Revisionary Notes on Neotropical Porcupines (Rodentia: Erethizontidae). 2. A Review of the Coendou vestitus Group with Descriptions of Two New Species from Amazonia

36 Pages·2001·1.4 MB·English

by  Robert S. Voss

PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY
CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024
Number 3351, 36 pp., 15 figures, 5 tables October 31, 2001
Revisionary Notes on Neotropical Porcupines
(Rodentia: Erethizontidae). 2. A Review of the
Coendou vestitus Group with Descriptions of Two
New Species from Amazonia
ROBERT S. VOSS1 AND MARIA N. F. DA SILVA2
CONTENTS
Abstract ....................................................................... 2
Resumen ....................................................................... 2
Resumo ........................................................................ 2
Introduction .................................................................... 2
Materials and Methods ........................................................ 3
Species Accounts ............................................................... 6
Coendou vestitus Thomas ...................................................... 6
Coendou pruinosus Thomas ................................................... 11
Coendou ichillus, new species ................................................. 17
Coendou roosmalenorum, new species ......................................... 24
Discussion .................................................................... 30
Acknowledgments ............................................................. 32
References .................................................................... 32
Appendix: Gazetteer ........................................................... 34
1DivisionofVertebrateZoology (Mammalogy),AmericanMuseumofNaturalHistory.E-mail:[email protected]
2InstitutoNacionaldePesquisasda Amazoˆnia,CP478, Manaus,AM69083,Brazil.E-mail:[email protected]
Copyright(cid:113)AmericanMuseumofNaturalHistory2001 ISSN0003-0082
2 AMERICAN MUSEUM NOVITATES NO. 3351
ABSTRACT
Neotropical porcupines of the Coendou vestitus group consist of four small-bodied species
from northwestern South America that are hypothesized to form a clade on the basis of their
unique dorsal pelage composition. In addition to soft wool and barbed quills—hair types
widely shared by other erethizontids—the dorsal pelage of vestitus-group porcupinesincludes
what may be called bristle-quills: long, thin, unbarbed quills with flexible tips. The group
includes Coendou vestitus Thomas, 1899; C. pruinosus Thomas, 1905; and two new species.
The latter provide the first documented records of small porcupines from western Amazonia,
where only large porcupines (C. prehensilis and C. bicolor) were previously known. Other
small porcupines probably remain to be discovered in Amazonia, where hydroelectric dam
projects offer unique (if unfortunate) faunal-sampling opportunities.
RESUMEN
El grupo de puercoespines neotropicales Coendou vestitus consiste en cuatro especies pe-
quen˜as del noroeste de Ame´rica del Sur, consideradas parte de un clado con base en la com-
posicio´n particular de su pelaje dorsal. Adema´s de lana suave y espinas con pu´as—tipos de
pelo compartidos ampliamente con otros eretizo´ntidos—el pelaje dorsal de los puercoespines
del grupo vestitus tiene lo que podr´ıa llamarse cerda-espinas: espinas largas, delgadasy ralas,
con puntas flexibles. Este grupo incluye a Coendou vestitus Thomas, 1899; C. pruinosus
Thomas, 1905; y dos nuevas especies. Estas u´ltimas constituyen los primeros registros docu-
mentados de puercoespines pequen˜os en la Amazonia occidental, donde previamente so´lo se
conoc´ıan puercoespines grandes (C.prehensilis yC.bicolor).Sediscutelaposibilidaddeque
otros puercoespines pequen˜os este´n por descubrirse en la Amazonia y se enfatizan las opor-
tunidadesu´nicasdemuestreodefaunaprovistasporlaconstruccio´nderepresashidroele´ctricas.
