

name combination.

Sciurus versicolor zuliae Osgood, 1910:26; type locality

“Orope, Zulia, Venezuela.”

Guerlinguetus hoffmanni quindianus J. A. Allen, 1914d:

587, type locality “Rio Frio (central Rio Cauca Valley,

altitude 3500 feet), western slope of Central (or Quindio)

Andes,” Magdalena, Colombia.

Sciurus saltuensis magdalenae J. A. Allen, 1914d:593, type lo-

cality “Banco (altitude 50– 100 feet) Rio Magdalena, a few

miles above mouth of Rio Caura,” Magdalena, Colombia.

Mesosciurus (Mesosciurus) hoffmanni quindianus: J. A.

Allen, 1915a:222; name combination.

Mesosciurus (Histriosciurus) gerrardi gerrardi: J. A. Allen,

1915a:236; name combination.

Mesosciurus (Histriosciurus) gerrardi zuliae: J. A. Allen,

1915a:246; name combination.

Mesosciurus (Histriosciurus) saltuensis saltuensis: J. A.

Allen, 1915a:247; name combination.

Mesosciurus (Histriosciurus) saltuensis bondae: J. A. Allen,

1915a:249; name combination.

Mesosciurus (Histriosciurus) saltuensis magdalenae: J. A.

Allen, 1915a:251; name combination.

Mesosciurus (Histriosciurus) gerrardi valdiviae J. A. Allen,

1915a:309; type locality “Puerto Valdivia (alt. 360 ft.

[600 ft; see Paynter 1997:352]), lower Rio Caura,” An-

tioquia, Colombia.

Mesosciurus hoffmanni quindanus: Cabrera and Yepes,

1940:194; name combination and incorrect subsequent

spelling of Guerlinguetus hoffmanni quindianus J. A.

Allen.

Sciurus [(Guerlinguetus)] hoffmanni quindianus: Ellerman,

1940:340; name combination.

Sciurus [(Guerlinguetus)] gerrardi gerrardi: Ellerman,

1940:341; name combination.

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38 Mammals of South America

Sciurus milleri J. A. Allen, 1912:91; type locality “Cocal

(altitude 4000 ft.),” Cauca, Colombia.

Sciurus variabilis choco Goldman, 1913:4, type locality

“Cana (altitude 3500 feet). [Darién,] Eastern Panama.”

Guerlinguetus hoffmanni manavi J. A. Allen, 1914d:589,

type locality “Manavi [= Manabí] (Rio de Oro, near

sea- level), Ec ua dor,” [= sea port of Bahía de Caráquez,

Manabí province; see Paynter 1993:15].

Sciurus gerrardi salaquensis J. A. Allen, 1914d:592, type lo-

cality “Rio Salaqui, [Chocó,] northwestern Colombia.”

Mesosciurus (Mesosciurus) hoffmanni manavi: J. A. Allen,

1915a:221; name combination.

Mesosciurus (Histriosciurus) gerrardi milleri: J. A. Allen,

1915a:241; name combination.

Mesosciurus (Histriosciurus) gerrardi versicolor: J. A.

Allen, 1915a:242; name combination.

Mesosciurus (Histriosciurus) gerrardi morulus: J. A. Allen,

1915a:243; name combination.

Mesosciurus (Histriosciurus) gerrardi choco: J. A. Allen,

1915a:244; name combination.

Mesosciurus (Histriosciurus) gerrardi salaquensis: J. A.

Allen, 1915a:245; name combination.

Mesosciurus (Histriosciurus) gerrardi baudensis J. A. Allen,

1915a:308; type locality “Baudo [= Serranía de Baudó]

(alt. 3500 ft.), [Chocó,] coast region of western Colombia.”

Sciurus (Guerlinguetus) variabilis choco: Elliot, 1917:19;

name combination.

Sciurus (Guerlinguetus) gerrardi choco: Goldman, 1920:

139; name combination.

Sciurus (Guerlinguetus) gerrardi morulus: Goldman, 1920:

140; name combination.

Sciurus gerrardi inconstans Osgood, 1921:40, replacement

name for Sciurus versicolor Thomas, 1900, said to be

preoccupied by Sciurus versicolor Zimmermann, 1777;

Hershkovitz (1947:36) stated that Zimmermann’s name

was invalid, but without specifying a reason.

Sciurus (Mesosciurus) gerrardi leonis Lawrence, 1933:369,

replacement name for Sciurus milleri J. A. Allen, 1912,

preoccupied by Sciurus epomophorus milleri H. C.

Robinson and Wroughton, 1911.

Sciurus [(Guerlinguetus)] hoffmanni manavi: Ellerman,

1940:340; name combination.

Sciurus [(Guerlinguetus)] gerrardi leonis: Ellerman, 1940:

341; name combination.

Sciurus [(Guerlinguetus)] gerrardi inconstans: Ellerman,

1940: 341; name combination.

Sciurus [(Guerlinguetus)] gerrardi morulus: Ellerman, 1940:

341; name combination.

Sciurus [(Guerlinguetus)] gerrardi choco: Ellerman, 1940:

341; name combination.

Sciurus [(Guerlinguetus)] gerrardi salaquensis: Ellerman,

1940:342; name combination.

Sciurus (Guerlinguetus) granatensis agricolae: Cabrera,

1961:362; name combination.

Sciurus (Guerlinguetus) granatensis gerrardi: Cabrera,

1961:363; name combination.

Sciurus (Guerlinguetus) granatensis granatensis: Cabrera,

1961:364; name combination.

Sciurus (Guerlinguetus) granatensis maracaibensis: Ca-

brera, 1961:365; name combination.

Sciurus (Guerlinguetus) granatensis noroslensis: Ca-

brera, 1961:367; name combination and incorrect

subsequent spelling of Sciurus granatensis norosiensis

Hershkovitz.

Sciurus (Guerlinguetus) granatensis perijae: Cabrera,

1961:367; name combination.

Sciurus (Guerlinguetus) granatensis saltuensis: Cabrera,

1961: 368; name combination.

Seiurus [sic] (Guerlinguetus) granatensis variabilis: Cabrera,

1961:369; name combination.

Sciurus (Guerlinguetus) granatensis zuliae: Cabrera, 1961:

370; name combination.

Sciurus (Guerlinguetus) granatensis norosiensis: Thorington

and Hoffmann, 2005:761; name combination.

Sciurus (Guerlinguetus) granatensis variabilis: Thorington

and Hoffmann, 2005:761; name combination.

The nominotypical subspecies is parapatric to N. g. moru-

lus, occurring at high elevations in the western Andes of Co-

lombia to the Sierra de Santa Marta, and across the central

and eastern cordilleras of Colombia to western Venezuela.

This subspecies also extends into the intervening lowlands,

including the area around Lake Maracaibo. Average size

larger than for all other subspecies; postauricular patches

present; dorsal color an “agouti” pattern, mostly brown

streaked with orange or red, or uniformly completely orange;

posterior dorsum may exhibit dark patterning, ranging from

a spot near the base of the tail to larger area; dorsal orange

or red tones may appear merely as a scapular patch or extend

farther posteriorly. Ventral color ranges from entirely red to

orange with white spots (of varying sizes) on chest and belly

to entirely white; white ventral spots may also extend as con-

tinuous lines across body at midventer. Entirely orange or red

individuals with a white venter predominate in the northern

parts of the range, on the slopes of the Serra de Santa Marta;

the “agouti” dorsal pattern with a mostly or entirely red ven-

ter is most common in southern samples.

N. g. morulus (Bangs, 1900)

synonyms:

Sciurus variabilis morulus Bangs, 1900b:43, type locality

“Loma del Leon, Panama.”

Sciurus versicolor Thomas, 1900g:385: type locality

“Cachabi [= Cachabí], Prov. Esmeraldas, N Ec ua dor,

alt. 160m.”

