Zootaxa 3636 (3): 421–438 ISSN 1175-5326 (print edition)

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Copyright © 2013 Magnolia Press
Article ZOOTAXA
ISSN 1175-5334 (online edition)
http://dx.doi.org/10.11646/zootaxa.3636.3.2
http://zoobank.org/urn:lsid:zoobank.org:pub:FE435AD6-66A4-46D8-95D3-7BFFD2ED28F1

A new species of porcupine, genus Coendou (Rodentia: Erethizontidae)

from the Atlantic forest of northeastern Brazil

ANTONIO ROSSANO MENDES PONTES1, JOSÉ RAMON GADELHA1, ÉVERTON R. A. MELO1,

FABRÍCIO BEZERRA DE SÁ2, ANA CAROLINA LOSS3, VILACIO CALDARA JUNIOR3,
LEONORA PIRES COSTA3 & YURI L. R. LEITE3
1
Universidade Federal de Pernambuco, Centro de Ciências Biológicas, Departamento de Zoologia, Laboratório de Estudo e Conser-
vação da Natureza. Rua Prof. Moraes Rego, 1235, Cidade Universitária, CEP: 50.740-620, Recife, PE, Brazil.
E-mail: mendespontes@gmail.com
2
Universidade Federal Rural de Pernambuco, Departamento de Morfologia e Fisiologia Animal, Laboratório de Anatomia. Rua Dom
Manoel de Medeiros, S/N, Dois Irmãos, CEP: 52.171-900, Recife, PE, Brazil
3
Universidade Federal do Espírito Santo, Departamento de Ciências Biológicas, Laboratório de Mastozoologia e Biogeografia. Av.
Marechal Campos, 1468, Maruípe, CEP: 29.043-900, Vitória, ES, Brazil

Abstract

We report the discovery of a new species of Coendou (Rodentia, Erethizontidae), here designated Coendou speratus sp.
nov. This small porcupine, locally known as coandu-mirim, is found in the Pernambuco Endemism Centre in the Atlantic
coast of northeastern Brazil north of the São Francisco river, one of the most important known biodiversity hotspots. The
geographic range of C. speratus overlaps with that of the larger, widespread C. prehensilis, but not with that of C. insidio-
sus from the southeastern Atlantic forest, nor with that of C. nycthemera, an eastern Amazonian species. Coendou speratus
is a small-bodied, long-tailed species that appears to be completely spiny because it lacks long dorsal fur. The dorsal quills
have conspicuously brownish red tips that contrast with the blackish dorsal background color. The new species is overall
similar to C. nycthemera, but the dorsal body quills are typically tricolored in the former and bicolored in the latter. The
new species is externally very distinct from C. insidiosus, especially because the latter has bicolored dorsal quills that are
almost completely hidden beneath longer and homogeneous pale or dark hairs.

Key words: Biodiversity hotspot, Coendou, Mammalia, Neotropics, taxonomy

Introduction

The Brazilian Atlantic forest is considered a world priority for biodiversity conservation and the fourth most
important hotspot of the planet (Myers et al. 2000). The Pernambuco Endemism Centre, a biogeographical unit of
the Brazilian Atlantic forest that lies north of the São Francisco river, harbors several endemic species, which
suggests that it is a hotspot within a hotspot (Punde et al. 2008; Carnaval et al. 2009). Unfortunately, this region has
so far lost 98% of its natural forest cover (Viana et al. 1997).
New World porcupines (Rodentia: Erethizontidae) are small- to medium-sized arboreal rodents that feed on
immature seeds, green or ripe fruit, palm nuts, bark and leaves (Emmons & Feer 1997; Lima et al. 2010; Passamani
2010). They currently comprise 15 species (Voss 2011), belonging to at least three genera (Coendou, Erethizon,
Chaetomys), depending on the author. Some authors recognize the hairy Sphiggurus as distinct from the spiny
Coendou (e.g., Husson 1978; Eisenberg & Redford 1999; Nowak 1999; Bonvicino et al. 2002; Woods & Kilpatrick
2005), whereas others consider Sphiggurus a junior synonym of Coendou (e.g., Handley & Pine 1992; Emmons &
Feer 1997; McKenna & Bell 1997; Voss & Angermann 1997; Voss 2011), arguing that they cannot be meaningfully
diagnosed as separate taxa. Here we follow this latter view. The number of species of Neotropical erethizontids is
still tentative, since species limits are poorly defined (Voss 2011), the application of some old names is uncertain
(Leite et al. 2011), and new species have been recently described (Voss & Silva 2001). The new species described

Accepted by M. Weksler: 18 Feb. 2013; published: 5 Apr. 2013 421

below had not been detected before in any of several sites recently surveyed in the region, and we found no
reference to this small porcupine in historical accounts, such as Marcgrave’s (1648), who carefully described this
regional fauna, including the larger porcupine, Coendou prehensilis Linnaeus (Leite et al. 2011).