RESUMO
Porcos-espinhos neotropicais do grupo Coendou vestitus sa˜o representados por quatro es-
pe´cies de pequeno porte, que hipotetizamos formar um clado, tendo por base a composic¸a˜o
u´nica de sua pelagem. Ale´m da pelagem lanosa e dos espinhos farpados—tipos de peˆlos
amplamente compartilhados por outros eretizont´ıdeos—a pelagem dorsal dos porcos-espinhos
dogrupovestitusincluemoquepodemoschamardecerdasespinhosas:longos,finos,espinhos
na˜o-farpadosdepontasflex´ıveis.OgrupoincluiCoendouvestitusThomas,1899;C.pruinosus
Thomas, 1905; e duas novas espe´cies. Estas fornecem o primeiro registro documentado de
porcos-espinhos de pequeno porte no oeste da Amazoˆnia, onde apenas porcos-espinhos de
grande porte (C. prehensilis e C. bicolor) eram anteriormente conhecidos. Discutimosaprob-
abilidade de que outras espe´cies de porcos-espinhos de pequeno porte sejam descobertas no
oeste amazoˆnico, e enfatizamos a importaˆncia u´nica de amostragens faun´ısticas criadas pela
construc¸a˜o de barragens hidroele´tricas.
INTRODUCTION bristle-quills(fig.1).Thelatteraremuchlon-
ger and thinner than conventional quills, but
Conspicuous among the Neotropical pre-
have the same distinctive root morphology
hensile-tailed porcupines currently referred
(described and illustrated by Chapman and
tothegenusCoendou(includingSphiggurus;
Handley and Pine,1992)isagroup ofsmall- Roze,1997),roundcrosssection,stiffcortex,
bodied species whose dorsal pelage contains and spongy internal matrix; unlike regular
three intermingled but distinctly different quills, however, the tips of bristle-quills are
structural types: soft fur, quills (defensive attenuated, flexible, wire-like filaments, and
spines with hard, sharp, barbed tips), and an they are not provided with microscopic
unusual structural type that may be called barbs.
2001 VOSS AND DA SILVA: NEOTROPICAL PORCUPINES 3
titus Thomas (1899), originally described on
thebasisof asinglespecimenfromColombia
purchased by the British Museum in 1854.
Thereafter, the species remained unknown
fromadditionalmaterialuntiltheearly1920s,
when Hermano Nice´foro Mar´ıa obtained sev-
en specimens in the western foothills of the
eastern Andes near Bogota´. Unfortunately,no
otherrecordsofthisobviouslyuncommon(or
elusive) porcupine have been reported in the
last 75 years, and it is not known whether or
not the species is still extant.
A second taxon, closely resemblingCoen-
dou vestitus but diagnosably different in col-
oration and craniodental characters, was de-
scribed as C. pruinosus by Thomas (1905).
Thomas’s type series consisted of five spec-
imens obtained between 2500 and 2600 m
elevation in the Venezuelan Andes near Me´-
rida, and most subsequently collected mate-
rial assignable to pruinosus is likewise from
montaneorfoothillhabitatsinVenezuelaand
Fig. 1. Three structural types found in the
dorsalpelageofmembersoftheCoendouvestitus Colombia. Although some authors (e.g., Ca-
speciesgroup:a,softwoolhairs;b,quills;c,bris- brera, 1961; Honacki et al., 1982; Concep-
tle-quills. Approximately life size (drawn from cio´n and Molinari, 1991; Woods, 1993; So-
pelage of AMNH 126171, holotype of C. ichil- riano and Ochoa, 1997) have regarded prui-
lus).
nosus as a subspecies or junior synonym of
vestitus,others(e.g.,Handley,1976;Handley
and Pine, 1992; Linares, 1998; Alberico et
al., 1999) have treated these taxa as valid
species. However, no substantive discussion
Although some species of Coendou have of character data has been provided to sup-
been incorrectly described as covered by port either usage.