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Suborder Sciuromorpha: Family Sciuridae 39

i cal trend. Head and back uniformly colored brown, heavily

streaked with orange; inconspicuous orange postauricular

patches present; ventral coloration sharply set off from the

dorsolateral coloration, with uniform reddish orange throat,

chest, and belly (hairs are entirely reddish orange). Color-

ation of tail variable, similar to that of dorsum on its proxi-

mal quarter to third but washed with reddish orange distally.

natural history: Nitikman (1985) summarized

the data on form and function, ontogeny and reproduc-

tion, ecol ogy, behavior, and ge ne tics of N. granatensis, with

most studies focused on Panamanian populations (e.g.,

Heaney 1978; Glanz et al. 1982; Glanz 1984).

remarks: Samples referred to N. granatensis range

from Costa Rica to Panama, and then south to Andean

South America, from western Venezuela to northern Ec ua-

dor. As the synonymies for each subspecies indicate, doz-

ens of names have been proposed for squirrels from this

vast area (Cabrera 1961, for example, listed 28 subspecies

in his concept of this polytypic species). One could eas-

ily conclude that the diversity of squirrels in this region

is extensive simply by reading the original descriptions

of these taxa. Indeed, based on our own examination of

hundreds of specimens deposited in the major U.S. and

Eu ro pe an museums as well as most of the type material,

both the nongeographic and geographic variation exhib-

ited by these samples is staggering. In fact, many subspe-

cies have been described from localities very close to one

another, without even a separation by elevation or river

basin. Hershkovitz (1947) personally collected dozens of

specimens in Colombia and described several new sub-

species, some of them with identical provenance! Other

authors have frequently described series in which different

Sciurus [(Guerlinguetus)] gerrardi baudensis: Ellerman,

1940: 342; name combination.

Sciurus granatensis salaquensis: Hershkovitz, 1947:35; name

combination.

Sciurus granatensis leonis: Hershkovitz, 1947:36; name

combination.

Sciurus granatensis versicolor: Hershkovitz, 1947:36; name

combination.

Sciurus granatensis manavi: Hershkovitz, 1947:36; name

combination.

Sciurus (Guerlinguetus) granatensis salaquensis: Goodwin,

1953:275.

Sciurus (Guerlinguetus) granatensis leonis: Goodwin, 1953:

276; name combination.

Sciurus (Guerlinguetus) granatensis manavi: Goodwin,

1953:276; name combination.

Sciurus (Guerlinguetus) granatensis choco: G. S. Miller

and Kellogg, 1955:257; name combination.

Sciurus (Guerlinguetus) granatensis morulus: G. S. Miller

and Kellogg, 1955:257; name combination.

Sciurus (Guerlinguetus) granatensis versicolor: Cabrera,

1961:370; name combination.

Notosciurus g. morulus is the oldest available name for

the subspecies occurring from the Panama Canal Zone to

western Colombia and northwestern Ec ua dor. Individuals

lack postauricular patches over most of this range, except

in Ec ua dor, where traces of a patch are present on speci-

mens. The mid- dorsum slightly to more strongly darker

than in other races, a characteristic more pronounced in

Colombian specimens than those of central and eastern

Panama; venter red, with hairs lacking a grayish base; tail

usually divided into three differently colored sections: a

dark proximal portion, a large middle red section, and a

dark (black) distal terminus.

N. g. nesaeus (G. M. Allen, 1902)

synonyms:

Sciurus nesaeus G. M. Allen, 1902:93; type locality “El

Valle, Margarita Island, [Nueva Esparta,] Venezuela.”

Mesosciurus (Mesosciurus) nesaeus: J. A. Allen, 1915a:233;

name combination.

Sciurus [(Guerlinguetus)] nesaeus: Ellerman, 1940:341;

name combination.

Sciurus granatensis nesaeus: Hershkovitz, 1947:37; name

combination.

Sciurus (Guerlinguetus) granatensis nesaeus: Cabrera, 1961:

367; name combination.

Notosciurus g. nesaeus is an insular subspecies known

only from Isla Margarita, Venezuela. It is easily distinguished

from mainland samples, but we have decided not to recog-

nize it as a distinct species on the same grounds as Hershko-

vitz (1947); that is, the color pattern ﬁ ts a general geo graph- Map 16 Selected localities for Notosciurus granatensis (●). Contour

line = 2,000 m.

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40 Mammals of South America

selected localities (Map 17): ARGENTINA:

Jujuy, Higuerilla (type locality of Sciurus [Mesosciurus] ar-

gentinius Thomas). BOLIVIA: Beni, Totaisal, 1 km SW of

Estación Biologica del Beni (AMNH 261917); Chuquisaca,

Sud Cinti, 15.6 km N of El Palmar, above Río Santa Marta

(LHE 1310); Cochabamba, Incachaca (FMNH 35224);

Río Chapare (FMNH 21547); La Paz, Charuplaya (BM

2.1.1.17), Mapiri (BM 1.1.1.58); Santa Cruz, Buenavista

(FMNH 21423), Santiago, Chiquitos (FMNH 105927);

Tarija, pié Sierra Santa Rosa, Itau (BM 25.2.1.18– 23), Ver-

mejo [= Bermejo] (AMNH 38994). BRAZIL: Acre, Ma-

noel Urbano (MZUSP 11235), Oriente, near Taumaturgo

(MPEG 758), Plácido de Castro (type locality of Sciurus

ignitus iquiriensis Vieira), Rio Branco (AMNH 149145);

Amazonas, Igarapé Grande [= Eirunepé], left bank Rio Ju-

ruá (MZUSP 4581); Mato Grosso, São João, cabeceiras do

Rio Aripuanã (type locality of Guerlinguetus rufus Moojen);

Rondônia, Ouro Preto d’Oeste (MPEG 20529), Samuel

(MPEG, ﬁ eld number 402– 8557). COLOMBIA: Antioquia,

4 km NE of Bellavista, above Río Ponce (FMNH 70038),

Urrao, Río Aná (FMNH 71106), 15 km E of Río Negrito,

Sonsón (FMNH 70043), 10 km S of Valdivia (FMNH

70048); Boyacá, Miraﬂ ores (AMNH 32173); Cauca,

Salento, west Quindio Andes (AMNH 32850); Cundina-

marca, El Roble, above Fusugasugá, east Andes (AMNH

34622), Panamá (AMNH 34680); Huila, Andalucáa, east-

ern Andes (AMNH 33700), La Candela (AMNH 33699),

San Adolfo, Acevedo (FMNH 71110); Nariño, La Laguna

(AMNH 42358); Valle del Cauca, vicinity of Palmira, cen-

tral Andes (AMNH 32175). PERU: Cusco, Cosñipata, Ha-

cienda Cadena (FMNH 65673), Ocabamba, near Cusco

(type locality of Sciurus aestuans cuscinus Thomas), Río

San Miguel (BM 22.1.1.21); Huánuco, Agua Caliente, Río

Pachitea (FMNH 55407), Tingo Maria (FMNH 24116);

Junín, Perené (AMNH 61814); Loreto, Puerto Punga, Río

Tapiche (AMNH 99203), Quebrada Pushaga, Río Morona,

upper Río Amazonas (FMNH 88974), San Fernando, Río

Yavari (FMNH 88981), Santa Cecilia, Río Maniti (FMNH

87173), Sarayacu, Río Ucayali (AMNH 76442); Madre

de Dios, Altamira, Manu (FMNH 98060), Río Inambari

(AMNH 16562); Pasco, Rumicruz (AMNH 60599); Puno,

Pampa Grande, Sandia (FMNH 79902); Ucayali, Lagarto,

upper Río Ucayali (AMNH 78943).

subspecies: We recognize the four subspecies of N.

pucheranii.

N. p. argentinius (Thomas, 1921)

synonyms:

Sciurus (Mesosciurus) argentinius Thomas, 1921k:609,

type locality “Higuerilla, 2000 m., in the Department of

Valle Grande, about 10 km. east of the Zenta range and

20 km. of the town Tilcara,” Jujuy, Argentina.

specimens from the same locality could be assigned to sep-

arate subspecies.