Material and methods

Study area and specimens. The biodiversity inventories that resulted in the discovery here reported have been
carried out since January 2002, through the line transect method (Buckland et al. 1993), adapted from Mendes
Pontes (2004), and have surveyed 30 forest fragments of the Pernambuco Endemism Centre (sensu Prance 1982;
1987; Silva & Casteletti 2003), or Pernambuco refugium (Punde et al. 2008; Carnaval et al. 2009), in the Atlantic
forest of northeastern Brazil (Figure 1). The studied forest fragments range in size from 4 to 3,642 ha (Silva Jr. &
Mendes Pontes 2008; Asfora & Mendes Pontes 2009) and are all subject to some degree of human interference
(such as hunting, feral dogs, forest clearing, selective logging, fires, etc.) and are cut through by roads used by
people and domestic animals.
The description of the new porcupine, locally known as coandu-mirim, is based on the analysis of five
individuals deposited at Museu Nacional, Universidade Federal do Rio de Janeiro, Brazil (MN), the mammal
collection of the Universidade Federal de Pernambuco, Recife, Brazil (UFPE), and the mammal collection of the
Universidade Federal do Espírito Santo, Vitória, Brazil (UFES). We also examined specimens of Coendou spp.
from the Museu de Zoologia, Universidade de São Paulo, Brazil (MZUSP) and Museu de Biologia Prof. Mello
Leitão, Santa Teresa, Brazil (MBML).
Measurements and morphometric analyses. All measurements are in millimeters, and weights are in grams.
Cranial and dental measurements were taken with digital calipers to the nearest 0.1 mm following Voss and Silva
(2001): LN = length of nasals; BB = breadth of braincase; LD = length of diastema; PZB = posterior zygomatic
breadth; APB = anterior palatal breadth; PPB = posterior palatal breadth; CIL = condylo-incisive length; ZL =
zygomatic length; BNA = breadth of nasal aperture; HIF = height of infraorbital foramen; LIF = length of incisive
foramen; BIF = breadth of incisive foramen; BIT = breadth of incisor tips; DI = depth of incisor; MTR = maxillary
toothrow length; LM = length of molars; BP4 = breadth of fourth pre-molar; BM1 = breadth of first molar. Three
additional cranial measurements were taken: LAB = length of auditory bulla (measured from the suture between
the auditory bulla and the basioccipital to the anterior extremity of the bulla); LDE = length of dentary (measured
from the posterior extremity of the angular process to the anterior margin of the incisor alveolus); HDE = height of
dentary (measured from the ventral margin of the angular process to the posterior extremity of the condyloid
process).
The new species was compared to specimens of Coendou nycthemera (Olfers) and C. insidiosus (Olfers) housed
at MBML, MN, MZUSP and UFES (Appendix 1); measurements of both taxa (including holotypes) were provided by
Voss & Angermann (1997). All specimens were classified into four heuristic age categories following Voss &
Angermann (1997). Only adult specimens were used in the morphological and morphometric analyses. Morphometric
data were analyzed using Statistica 7.0 for Windows (Statsoft Inc. 2004). Two-tailed t-tests were used to evaluate the
significance of mean differences between species. Discriminant function analysis (DFA) was applied to cranial
morphometric data after measurement values were log10 transformed (following Sokal & Rohlf 1995).
Molecular data and phylogenetic analyses. We extracted total genomic DNA from liver tissues preserved in
95% ethanol using the salt protocol and proteinase K (Bruford et al. 1992). Primers MVZ05 and MVZ16 (Smith &
Patton 1993) were used in the polymerase chain reaction (PCR) to amplify the initial 801 base pairs (bp) of the
mitochondrial cytochrome b (cyt-b) gene. PCR products were purified using ExoSAP enzymes (GE Healthcare
Life Sciences). In the cycle-sequencing reactions, we used primers MVZ05 and MVZ127 (Smith & Patton 1993;
Leite & Patton 2002) for the light strand and MVZ04 and MVZ16 (Smith & Patton 1993) for the heavy strand with
BigDye Terminator 3.1 (Applied Biosystems, Inc.). Sequences were read on an ABI-310 capillary automated
sequencer (Applied Biosystems, Inc.) and aligned using Clustal W (Thompson et al. 1994) as implemented in
MEGA version 4.0 (Tamura et al. 2007). Genetic distances were estimated in MEGA, using Kimura's 2-parameter
model (K2P). Additional sequences were obtained from GenBank (http://www.ncbi.nlm.nih.gov/Genbank/),
including those from Erethizon dorsatum (Linnaeus) and Chaetomys subspinosus (Olfers), which were used as
outgroups (Table 1).