quills only (e.g., C. prehensilis, C. bicolor; Herein we redescribe Coendou vestitus
Ellerman, 1940), a sparse underfur of short and C. pruinosus based on our examination
wool hairs is always present among the quill of typesand otherspecimensinNorthAmer-
bases of such forms. Other species of Coen- ican and European museums. Clarifying the
dou are superficially coveredbylongfur,but diagnostic characteristics of vestitus and
these taxa always possess an underlying de- pruinosus is necessary in order to diagnose
fensivecoatofsharpquills.Infact,allCoen- two new species that significantly extendthe
dou species possess both soft fur and quills; ecogeographic distribution of vestitus-like
the distinctive feature of the species treated porcupines into the lowland rainforests of
in this report consists in the additional pres- Amazonia. Unanswered questions concern-
ence of thin bristle-quills. Because the latter ing erethizontid distributions in Amazonia
are not known to occur among any other er- are briefly considered in a concluding dis-
ethizontids,thesimplestexplanationfortheir cussion. This paper is the second in a series
taxonomic distribution is that they evolved of preliminary reports (initiated by Voss and
uniquely in the common ancestor of the Angermann, 1997) with the object of resolv-
group of species described below. ing key taxonomic problems in advance of a
The oldest name for any Neotropical por- comprehensive revision of Neotropical por-
cupinewiththinbristle-quillsisCoendouves- cupines currently in preparation.
4 AMERICAN MUSEUM NOVITATES NO. 3351
MATERIALS AND METHODS alveolar margin to the crown of the ipsilat-
eral P4 [dP4 of subadults].
SPECIMENS: Specimens that we examined LIF, Length of Incisive Foramen: Inside
and measured for this study—and others re- length of one incisive foramen (or both, if
ferred to in the text—are in the American the foramina are recessed in a common fos-
Museum of Natural History, New York sa).
(AMNH);theNaturalHistoryMuseum,Lon- BIF, Breadth of Incisive Foramina: Inside
don (BMNH); the Coleccio´n de Vertebrados breadth across both incisive foramina.
de la Universidad de los Andes, Me´rida MTR, Maxillary Tooth Row: Greatest
(CVULA); the Museo de la Estacio´n Biolo´- crown length from P4 [dP4 of subadults] to
gica de Rancho Grande, Maracay (EBRG); M3.
the Departamento de Biolog´ıa de la Escuela LM, Length of Molars: Greatest crown
Polite´cnica Nacional, Quito (EPN); the Field length of the upper molar series (M1–M3).
Museum of Natural History, Chicago BP4, Breadth of P4: Greatest crown
(FMNH); the InstitutoNacionaldePesquisas breadth of the permanent upper premolar.
daAmazoˆnia,Manaus(INPA);theMuseode BM1, Breadth of M1: Greatest crown
Biolog´ıa de la Universidad Central de Ven- breadth of the first upper molar.
ezuela, Caracas (MBUCV); the Museum of APB, Anterior Palatal Breadth: Measured
ComparativeZoologyofHarvardUniversity, betweenthecrownsofthefirstuppermolars.
Cambridge (MCZ); the Museo de Historia PPB, Posterior Palatal Breadth: Measured
Natural La Salle, Caracas (MHNLS); the between the crowns of the third upper mo-
Muse´um National d’Histoire Naturelle,Paris lars.
(MNHN); the National Museum of Natural PZB, Posterior Zygomatic Breadth:Great-
History, Washington, D.C. (USNM); and the est breadth across the zygomatic arches be-
Museum fu¨r Naturkunde der Humboldt- hind the orbits.
Universita¨t zu Berlin (ZMB). HIF, Height of the Infraorbital Foramen:
MEASUREMENTS: All reported measure- Measured as the greatest inside diameter,
ments are in millimeters (mm) and all re- usually at an angle of about 30–40(cid:56) fromthe
ported weights are in grams (g). Because midsagittal plane.
head-and-body length (HBL) and length of ZL, Zygomatic Length: Measured from
tail (LT) were seldom recorded by collectors the posterior margin of the infraorbital fora-
for the specimens treated in this report, we men to the posterolateral corner of the zy-
obtained approximate values (prefixed by gomatic arch.