Here, we offer a brief summary of geographic trends

expressed in N. granatensis, and we attach names that we

believe best represent the major axes of this variation. In so

doing, we reduce the number of subspecies to only ﬁ ve for

the South American part of the species’ distribution. The

description we provide above for each one can, with be-

nevolence, be termed “impressionistic.” However, we can-

not accept a single name for all such squirrels at the risk of

obscuring a most bafﬂ ing but manifest evolutionary phe-

nomenon. In fact, Hershkovitz (pers. comm. to MdV) ﬁ rst

devised his “metachromism theory” (Hershkovitz 1968,

1970a) because he wanted to explain the color variation he

saw in Colombian squirrels. Keeping at least some of the

plethora of names as “valid” will, we hope, encourage re-

searchers to study these squirrels further and discover how

and why they vary so much.

Notosciurus pucheranii (Fitzinger, 1867)

Pucheran’s Squirrel, Ardija, Quatipuru

synonyms: See under subspecies.

description: Variable squirrel of medium size, with

head and body length 151– 212 mm (N = 121 adult speci-

mens), tail length 135– 225 mm (N = 117), hindfoot length

34– 58 mm (N = 125), ear length 15– 29 mm (N = 109), and

mass 136– 242 g (N = 6). Dorsal color of head and back

brown, streaked with orange, sometimes with deﬁ ned

mid- dorsal black stripe, and darker than that of sides;

cheeks brownish orange; orange eye- ring present, and or-

ange postauricular patches conspicuous. Ventral surfaces

distinct in color from dorsolateral region, with throat

white or the same white, yellow, or orange color as chest;

belly orange; ventral hairs usually have gray base. Didier

(1955) illustrated the baculum for specimens from eastern

Peru.

distribution: Notosciurus pucheranii is a polytypic

species of primarily Andean distribution, ranging from the

central and eastern Andean forests of Colombia to the east-

ern Andes of Peru, and from the Peruvian lowlands to west-

ern Amazonian Brazil, Bolivia, and northwestern Argentina.

Their ecological range spans upper montane forests, low-

land rainforest, and lowland seasonally dry forests, across a

broad elevational range from 100 to 3,200 m. A geographic

gap in this range is apparent between the Nariño depart-

ment in southern Colombia and the Loreto department in

northern Peru, with no specimens as yet recorded from Ec-

ua dor. It is unclear whether this gap is real or the result of

inadequate sampling. Most collecting localities are Andean

or situated in Andean foothills; there are fewer localities at

lower elevations east of the Andean slopes.

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Suborder Sciuromorpha: Family Sciuridae 41

Leptosciurus ignitus ignitus: J. A. Allen, 1915a:204; name

combination.

Leptosciurus ignitus irroratus: J. A. Allen, 1915a:206;

name combination.

Sciurus irroratus ochrescens: Osgood, 1916:204; name

combination.

Sciurus ignitus irroratus: Thomas, 1927f:599; name

combination.

Sciurus [(Guerlinguetus)] ignitus ignitus: Ellerman,

1940:344; name combination.

Sciurus [(Guerlinguetus)] ignitus irroratus: Ellerman,

1940:344; name combination.

Guerlinguetus rufus Moojen, 1942:14; type locality “S. João

[São João] (cabeceiras do Aripuanã) M. [Mato] Grosso,”

Brazil; preoccupied by Sciurus rufus Kerr, 1792.

Sciurus (Guerlinguetus) rufus: W. P. Harris, 1944:12; name

combination.

Sciurus rufus: Moojen, 1952b:28; name combination.

Sciurus ignitus ignitus: Sanborn, 1951:18; name

combination.

Sciurus ignitus iquiriensis C. O. da C. Vieira, 1952:28, type

locality “Placido de Castro, Rio Abunã, Acre,” Brazil.

Sciurius [sic] ignitus: Didier, 1955:423; name combination,

but typographic error.

Sciurus (Guerlinguetus) ignitus iquiriensis: C. O. da C.

Vieira, 1955:408; name combination.

Sciurus cabrerai Moojen, 1958:50; replacement name for

Guerlinguetus rufus Moojen, 1942.

Sciurus [(Guerlinguetus)] ignitus cabrerai: Cabrera, 1961:

372; name combination.

Sciurus [(Guerlinguetus)] ignitus: R. S. Hoffmann, Anderson,

Thorington, and Heaney, 1993:441; name combination.

Guerlinguetus ignitus: J. A. Oliveira and Bonvicino, 2006:

348; name combination.

This subspecies occurs on the eastern Andean slopes

and in the Amazon Basin from northern to southern Peru,

extending eastward to the states of Acre and Amazonas in

Brazil.

N. p. pucheranii (Fitzinger, 1867)

synonyms:

Sciurus rufoniger Pucheran, 1845:336; type locality “Co-

lombie (Santa- Fé de Bogotá),” Cundinamarca, Colom-

bia; preoccupied by Sciurus rufonigra Gray, 1842 [see

Fitzinger 1867a].

Funambulus Pucheranii Fitzinger, 1867a:487; replacement

name for Sciurus rufoniger Pucheran [preoccupied by

Sciurus rufonigra Gray, 1842]; type locality given as

“Am. Columbien.”

Macroxus griseogena Gray, 1867:429; at least in part; no

explicit type locality was given, but “Santa Fé de Bo-

gotá” was listed among the several localities cited.

Sciurus argentinius: Thomas, 1925b:577; name combination.

Leptosciurus argentinius: Cabrera and Yepes, 1940:193;

name combination.

Sciarus [sic] [(Guerlinguetus)] ignitus argentinius: Cabrera,

1961:370; name combination.

This subspecies occurs in extreme southern Bolivia and

adjacent northwestern Argentina.

N. p. boliviensis (Osgood, 1921)

synonyms:

Macroxus leucogaster Gray, 1867:430, type locality “Bo-

livia, Santa Cruz de la Sierra”; preoccupied by Sciurus

leucogaster F. Cuvier, 1831:300 (= Sciurus aureogaster

F. Cuvier, 1829c).

Leptosciurus leucogaster: J. A. Allen, 1915a:207; name

combination.

Sciurus boliviensis Osgood, 1921:39, replacement name

for Macroxus leucogaster Gray, 1867, which is preoc-

cupied by Sciurus leucogaster F. Cuvier, 1831.

Sciurus [(Guerlinguetus)] boliviensis: Ellerman, 1940:344;

name combination.

Sciurus sanborni Osgood, 1944:191; type locality “La

Pampa, between Rio Inambari and Rio Tambopata,

about twenty miles north of Santo Domingo, Madre de

Dios, Peru. Altitude about 1900 feet.”

Sciurus ignitus boliviensis: Sanborn, 1951:18; name

combination.

Sciurus [(Guerlinguetus)] ignitus boliviensis: Cabrera,

1961:371; name combination.

Sciurus [(Guerlinguetus)] sanborni: Cabrera, 1961:373;

name combination.

Notosciurus p. boliviensis is distributed from southern

Peru to central Bolivia.

N. p. ignitus (Gray, 1867)

synonyms:

Macroxus ignitus Gray, 1867:429, type locality “Bolivia.”

Macroxus irroratus Gray, 1867:431, type locality “Brazil,

Upper Ucayali.”

Sciurus aestuans cuscinus Thomas, 1899a:40, type locality

“Ocabamba, Cuzco,” Peru.

S[ciurus]. irroratus: Thomas, 1899a:40; name combination.

Sciurus cuscinus: Thomas, 1902b:129; name combination.

Sciurus irroratus: Ihering, 1904:420; name combination.

[Sciurus (Guerlinguetus)] aestuans cuscinus: E.- L. Troues-

sart, 1904:327; name combination.

[Sciurus (Guerlinguetus)] irroratus: E.- L. Trouessart,

1904:328; name combination.