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TABLE 1. Specimens used in the phylogenetic analyses of cytochrome b, specimen number, GenBank accession
number, collecting locality and reference data.
Species Specimen number GenBank Locality Reference
accession
C. speratus MN 72045* KC261595 Usina Trapiche, Pernambuco this study
C. speratus MN 72046 KC261594 Usina Trapiche, Pernambuco this study
C. speratus UFPE 1708 KC261592 Usina Trapiche, Pernambuco this study
C. speratus UFPE 1709 KC261593 Usina Trapiche, Pernambuco this study
C. speratus UFES 1184 KC261596 Usina Trapiche, Pernambuco this study
C. nycthemera UFES 2079 KC261597 UHE Estreito, Tocantins this study
C. insidiosus UFES 136 KC261591 Nova Viçosa, Bahia this study
C. melanurus UFPB 3001 AF411583 São João da Baliza, Roraima Bonvicino et al. 2002
C. prehensilis MN 73383** HM 462243 Usina Trapiche, Pernambuco Leite et al. 2011
C. prehensilis Manso 138 AF411582 UHE Manso, Mato Grosso Bonvicino et al. 2002
C. prehensilis Manso 212 AF411581 UHE Manso, Mato Grosso Bonvicino et al. 2002
C. prehensilis Manso 849 AF411584 UHE Manso, Mato Grosso Bonvicino et al. 2002
C. prehensilis MNFS 1016 U34851 Fazenda Santa Fé, Acre Lara et al. 1996
C. prehensilis MNFS 439 U34852 Eirunepé, Amazonas Lara et al. 1996
C. spinosus CIT 1326 EU544661 UHE Rosal, Espírito Santo Vilela et al. 2009
C. spinosus MN 46937 AF411580 Sumidouro, Rio de Janeiro Bonvicino et al. 2002
C. spinosus MN 46938 AF407277 Rio das Ostras, Rio de Janeiro Bonvicino et al. 2002
C. spinosus CIT 2113 EU544662 Biritiba Mirim, São Paulo Vilela et al. 2009
E. dorsatum – FJ357428 – Vilela et al. 2009
Ch. subspinosus – EU544660 Salvador, Bahia Vilela et al. 2009

* holotype; ** neotype.

We examined hierarchical relationships among cyt-b haplotypes using maximum parsimony and Bayesian
inference. Maximum parsimony analysis with heuristic search was performed in PAUP* 4.0 beta 10 (Swofford
2003) with stepwise addition and a limit of 100 rearrangements of TBR (tree-bisection-reconnection) used in the
branch-swapping algorithm. Statistical support was estimated using the bootstrap with 500 pseudo-replicates, and
we considered robust only nodes with bootstrap support above 80%. Bayesian inference was performed in
MrBayes 3.1.2 (Huelsenbeck & Roquist 2001) running for 106 generations and sampling one tree every 103
generations, resulting in 103 trees. We eliminated the first 250 trees as burn-in and estimated a 50% majority-rule
consensus from the remaining trees. Only nodes with Bayesian posterior probability (BPP) greater than 95% were
considered robust. The program jModelTest0.1.1 (Posada 2008) was used to establish the best evolutionary model
according to the data, using the Bayesian Information Criterion (BIC). The Hasegawa et al. (1985) model was
selected, with gamma distribution shape parameter (HKY+G) of 0.2530, transition/transversion ratio of 7.1994,
and base frequencies A = 0.3099, C = 0.2816, G = 0.1060 and T = 0.3025.

Results and discussion

A new species of small porcupine was discovered during regular field surveys within a privately owned area in the
state of Pernambuco, northeastern Brazil. Although this species had not been sighted in 23 other forest fragments
previously surveyed in the same region, it has regularly been sighted at Usina Trapiche, suggesting that the species
is relatively common in that particular area.

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FIGURE 1. The location of the Pernambuco Endemism Centre in northeast Brazil; former and current forest remnants, and the
location of the study site at Usina Trapiche; exact locations of the sightings and collections of Coendou speratus at Usina
Trapiche, Brazil.

Coendou speratus sp. nov.