‘‘ca.’’) for these dimensions by measuring LN, Length of Nasals: Greatest length of
dried skins to the nearest 5 mm with a flex- one nasal bone (the longest if right and left
ible rule. Length of the hindfoot (HF), re- elements are unequal).
corded to the nearest millimeter,includesthe BNA, Breadth of Nasal Aperture:Greatest
claws (c.u.) and was either measured by col- transverse dimension of the nasal orifice, al-
lectors in the field and/or remeasured by us ways at or near the nasal/premaxillary su-
on dried skins (only minimally distorted tures.
dried hindfeet containing the intact pedal BB, Breadth of Braincase: Transverse di-
skeleton wereremeasured).Measurementsof mension of the braincase, measured by plac-
the skull and dentition, taken with dial or ingthecaliperjawsjustabovethesquamosal
digital calipers and recorded to the nearest zygomatic root on each side.
0.1 mm, are abbreviated and defined as fol- DI,DepthofIncisor:Distancebetweenthe
lows (see fig. 2): greater and lesser curvatures of an upper
CIL, Condylo-incisive Length: Measured tooth.
from the articular surface of one occipital BIT,BreadthoftheIncisorTips:Measured
condyle to the greater curvature of the ipsi- across the enameled tips of both upper teeth.
lateral upper incisor. AGE CLASSIFICATION: We used maxillary
LD, Length of Diastema: Measured from tooth eruption, cranial suture closure, and
thelessercurvatureofanupperincisoratthe pelage maturation to define a heuristic age
2001 VOSS AND DA SILVA: NEOTROPICAL PORCUPINES 5
Fig. 2. Anatomical limits of 18 craniodental measurements defined in the text.
6 AMERICAN MUSEUM NOVITATES NO. 3351
classification as follows (after Voss and An- foot; the left hindfoot is detached but intact.
germann, 1997). The skull lacks the occiput, a common con-
Juveniles—Maxillary toothrow incom- sequence of early to mid-19th century spec-
plete (three or fewer teeth erupted); all cra- imen preparation methods.Theoriginallabel
nialsuturesopen;pelageoftenconspicuously gives the place of origin only as‘‘N[ouvel]le
immature, including long fur even in species Grenade’’ ((cid:53) Colombia). According to
that lack visible fur as adults. Thomas (1899), the specimen waspurchased
Subadults—Immature maxillary dentition in 1854.
(dP4–M3) completely erupted, or dP4 shed GEOGRAPHIC DISTRIBUTION: Coendou ves-
and P4 incompletely erupted; all cranial su- titus is only known from two definite local-
tures still visible; pelage always appearsma- ities, both of which are in the western foot-
ture. hills of the eastern Andean cordillera of Co-
Adults—Permanent maxillary dentition lombia. The first, where Hermano Nice´foro
(P4–M3) fully erupted, with light to moder- Mar´ıa collected six specimens from 1923 to
ate wear (teeth usually not worn below wid- 1925, is San Juan de R´ıo Seco, a small vil-
est part of crown and almost always with at lage located about 60 km WNW of Bogota´
least some occlusal detail remaining); some
at ca. 1300 m elevation. The second, where
cranial sutures usually obliterated.
the same collector took a single specimen in
Old adults—Cheekteeth worn below wid-
1925, is Quipile, a village only about 15 km
est part of crown (and therefore not measur-
SSE of San Juan and at approximately the
able),with littleornoocclusaldetailremain-
same altitude.
ing on M1 and M2; only nasal sutures (if
DESCRIPTION: External—The dorsal fur,
any) usually visible.