Sciurus cuscinus ochrescens Thomas, 1914a:362, type

locality “Bolivia, in upperparts of Beni and Mamoré

Rivers. Type from Astillero, [La Paz,] 67°W, 16°S. Alt.

2700 m.”

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42 Mammals of South America

Sciurus aestuans rufoniger: Rode, 1943:384; name com-

bination, and required elimination of hyphen used in

original description (ICZN 1999:Art. 32.5.2.2).

Sciurus pucherani caucensis: Goodwin, 1953:277; name

combination, and incorrect subsequent spelling of Fu-

nambulus pucheranii Fitzinger.

Sciurus pucheranii salentensis: Goodwin, 1953:277; name

combination.

Sciurus pucherani: Didier, 1955:416; name combination

and incorrect subsequent spelling of Funambulus puch-

eranii Fitzinger.

Sciurus pucheranii: Didier, 1955:422; name combination.

Sciurus [(Guerlinguetus)] pucheranii santanderensis

Hernández- Camacho, 1957:219, type locality “Meseta

de los Caballeros, Santander,” Colombia.

Sciurus [(Guerlinguetus)] pucheranii caucensis: Cabrera,

1961:372; name combination.

Sciurus [(Guerlinguetus)] pucheranii medellinensis: Ca-

brera, 1961:372; name combination.

Sciurus [(Guerlinguetus)] pucheranii pucheranii: Cabrera,

1961:373; name combination.

Sciurus [(Guerlinguetus)] pucheranii santanderensis: Ca-

brera, 1961:373; name combination.

Sciurus [(Guerlinguetus)] pucheranii: Eisenberg, 1989:333;

name combination.

The nominotypical subspecies is distributed in the An-

dean cordilleras of central and southern Colombia. It is geo-

graph i cally separated by a large gap from the cis- Andean dis-

tributions of the other three subspecies, as noted previously.

natural history: Leonard et al. (2009) summa-

rized the natural history, behavior, and other aspects of the

biology of this species, restricting their compilation to the

trans- Andean Colombian populations. Otherwise, little

has been published on this species, surprisingly so given

its broad cis- Andean range. Emmons and Feer (1997) pro-

vided a few ecological and behavioral details for argentin-

ius, ignitus, and sanborni, which they regarded as separate

species.

remarks: Both Funambulus pucheranii Fitzinger and

Macroxus ignitus Gray were published in 1867, but Fitz-

inger’s paper appeared in March (“21 März 1867”) and

Gray’s in December. The species name pucheranii Fitzinger

thus has priority over ignitus Gray.

Notosciurus pucheranii is highly variable throughout

its range, and many supposed species and subspecies have

been described that we treat either as subspecies rather than

species or as ju nior synonyms. All names proposed have

been largely based on coat color, which, although variable,

presents a reasonably deﬁ ned geo graph i cal trend, as follows.

Animals with denser and longer fur are found at higher el-

evations, while specimens with less dense and shorter pel-

age come from lower elevations. In northern Colombia

Macroxus tephrogaster Gray, 1867:431; probably part (?);

while no explicit type locality was given, “Bogotá” was

listed among the several localities cited. Thorington and

Hoffman (2005) treated tephrogaster as a synonym of

S. deppei.

Macroxus medellinensis Gray, 1872:408; type locality

“Medellin, Antioquia,” Colombia.

S[ciurus]. aestuans var. rufoniger: J. A. Allen, 1877:669; part.

Sciurus tephrogaster: J. A. Allen, 1877:763; name combina-

tion.

Sciurus griseogenys: Alston, 1878:667, part; name combi-

nation.

Sciurus deppei: Alston, 1878:668, part; name combination.

Sciurus rufo- niger: Alston, 1878:669; name combination.

Sciurus aestuans rufo- niger: True, 1884:595, part; name

combination.

S[ciurus]. Pucherani: Thomas, 1898f:266; name combina-

tion and incorrect subsequent spelling of Funambulus

pucheranii Fitzinger.

S[ciurus]. (Guerlinguetus) caucensis E. W. Nelson, 1899a:79,

type locality “Lima River, upper Cauca Valley, [Valle del

Cauca?; see Paynter 1997:252,] Colombia (alt. 6000

feet).”

[Sciurus] medellinensis: E.- L. Trouessart, 1904:327; name

combination.

[Sciurus (Guerlinguetus)] caucensis: E.- L. Trouessart,

1904:328; name combination.

Guerlinguetus pucheranii salentensis J. A. Allen, 1914d:587,

type locality “near Salento (altitude 9000 ft.), Central

Andes, [Cauca,] Colombia.”

Leptosciurus pucheranii pucheranii: J. A. Allen, 1915a:200;

name combination.

Leptosciurus pucheranii medellinensis: J. A. Allen,

1915a:202; name combination.

Leptosciurus pucheranii caucensis: J. A. Allen, 1915a:203;

name combination.

Leptosciurus pucheranii salentensis: J. A. Allen, 1915a:203;

name combination.

Sciurus (Leptosciurus) pucheranii medellinensis: Anthony,

1923a:5; name combination.

Leptosciurus pucherani: Cabrera and Yepes, 1940:192;

name combination and incorrect spelling of pucheranii

Fitzinger.

Sciurus [(Guerlinguetus)] pucherani pucherani: Ellerman,

1940:343; name combination and incorrect spelling of

pucheranii Fitzinger.

Sciurus [(Guerlinguetus)] pucherani medellinensis: Eller-

man, 1940:343; name combination and incorrect spell-

ing of pucheranii Fitzinger.

Sciurus [(Guerlinguetus)] pucherani salentensis: Ellerman,

1940:343; name combination and incorrect spelling of

pucheranii Fitzinger.

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Suborder Sciuromorpha: Family Sciuridae 43

with a gray base; this is the phenotype of N. p. boliviensis.

The name Sciurus sanborni Osgood has been applied to

animals with this color pattern; we treat this name as a ju-

nior synonym of boliviensis. Southern Bolivian and north-

ern Argentinean samples belong to N. p. argentinius, with

the main distinction between them and N. p. boliviensis

being in the color of the venter, which is self- colored or-

ange in the former and white with a gray base in the latter.

Specimens from Bermejo, Tarija department, Bolivia, have

a white venter (characteristic of typical boliviensis), but

this locality is situated in the middle of the range of N. p.

argentinius.

The reason that we recognize four subspecies in N.

pucheranii rather than a single, monotypic species or

four separate species is that the characters that would

identify distinct taxa do not segregate in a clear- cut man-

ner. Samples frequently contain specimens with mixed

characteristics in regions where one subspecies replaces

another.

Throughout its distribution, specimens of N. pucher-

anii possess postauricular patches and three pairs of mam-

mae. However, L. H. Emmons (pers. comm. to MdV) col-

lected a female with four pairs of mammae from Sud Cinti,

Chuquisaca, Bolivia (LHE 1313). Because all females of

Notosciurus we have examined have three pairs of mam-

mae, Emmons’s specimen may be an anomaly or perhaps

has been misidentiﬁ ed.

Genus Simosciurus J. A. Allen, 1915

Simosciurus is a genus of large, trans- Andean squirrels that

occurs in western Ec ua dor and northwestern Peru. We recog-

nize two species, both of which inhabit dry to moist forests

from sea level to about 2,300 m. Simosciurus can be distin-

guished from most South American squirrels by their coarser

pelage and larger size (relative to Sciurillus, Microsciurus,

Guerlinguetus, Notosciurus, and Syntheosciurus). Although

comparable in size, Simosciurus is distinguished from Had-

rosciurus by its very short and broad rostrum and distinctly

coarse fur. The rostral character is difﬁ cult to quantify, but it

can be visualized most readily in dorsal view comparisons.

Not only are the nasals wider but also the premaxillae re-

main visible distally to the tip of the rostrum. In Hadrosci-

urus, the premaxillae are narrower and disappear from view

before the rostral tip. The respective trans- versus cis- Andean

distributions of the two genera circumvent the possibility of

their misidentiﬁ cation based on geography alone.

J. A. Allen (1915a) erected the genus Simosciurus to in-

clude Sciurus stramineus P. Gervais, with three subspecies

(nebouxii I. Geoffroy St.- Hilaire; guayanus Thomas; and

zarumae J. A. Allen) in addition to the nominotypical form.

Moore (1959) retained stramineus in Simosciurus (as a subge-

(Antioquia and Quindio departments), most specimens have

a well- deﬁ ned mid- dorsal black stripe (characteristic of N.

p. pucheranii), but in samples from Cundinamarca, Huila,

and Nariño this stripe becomes less pronounced and in some

series disappears entirely. In Peru, from Loreto department

southward, the N. p. ignitus phenotype predominates, with

animals having dark brown dorsal parts, without a mid-

dorsal stripe, and a grayish white throat and orange chest

and belly, with ventral hairs gray at the base. This same

phenotype extends to adjacent Brazil, in the states of Ama-

zonas and Acre. Specimens from localities in central and

southern Peru have the same general color as typical N. p.

ignitus, but with less demarcation of the throat from that

of the chest and belly, and an overall ventral color ranging

from pale yellow to orange.

Southern Peruvian and adjacent Bolivian samples are

generally similar, although some Bolivian samples have

more white on the venter. By central Bolivia, this gen-

eral trend of lightening of the venter reaches the extreme,

where specimens have white bellies but with hairs still

Map 17 Selected localities for Notosciurus pucheranii (●); cis- and trans- Andean

distributions are mapped as disjunct (see Remarks). Contour line = 2,000 m.

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44 Mammals of South America

thought that J. A. Allen’s (1877) Sciurus hypopyrrhus was

the same as the Ec ua dorean taxon, an error probably re-

sulting from the unusually coarse pelage of Simosciurus,

which resembles that of some North American species and

is completely different from that of any other South Ameri-

can form. Because Sciurus hypopyrrhus is ﬁ rmly in the syn-

onymy of Sciurus aureogaster F. Cuvier, 1829 (Thorington

and Hoffmann 2005), the name Echinosciurus is not avail-

able for this genus of squirrels.

KEY TO THE SPECIES OF SIMOSCIURUS:

1. Nuchal patch present, colored white to pale gray, slightly

tinged with very pale yellow; dorsal color strongly

streaked with yellowish gray anteriorly (same color as

nuchal patch) and more vivid pale yellow posteriorly;

head about the same color as other anterior parts of

body; venter mostly pale yellowish gray. . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . Simosciurus nebouxii

1′. No nuchal patch; dorsal color usually brown heavily

streaked anteriorly with gray, sometimes yellowish gray

posteriorly; head distinctly darker than rest of dorsum,

with brownish tones; venter dark brown, sometimes

with tinge of gray . . . . . . . . . . .Simosciurus stramineus

Simosciurus nebouxii (I. Geoffroy St.- Hilaire, 1855)

White- naped Squirrel

synonyms:

Sciurus Nebouxii I. Geoffroy St.- Hilaire, 1855:165, plate

12, a– d; type locality “Pérou, à Payta [= Paita], [Piura].”

Sciurus stramineus: Alston, 1878:664; part; S. nebouxii

I. Geoffroy St.- Hilaire placed in synonymy.

Macroxus nebouxi: E.- L. Trouessart, 1897:428; name

combination, and incorrect subsequent spelling of Sci-

urus nebouxii I. Geoffroy St.- Hilaire.

Sciurus stramineus nebouxii: Thomas, 1900a:151; name

combination.

Sciurus stramineus nebouxi: E.- L. Trouessart, 1904:326 ;

name combination, and incorrect subsequent spelling of

Sciurus nebouxii I. Geoffroy St.- Hilaire.

Sciurus nebouxi: Neveu- Lemaire and Grandidier, 1911:8;

incorrect subsequent spelling of Sciurus nebouxii I.

Geoffroy St.- Hilaire.

Sciurus stramineus zarumae J. A. Allen, 1914d: 597; type

locality “Zaruma, [El Oro,] southwestern Ec ua dor; al-

titude 6000 ft.”

Simosciurus stramineus nebouxii: J. A. Allen, 1915a:280;

name combination.

Simosciurus stramineus zarumae: J. A. Allen, 1915a:284;

name combination.

Sciurus [(Guerlinguetus)] stramineus nebouxi: Ellerman,

1940:343; name combination, and incorrect subsequent

spelling of nebouxii I. Geoffroy St.- Hilaire.

nus of Microsciurus), but Ellerman (1940), Cabrera (1961),

and Thorington and Hoffmann (2005) placed the species in

Sciurus, subgenus Guerlinguetus. Allen (1915a:280– 281)

recognized that two of these taxa had a white nuchal patch

(nebouxii and zarumae) and two did not (stramineus and

guayanus). Emmons and Feer (1997) suggested that animals

possessing the nuchal patch were only a highland color vari-

ant, a position that supported Allen’s initial recognition of a

single species. Our study of more than 140 specimens, how-

ever, indicates clear geo graph i cal separation of specimens

with and without a distinct nuchal patch. We interpret this

pattern as indicative of species, rather than subspecies, and

thus recognize S. nebouxii I. Geoffroy St.- Hilaire and S. stra-

mineus P. Gervais. No mea sure ments of mass are available in

the material we examined, but the two species are of similar

size, with an average head and body length of 260 mm, tail of

about 294 mm, ear of 35 mm, and hindfoot of 63 mm.

The vernacular name “Guayaquil Squirrel” (or ardilla

de Guayaquil, Tirira 2004, 2007) has been used in the re-

cent accounts that recognize a single species of these trans-

Andean squirrels. However, because the main difference

between the two species we recognize lies in the presence

(in S. nebouxii) or absence (in S. stramineus) of a nuchal

patch, we refer to the former as the “White- naped Squir-

rel,” the En glish equivalent of the local Spanish name, ar-

dilla de nuca blanca (Tirira 2007).

synonyms:

Macroxus: Lesson, 1842:112, part (listing of stramineus);

not Macroxus F. Cuvier, 1823.

Sciurus: I. Geoffroy St.- Hilaire, 1855:165; part (inclusion

of nebouxii); not Sciurus Linnaeus, 1758.

Echinosciurus E.- L. Trouessart, 1880a:292; part (listing

of stramineus); proposed as a subgenus of Sciurus; type

species Sciurus hypopyrrhus Wagler, 1831, a synonym

of Sciurus aureogaster F. Cuvier, 1829.

Simosciurus J. A. Allen, 1915a:280; type species Sciurus

stramineus P. Gervais, 1841.

[Sciurus] (Guerlinguetus): Ellerman, 1940:343; name

combination.

Sciurus (Simosciurus): Goodwin, 1953:278; name

combination.

Microsciurus (Simosciurus): Moore, 1961a:15, table 2;

name combination.

Guerlinguetus (Simosciurus): Moore, 1961b:15; name

combination.

remarks: J. A. Allen (1915a) proposed the genus Si-

mosciurus with type species Sciurus stramineus P. Gervais,

1841 (a name often attributed to Eydoux and Souleyet,

1841, but see Remarks under the Simosciurus stramineus).

The name Echinosciurus E.- L. Trouessart actually predates

Simosciurus, but the type species of Echinosciurus is Sci-

urus hypopyrrhus Wagler, 1831. E.- L. Trouessart (1880a)

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Suborder Sciuromorpha: Family Sciuridae 45

natural history: Little is known about the ecol-

ogy and population biology of this species. One specimen

(FMNH 53864) was found in a grass nest on a canyon wall.

remarks: We regard Simosciurus stramineus and S.

nebouxii as distinct species based on their different color

patterns and separate geographic ranges. However, since

Alston (1878:665) ﬁ rst noted the appearance of “irregu-

lar tufts of pure white hairs, . . . and sometimes uniting in

larger patches,” others have viewed these differences as

constituting individual variation rather than species- group

characters (e.g., Emmons and Feer 1997). Individuals of

both color patterns have been obtained at the same locality

in southern Ec ua dor (Arenillas, El Oro province; specimens

in FMNH). Two of the three specimens available possess

the nuchal patch characteristic of S. nebouxii, but the third

has no nuchal patch, as is diagnostic of S. stramineus.