soft in texture and dull in appearance,isuni-
formly blackish brown and averages about
SPECIES ACCOUNTS
50–60mmmiddorsally;thefurisshorterand
The specimens examined for this report sparseralongtheflanks,butitislongenough
can be sorted into four distinct groups on the to conceal most of the quills except on the
basis of external and osteological characters. face. The quills (ranging in length from
Whereas two of these groups have available about 25 to 35 mm middorsally) are bicol-
names based on extant holotypes, the others ored, pale yellow or ivory-white basally and
are unnamed. The two named forms are re- dark-brown or blackish distally; usuallyonly
described below in chronological order of the distalmost 1⁄ or less of each quill is dark,
theirpublication,followedbyaccountsofthe so the pale quill bases are exposed when the
new species. Specimen records are mapped fur is parted or ruffled. None of the quills
and documented by geographic coordinates anywhere on the body is pale-tipped. Scat-
in the appendix. tered abundantly but inconspicuously
throughoutmostofthedorsalpelagearelong
Coendou vestitus Thomas (to 70–80 mm) polished bristle-quills that
Figures3, 4,5A,6A emerge from thefur like finewires;theseare
yellowish basally, but the emergent tips are
CoendouvestitusThomas,1899:284(originalde-
dark. No bristle-quills occur over the rump,
scription).
Coendou (Sphiggurus) vestitus: Tate, 1935: 307 however, which is clothed only by short,
(new name combination). sharp quills and fur. The ventral surface of
Coendou (Sphiggurus) vestitus vestitus: Cabrera, the body is densely covered with soft and
1961: 603 (new name combination). uniformly dark-brown fur from chin to anus.
Sphiggurus vestitus: Honacki et al., 1982: 572 Two classes of hairs can be distinguished in
(new name combination).
the ventral fur (fine, wavy wool hairs and
TYPE MATERIAL: The holotype only, coarser, straighter guard hairs), but there are
BMNH 54.6.26.1, consisting of the skin, no conspicuously thickened spinous hairs
skull, and mandibles of a subadult animal of grouped in triads or other clusters.
unknown sex. The skin, much faded with The tail is short, apparently averaging
age, is understuffed and lacks the left fore- about half the length of the combined head-
2001 VOSS AND DA SILVA: NEOTROPICAL PORCUPINES 7
Fig. 3. Dorsal view of the skin of Coendou vestitus (BMNH 24.2.21.2). Approximately (cid:51)0.4.
8 AMERICAN MUSEUM NOVITATES NO. 3351
Fig.4. Dorsal,ventral,andlateralcranialviewsofCoendouvestitus(MNHN1929–632).Allviews
approximately (cid:51)1.5.
2001 VOSS AND DA SILVA: NEOTROPICAL PORCUPINES 9
Fig.5. VentralcranialviewsofCoendouvestitus(A,MNHN1929–632)andC.pruinosus(B,MCZ
18738)showingspeciesdifferencesinpalatalmorphology.InCoendouvestitus,thebonypalatebetween
the toothrows is marked by a high median keel that is flanked by deep lateral grooves or gutters (lg);
in addition, the mesopterygoid fossa (mpf) extends anteriorly between the second molars, and the roof
ofthefossaisperforatedbylargesphenopalatinevacuities(spv).InC.pruinosus,thepalateissometimes
weakly keeled but never has deep lateral gutters; also, the mesopterygoid fossa extends anteriorly only
betweenthethirdmolars,andtheroofofthefossaisnotperforatedbydistinctsphenopalatinevacuities.