Arenillas is the northernmost locality for S. nebouxii,

and this mixture of characters may be due to unusual poly-

morphism, may indicate intergradation with the northern

S. stramineus, or may actually represent an area of distri-

butional overlap between the two species. Nevertheless, a

gap of more than 150 km exists in the known ranges of

two species between Arenilles in the south (nebouxii) and

the Guayaquil area in the north (stramineus). Collecting

efforts directed to this region, including the application

of molecular ge ne tic methods, will aid in determining the

degree of intergradation, if any, and ultimately the proper

number of species to recognize.

Sciurus [(Guerlinguetus)] stramineus zarumae: Ellerman,

1940:343; name combination.

Simosciurus stramineus zaruma: Cabrera and Yepes, 1940:

196; name combination.

Sciurus (Simosciurus) stramineus zarumae: Goodwin, 1953:

278; name combination.

Sciurus (Guerlinguetus) stramineus: Cabrera, 1961:373;

part; placed S. nebouxii I. Geoffroy St.- Hilaire and S. s.

zarumae J. A. Allen in synonymy.

Sciurus nebouxiii: Soukup, 1965:360; incorrect subsequent

spelling of Sciurus nebouxii I. Geoffroy St.- Hilaire;

placed in synonymy of S. stramineus P. Gervais.

description: Medium- sized squirrel with head

and body length 230– 281 mm (N = 52), tail length 264–

330 mm (N = 52), hindfoot length 52– 70 mm (N = 53),

and ear length 26– 43 mm (N = 24). Body color light gray,

with posterior dorsum and posterior parts of forelegs and

hindlegs pale yellow; head, including cheeks, same color

as body; ears distinctly darker than head, pinnae rimmed

with elongated black hairs; nuchal patch present, white

to grayish white; forefeet and distal parts of forelimbs

darker than body; hindfeet and distal parts of hindlegs

dark brown; venter yellow streaked with gray. Tail black

to light gray, except near yellowish base.

distribution: This species ranges from extreme

southwestern Ec ua dor, in El Oro and Loja provinces, to

northwestern Peru, in Lambayeque, Tumbes, and Piura de-

partments, from sea level to 2,300

m. Jessen et al. (2010)

reported the presence of this species (as S. stramineus) in

and around Lima, Peru, but considered these animals a

probable human introduction.

selected localities (Map 18): EC UA DOR:

El Oro, Arenillas (FMNH 53258), Río Pindo (AMNH

60474), Zaruma (type locality of Sciurus stramineus

zarumae J. A. Allen); Loja, Guainche [not located; see

Paynter 1993] (AMNH 61375), Los Posos (AMNH

67680), Malacatos (FMNH 54285). PERU: Laybayeque,

Chongoyape, Cabache (FMNH 80995); Piura, Cerro

Prieto, Lancones, Encuentros (FMNH 83437), Haci-

enda Bigotes, Salitral (FMNH 80908), Le Payta [= Paita]

(type locality of Sciurus nebouxii I. Geoffroy St.- Hilaire),

Monte Grande, 14 km N and 25 km E of Talara (MVZ

135638), near Talara, Pariñas Valley (FMNH 53864),

Palambla (AMNH 63701); Tumbes, Huasimo (FMNH

80991).

subspecies: We treat Simosciurus nebouxii as

monotypic. J. A. Allen (1915a:280) distinguished zaru-

mae, which he named from southwestern Ec ua dor, from

the nominotypical form from adjacent Peru on the basis

of dorsal and ventral color differences. Our examina-

tion of larger series fails to substantiate this geographic

segregation. Map 18 Selected localities for Simosciurus nebouxii (●). Contour line = 2,000 m.

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46 Mammals of South America

selected localities (Map 19): EC UA DOR:

Guayas, Chongoncito (AMNH 60548), Daulé (AMNH

34684), Guayaquil (type locality of Macroxus fraseri Gray);

Los Ríos, Hacienda Pijigal, Vinces (AMNH 62886), Pimo-

cha, Río Babahoyo (MNHN 1932/2893); Manabí, Balzar

Mountains, upper Río Palenque (type locality of Sciurus

stramineus guayanus Thomas); Tama [= Jama] (FMNH

53256); Santa Elena, Cerro Manglar Alto [= Manglaralto,

placed at Cerros de Colonche following Paint er 1993]

(AMNH 66640).

subspecies: We treat Simosciurus stramineus as

monotypic. However, J. A. Allen (1915a:281, 284) recog-

nized two subspecies among those geographic samples with-

out a nuchal patch, with guayanus Thomas “much lighter

colored throughout than true stramineus.”

natural history: Tirira (2007) summarized the

natural history attributes of this species; however, his map

of S. stramineus also included the range of S. nebouxii, so

his account possibly includes both species. Simosciurus

stramineus inhabits both primary and secondary forests

and disturbed areas, including both coffee and citrus plan-

tations where individuals cause damage to crops. The nat-

ural habitat is primarily dry forest, but the species can be

found in adjoining humid forests as well. This species is di-

urnal and arboreal, and in contrast with the solitary nature

of most other squirrels, this species may be found in groups

of three to ﬁ ve individuals. They feed on a variety of seeds

taken directly from trees but occasionally eat insects, fungi,

green leaves, and tree bark. Their activity is concentrated

in the early daylight hours. They take refuge in treetops, in

hollow logs, or in foliage. Their nests are made from leaves

and twigs.

remarks: P. Gervais (1841) described stramineus as a

member of the genus Sciurus. The name, however, has been

commonly attributed to the naturalists J. F. T. Eydoux and

L. F. A. Souleyet (e.g., J. A. Allen 1915a; Cabrera 1961;

Thorington and Hoffmann 2005; Thorington et al. 2012),

although these authors explicitly stated that Paul Gervais

both identiﬁ ed and described all mammals obtained dur-

ing the voyage of French corvette La Bonite (Eydoux and

Souleyet 1841:ii; 3, footnote).

P. Gervais (1841:39) gave the type locality of Simosci-

urus stramineus as “Omatopé, Pérou.” There is no locality

with this spelling in Peru; the locality referred to is most

likely Amotape, Piura, Peru, about 25 km northeast of the

port town of Paita (= Payta as given in Eydoux and Souleyet

1841). “Omatopé” (= Amotape) is also the type locality

of Eptesicus innoxius (P. Gervais 1841), collected on the

same expedition (see A. L. Gardner 2008:449). However,

Amotape is located in the center of the geographic range of

Simosciurus nebouxii I. Geoffroy St.- Hilaire, and all speci-

mens that we have examined of this species have the nuchal

Simosciurus stramineus (P. Gervais, 1841)

Guayaquil Squirrel

synonyms:

Sciurus stramineus P. Gervais, 1841:37, plate 9; type lo-

cality “Omatopé, ou Pérou” [= Amotape, Piura, Peru];

here corrected to Guayaquil, Guayas, Ec ua dor (see Re-

marks, below).

Macroxus stramineus: Lesson, 1842:112; name combi-

nation.

Macroxus fraseri Gray, 1867:430; type locality given as

“Republic of Ec ua dor.”

Sciurus hypopyrrhus: J. A. Allen, 1877:746; part (material

from Guayaquil, Ec ua dor).

Sciurus tephrogaster: J. A. Allen, 1877:763; part (Mac-

roxus fraseri Gray treated as a synonym).

Sciurus variabilis: J. A. Allen, 1877:768; part (Sciurus stra-

mineus P. Gervais treated as a synonym).

Sciurus (Echinosciurus) stramineus: E.- L. Trouessart,

1880b:202; name combination.

Sciurus stramineus guayanus Thomas, 1900a:150; type

locality “Balzar mountains, upper Río Palenque,

[Manabí,] Ec ua dor.”

Sciurus stramineus stramineus: Thomas, 1900a:151; name

combination.

Simosciurus stramineus stramineus: J. A. Allen, 1915a:281;

name combination.

Simosciurus stramineus guayanus: J. A. Allen, 1915a:283;

name combination.

Sciurus [(Guerlinguetus)] stramineus stramineus: Eller-

man, 1940:343; name combination.

Sciurus [(Guerlinguetus)] stramineus guayanus: Ellerman,

1940:343; name combination.

Simosciurus stramineus: Moore, 1959:204; name combi-

nation.

Sciurus (Guerlinguetus) stramineus: Cabrera, 1961:373; part.

Microsciurus (Simosciurus) stramineus: Moore, 1961a:15,

table 2; name combination.

Guerlinguetus (Simosciurus) stramineus: Moore, 1961b:15;

name combination.

description: Similar in size to S. nebouxii, with

head and body length 241– 327 mm (N = 40), tail length

250– 380 mm (N = 40), hindfoot length 50– 69 mm (N = 40),

ear length 20– 38 mm (N = 7). Body color brownish gray,

with posterior dorsum and posterior parts of forelegs

and hindlegs yellowish; crown, cheeks, and ears distinctly

darker than body; no nuchal patch; forefeet and distal parts

of forelimbs from darker than body to whitish gray; hind-

feet and distal parts of hind legs dark brown; venter brown-

ish gray; tail black and gray, except for the yellowish base.

distribution: Central Ec ua dor west of the Andes,

where it ranges from sea level to about 300 m, in Manabi,

Los Ríos, and Guayas provinces.

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Suborder Sciuromorpha: Family Sciuridae 47

Martín department, Peru, the ﬁ rst record of the genus for

South America. Syntheosciurus brochus occurs in evergreen

montane forests of the Cordillera Talamanca of Costa Rica

and Panama at elevations above 1,700 m. The Peruvian

specimen was taken at a single locality in Parque Nacional

Río Abiseo, an area covered by montane rainforest at eleva-

tions above 1,500 m on the eastern slope of the Andes (S.

C. Ramirez and Vela 2003). The large geographic hiatus

between the known localities of the Central American S.

brochus and the Peruvian record appears to be real, because

intervening areas in the northern Andes of Ec ua dor and

Colombia with the appropriate habitat and elevations have

been extensively collected, with large series of other taxa of

squirrels recorded with no Syntheosciurus taken.

Syntheosciurus can be distinguished from other South

American squirrels by the following combination of charac-

teristics: size intermediate between that of Microsciurus and

Guerlinguetus, with head and body length approximately

180 mm and tail length 150 mm; the single weight available

among specimens we examined was that of a adult male, re-

corded as 244 g. Externally, Syntheosciurus is distinguished

by its small ears, which are, on average, about 15 mm in

length. Females have three pairs of mammae. The temporal

portion of the orbitotemporal fossa is similar to that of the

larger South American squirrels, with the postorbital pro-

cess more or less equally dividing it from the orbital por-

tion. This is different from the condition in Microsciurus,

in which the temporal portion of the orbitotemporal fossa

is greatly reduced, and from that in Sciurillus, in which it is

obliterated. The incisors are clearly proodont, PM3 is pres-

ent, and the cristae of PM4 to M3 are oriented obliquely.

In the original description of the genus, Bangs (1902:25)

stated that the incisors have well- developed grooves on the

buccal surface; this characteristic is present in the holotype

of S. brochus but is absent in other specimens examined

(Goodwin 1943; Heaney and Hoffmann 1978).

synonyms:

Syntheosciurus Bangs, 1902:25; type species Syntheosci-

urus brochus, by monotypy.

Synthetosciurus Elliot, 1904:91; incorrect subsequent

spelling of Syntheosciurus Bangs.

Sciurus: Goodwin, 1943:1, part (description of poasensis).

Syntheosciurus (Syntheosciurus): Moore, 1959:179; name

combination.

Syntheosiurus Villalobos and Cervantes- Reza, 2007:31;

incorrect subsequent spelling of Syntheosciurus Bangs.

remarks: Phyloge ne tic relationships of Syntheosci-

urus to other American squirrels remain unclear, but Mer-

cer and Roth (2003) proposed that Syntheosciurus was a

sister group to Microsciurus, based on DNA sequence anal-

yses. This hypothesis agrees with the earlier, more superﬁ -

cial assessments of Bangs (1902) and Hall (1981). Moore

patches. But P. Gervais’s (1841:38– 39) detailed description

of color characteristics of his Sciurus stramineus includes no

mention of this very distinctive character. As can be seen in

Maps 18 and 19, the two species of Simosciurus we recog-

nize are allopatric, with their ranges apparently separated

by about a 150- km gap. Therefore, we believe that the lo-

cality of the type specimen was originally recorded in er-

ror or that some other mistake occurred during the voyage

and/or after arrival of the specimens in Paris. The ship Le

Bonite remained anchored at Paita for a few days and then

proceeded to Guayaquil, further north, where it stayed for

several more days before departing. Because Guayaquil is

an area inhabited by Simosciurus stramineus, we hereby

designate Guayaquil, Provincia de Guayas, Ec

ua dor as the

type locality of the species (ICZN 1999:Art. 76A).

P. Gervais (1841:38– 39) described the species as if he

had an actual specimen in hand, but the holotype could not

be located by one of us (MdV) in the Muséum National

d’Histoire Naturelle in Paris in 1992, nor has any other au-

thor reported on the holotype since the original description.

It may have been lost.

Genus Syntheosciurus Bangs, 1902

Syntheosciurus is a genus of small squirrels known until

now by the single species S. brochus from Central America.

We report what is likely an undescribed species from San

Map 19 Selected localities for Simosciurus stramineus (●). Contour

line = 2,000 m.

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48 Mammals of South America

ﬁ ed as Microsciurus sp., and it was similarly listed in recent

compilations of the mammal fauna of Peru (Pacheco et al.

1995, 2009). Several additional specimens may exist in Pe-

ruvian collections (V. Pacheco, pers. comm.).

Suborder Castorimorpha Wood, 1955

Infraorder Geomorpha Thaler, 1966

Superfamily Geomyoidea Bonaparte, 1845

James L. Patton

Superﬁ cially dissimilar in body form and habits, the pocket

gophers (Geomyidae) and kangaroo rats and pocket mice

(Heteromyidae) have been united as a recognizable mono-

phyletic assemblage for many de cades. All genera share ex-

ternal fur- lined cheek pouches, four cheek teeth (dental for-

mula I 1/1, C 0/0, PM 1/1, M 3/3 = 20) with a bilophodont

or more simpliﬁ ed occlusal pattern, and an infraorbital fora-

men positioned anterior to the zygomatic plate on the side

of the rostrum, opening anterolaterally in the Geomyidae,

(1959) included Syntheosciurus in his subtribe Microsciu-

rina, together with Microsciurus and Leptosciurus [= No-

tosciurus herein] (sensu J. A. Allen 1915a) and Simosciurus.

A phyloge ne tic hypothesis based on 57 morphological char-

acters (Villalobos and Cervantes- Reza 2007) placed Synthe-

osciurus in a group with other Central American taxa, and

separate from squirrels with more obvious South American

afﬁ nities such as Microsciurus and Notosciurus.

In keeping with our adoption of most of J. A. Allen’s

(1915a) arrangement for the genera of South American

Sciuridae, we employ Syntheosciurus as the generic name

for these animals. See Hall (1981) for a list of synonyms

and distributional rec ords for the Central American S.

brochus.