and-body (see measurements in table 1).The Skull—The frontal and nasal sinuses are
proximal half of the tail is clothed dorsally uninflated, resulting in a flat dorsal profile
with quills and woolly fur like therump,and from the nasal tips to the midparietal region
the tip has a naked, calloused dorsal prehen- in all the specimens examined. The rostrum
sile surface, but the rest of the tailiscovered is short and tapering in younger animals,but
above and below with blackish bristles; the old adults have proportionately longer ros-
bristles under the base of the tail are much trums with prominent lateral excavations for
stiffer and denser than those on the lateral the origin of the infraorbital muscle. The na-
anddorsalsurfaces.Thedorsalsurfaceofthe salbonesaremore-or-lessparallel-sided,nei-
hands and feet are densely covered with ther increasing nor decreasing posteriorly in
coarse brownish hairs. breadth, with rounded posterior margins that
Thelongmystacialvibrissaeareuniformly extend well behind the premaxillae. Viewed
blackish and extend behind the pinnae when from above, the zygomatic arches converge
laid back alongside the head. Supraorbital anteriorly from their widest pointat thelevel
(superciliary),genal,submental,andpostcra- of the squamosal roots with only a slightlat-
nial vibrissae are also present. The postcra- eral deflection at the level of the orbits. The
nial vibrissae occur on the forelimb between jugals are slender elements, lackinganycon-
the elbow and wrist, on the hindlimb be- spicuous postorbital expansion. The dorso-
tweenthekneeandankle,andalongtheven- lateral contours of the braincase are strongly
tralsurface ofthebodybetweentheforelimb sculpted by the origin of thetemporalismus-
and hindlimb. cle, but the left and right temporal scars are
10 AMERICAN MUSEUM NOVITATES NO. 3351
Fig. 6. Ventrolateral cranial views of Coendou vestitus (A, AMNH 71360) and C. pruinosus (B,
MCZ 18738) showing species differences in alisphenoid morphology. In Coendou vestitus, the alisphe-
noid is incompletely ossified, resulting in an open sphenopterygoid canal and a buccinator-masticatory
foramen(bmf)thatisconfluentwiththeforamenovale(fo).Bycontrast,thealisphenoidisfullyossified
in C. pruinosus, whose sphenopterygoid canal (arrow) is laterally enclosed and whose buccinator-mas-
ticatory and oval foramina are separate.
widely separatedanddonotjoinmiddorsally are small (ca. 14–15 mm) rounded capsules
to form a sagittal crest. that are well separated from the base of the
The small incisive foramina are complete- paroccipital process on each side. The dorsal
ly contained in the premaxillae (e.g.,BMNH roof of the external auditory meatus has an
24.2.21.2) or shallowly contact the maxillae indistinct bony ridge that is less well devel-
(e.g., AMNH 71359), but do not deeplypen- oped than that seen in some congeneric taxa
etrate between the latter bones; the left and (e.g., Coendou melanurus; Voss et al., 2001:
right foramina are incompletely separated fig. 70A).
and are recessed in a common fossa in some
The mandible is distinctive in the absence
specimens, but in others they are completely
of a well-defined coronoid process, which is
separated and are not recessed together. The
represented only as a rounded convexity at
posterior diastema is marked by shallow and
the base of the ascending ramus in all spec-
widely separated lateral sulci. In all of the
imens examined.
specimens examined, the palatal bridge (be-
Dentition—The upper incisors have pale
tween the toothrows) has a well developed
yellow-orange enamel bands and are mod-
centralkeelanddeeplateralgutters(fig.5A).
erately procumbent. The cheekteeth essen-
The mesopterygoid fossa penetrates anteri-
tially resemble those of other erethizontids
orly to or between the second molars, and
(except Chaetomys) in occlusal morphology,
the bony roof of the fossa is perforated by
well-formed sphenopalatine vacuities ((cid:46)1 and the toothrows are subparallel to weakly
mm in diameter) in all of the specimens at convergent. The permanentfourthupperpre-
hand. The alisphenoid is incompletely ossi- molarisonlyslightlylargerthanthefirstmo-
fied, with the result that the sphenopterygoid lar in some specimens (e.g., AMNH 70529),
canal is open laterally and the buccinator- but P4 is conspicuously larger than M1 in
masticatoryforamenisconfluentwiththefo- others (e.g., USNM 240035).
ramen ovale (fig. 6A). The auditory bullae COMPARISONS: See the accounts for Coen-