KEY TO THE SPECIES OF SYNTHEOSCIURUS:

1. Body color brown punctuated with orange; belly pale

orange; eye- ring present and conspicuous; no postau-

ricular patch . . . . . . . . . . . . . . .Syntheosciurus brochus

1′. Body color brown punctuated with ochraceous; belly

orange red; no eye- ring; postauricular patch present. . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . .Syntheosciurus sp.

Syntheosciurus sp.

Peruvian Montane Squirrel

synonyms: None.

description: In comparison to Central American S.

brochus, the Peruvian specimen has longer palmar pads;

belly tinted more reddish than orange; tail hairs fringed

with orange; dorsum brown streaked with ochraceous rather

than orange; no eye- ring, but postauricular patch present.

Skull shorter in length, and bullae have two well- separated

rather than closely appressed and parallel septa.

distribution: This species is only known from

a single locality in Andean montane forest at 2,800 m in

northern Peru.

selected localities (Map 20): PERU: San Mar-

tín, Vilcabamba del Pajate, 31 km NE of Pataz [La Liber-

tad] (USNM [MML 279]).

subspecies: None.

natural history: Nothing is known about the

ecol ogy or other aspects of the natural history of this ani-

mal. The single known locality is in upper montane forest

(S. C. Ramirez and Vela 2003).

remarks: This species is here reported thanks to the

late Dr. Charles O. Handley Jr., then at the Smithsonian In-

stitution, National Museum of Natural History, Washington

D. C., who showed a single specimen to one of us (MdV).

Mariella Leo collected this specimen in 1989 during an in-

ventory in the Parque Nacional del Río Abiseo (see S. C.

Ramirez and Vela 2003). In that report, made to support

a management plan for the park, the specimen was identi-

Map 20 Single known locality for Syntheosciurus sp. (●). Contour line = 2,000 m.

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Trypanosoma cruzi in domestic and wild mammals in the northeast region of Colombia

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Rodentia Life History

Chapter

May 2022

Gisela Sobral

264 km N of Xavantina (BM 79

203-207

Mato Grosso
Serra Do Roncador

Mato Grosso, Serra do Roncador, 264 km N of Xavantina (BM 79.203– 207);

Gradaús, right bank Rio Fresco (MZUSP 20399), mouth of Rio Bacajá, right bank Rio Xingú (MZUSP 25455

Belém Pará

Pará, Belém (MPEG 710), Gradaús, right bank Rio Fresco (MZUSP 20399), mouth of Rio Bacajá, right bank Rio Xingú (MZUSP 25455), Portel, Rio Procupi (MPEG 606), Rio Riosinho, left bank Rio Fresco (MPEG 1039),

São Lourenço (type locality of Sciurus Roberti Thomas)

Pernambuco

Pernambuco, São Lourenço (type locality of Sciurus Roberti Thomas);

type locality of Sciurus ingrami sebastiani P. Müller and Vesmanis

São Paulo
Ilha De
São Sebastião

São Paulo, Ilha de São Sebastião (type locality of Sciurus ingrami sebastiani P. Müller and Vesmanis ), Ilha do Cardoso (MZUSP 3905), Porto Cabral, Rio Paraná (MZUSP 6021), Presidente Epitácio, left bank Rio Paraná (MZUSP 3733), Santos (MZUSP 3699), Serra da Juréia (MZUSP 12835). s u b s p e c i e s : Guerlinguetus brasiliensis is polytypic:

Microsciurus fl aviventer 2′. Dorsum dark brown, with mid-dorsal dark stripe Microsciurus santanderensis

Red.............................................................. Dorsum Brown

Dorsum brown, streaked with red................ ........................ Microsciurus fl aviventer 2′. Dorsum dark brown, with mid-dorsal dark stripe.... ..................... Microsciurus santanderensis

Microsciurus isthmius 4′. Venter washed with red on throat and chest; mixed brown

Venter Orange
................................. Mixed With Brown On Belly

Venter orange, mixed with brown on belly.......... .......................... Microsciurus isthmius 4′. Venter washed with red on throat and chest; mixed brown, dark gray, and red on belly........Microsciurus otinus

Estirão do Equador, right bank Rio Javari (MZUSP 20350), Lago do Mapixi, east of Rio Purus (FMNH 38880); Rondônia, Cachoeira Nazaré, left bank Rio Ji-Paraná (MPEG 20826) COLOM- BIA: Caquetá, Florencia (type locality of M

Yensen Bravo

Bravo, Yensen et al. 2002). BRAZIL: Acre, Oriente (= Seringal Oriente), near Taumaturgo, Rio Juruá (MZUSP 20351); Amazonas, Acajutuba, Rio Negro (type locality of M. manarius Thomas), Codajás, left bank Rio Solimões (MZUSP 4207), Eirunepé, left bank Rio Juruá (MZUSP 4621), Estirão do Equador, right bank Rio Javari (MZUSP 20350), Lago do Mapixi, east of Rio Purus (FMNH 38880), Manacapuru, Rio Solimões (BM 26.5.5.36), Paraná do Aiapuá, Rio Purus (BM 27.8.11.61); Rondônia, Cachoeira Nazaré, left bank Rio Ji-Paraná (MPEG 20826). COLOM- BIA: Caquetá, Florencia (type locality of M. fl orenciae J. A. Allen); Putumayo, La Tagua, Tres Troncos, Río Caquetá (FMNH 71118); Vaupés, Río Vaupés, in front of Tahuapunto (AMNH 78625). EC UA DOR: Loja, El Porotillo, San José (MNHN 1934/1210);

AMNH type locality of Sciurus (Microsciurus) peruanus J. A. Allen), near Huampani, Río Cenepa (MVZ 154929), 12 km E of La Peca Nueva (LSUMZ 21876); Cusco, Quincemil (FMNH 75210); Junin, Chanchamayo (type locality of Microsciurus rubrirostris

Jan 1953
275

Guayabamba Amazonas

PERU: Amazonas, Guayabamba (AMNH type locality of Sciurus (Microsciurus) peruanus J. A. Allen), near Huampani, Río Cenepa (MVZ 154929), 12 km E of La Peca Nueva (LSUMZ 21876); Cusco, Quincemil (FMNH 75210); Junin, Chanchamayo (type locality of Microsciurus rubrirostris J. A. Allen); Loreto, boca Río Curaray (AMNH Sciurus (Guerlinguetus) granatensis bondae: Goodwin, 1953: 275; name combination.

La Guajira,] Colombia (altitude 8000 ft [2,000 ft; see Paynter 1997:139]) Sciurus saltuensis bondae J. A. Allen, 1899b:213, type locality

Jan 1904
185431326

Sciurus variabilis saltuensis Bangs, 1898b:185, type locality "Pueblo Viejo [= El Pueblito], [north slope Sierra Nevada de Santa Marta, La Guajira,] Colombia (altitude 8000 ft [2,000 ft; see Paynter 1997:139])." Sciurus saltuensis bondae J. A. Allen, 1899b:213, type locality "Bonda, Santa Marta District, [Magdalena,] Colombia." Sciurus saltuensis: J. A. Allen, 1904c:431; name combination. Sciurus variabilis gerrardi: E.-L. Trouessart, 1904:326; name combination.

1910:26; type locality "Orope, Zulia, Venezuela Guerlinguetus hoffmanni quindianus J. A. Allen, 1914d: 587, type locality "Rio Frio (central Rio Cauca Valley, altitude 3500 feet), western slope of Central (or Quindio) Andes

Zuliae Sciurus Versicolor
Osgood

Sciurus versicolor zuliae Osgood, 1910:26; type locality "Orope, Zulia, Venezuela." Guerlinguetus hoffmanni quindianus J. A. Allen, 1914d: 587, type locality "Rio Frio (central Rio Cauca Valley, altitude 3500 feet), western slope of Central (or Quindio) Andes," Magdalena, Colombia.

Family Sciuridae G. Fischer, 1817

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