Figure 2

Holotype. MN 72045, a pregnant adult female collected by A. R. Mendes Pontes (field number ARMP 61) on 25
March 2009. The holotype consists of a flat skin (Figure 3) and skull (Figure 4) accompanied by liver, heart and
muscle samples preserved in 95% ethanol.
Type locality. Mata Tauá, Usina Trapiche, municipality of Sirinhaém, state of Pernambuco, Brazil,
8°33'46.13"S 35°10'9.09"W, elevation 85 m. Mata Tauá is a 280.33 ha isolated fragment of Submontane Tropical
Rainforest (Oliveira & Fontes 2000) within Usina Trapiche, a privately owned sugar plantation.
Paratypes. All four paratypes were collected by Mendes Pontes, Gadelha and Melo in the Usina Trapiche
area, Sirinhaém, state of Pernambuco, Brazil. Like the holotype, these specimens consist of flat skins and skull
with liver, heart and muscle tissue preserved in 95% ethanol. Three paratypes were collected at Boca da Mata
(8°31'56.64"S 35°5'39.85"W): of these, MN 72046 (=ARMP 60) was a pregnant adult female collected on 15
February 2009; UFPE 1708 (=ARMP 56) was an adult male collected on 22 December 2008; and UFPE 1709
(=ARMP 57) was a pregnant adult female collected on 22 January 2009, with quills on the rump and tail damaged
by fire. The fourth paratype, UFES 1184 (=ARMP 62), was an adult male collected at Mata da Barragem
(8°37’49”S 35°11’48”W), on 27 April 2009.
Additional records. Two specimens of Coendou speratus were rescued by the Instituto Chico Mendes de
Conservação da Biodiversidade (ICMBio) on two different occasions. The first was rescued on 20 September 2009
from a house close to a forest fragment in Curado IV (8°3'54.84"S 34°59'55.76"W), city of Jaboatão dos
Guararapes, approximately 50 km from Usina Trapiche; this animal was subsequently released in the Mata Xanguá
(8°39'13.14"S 35°10'11.78"W) on 24 September 2009 by ICMBio employees (release # 38 of Centro de Triagem
de Animais Silvestres, CETAS). The second animal was rescued on 28 September 2009 in Várzea (8°2'39.46"S
34°57'40.10"W), city of Recife, approximately 60 km from Usina Trapiche; this animal was released in an urban
protected forest fragment, Mata Brennand (8°3'13.29"S 34°58'52.52"W), Várzea, Recife, on 30 September 2009,
by ICMBio employees (release # 41 of CETAS).
In addition, a specimen housed at the Museu Nacional, Rio de Janeiro, Brazil, also matches the description of
C. speratus. This specimen apparently lacks a catalog number but it has field number SNP1 (for “Serviço Nacional
de Peste” #1) and consists of the skin and skull of an adult female. It was collected during a plague survey in the

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vicinity of Viçosa, state of Alagoas, northeast Brazil, just 150 km southwest of the type locality of C. speratus. The
common name coandu-mirim is written on the skin tag, and the collecting date is unknown, but it probably took
place between 1952 and 1956 (Oliveira & Franco 2005), and we identify it as belonging to this new species.
Diagnosis. Coendou speratus is a small-bodied, long- tailed species (Table 2) that appears to be completely
spiny because it lacks long fur (Figures 3, 5). The dorsal quills have conspicuously brownish red tips (Figure 6) that
contrast with the blackish dorsal background color. The sigmoid notch of the mandible is shallow; the condylar and
coronoid processes reach the same height, delimiting a shallow and squarish sigmoid notch (Figure 4). The kidney-
shaped auditory bullae are inflated anteriorly, and each one has a small external auditory meatus. The
sphenopalatine vacuities are present in most specimens examined.

TABLE 2. External measurements (mm) and body weight (g) of the type series of Coendou speratus. M=male, F=female.
UFPE 1708 UFPE 1709 MN 72046 MN72045 UFES 1184
(=ARMP 56) (=ARMP 57) (=ARMP 60) (=ARMP 61) (=ARMP 62)
paratype (M) paratype (F)* paratype (F)* holotype (F)* paratype (M)

Head and body length (HBL) 330 370 NA 440 340

Tail length (TAL) 320 310 NA 286 314
Ear length (EAR) 22 16 NA 27 24
Hind foot length (HFL) 56 58 NA 50 51
Weight (W) 1650 1050 NA 1600 1450

* pregnant; NA = not available

Geographic distribution. The new species is probably endemic to the Pernambuco Endemism Centre or
Pernambuco refugium (Punde et al. 2008; Carnaval et al. 2009) in the northeast sector of the Brazilian Atlantic
forest (north of the São Francisco river), which encompasses the coastal region of the states of Alagoas,
Pernambuco, Paraíba and Rio Grande do Norte. However, we did not find the new species during 5-year surveys in
23 other areas, including the largest and best-preserved forest fragment, the Mata Coimbra (3,642 ha), not far from
the type locality at Usina Trapiche. Further surveys along the Atlantic coast of northeastern Brazil, are needed in
order to locate additional populations and determine the geographic range of this species.
The geographic range of Coendou speratus overlaps with that of C. prehensilis, but not with that of C.
insidiosus (which only occurs to the south of the São Francisco River mouth; Oliver & Santos 1991; Caldara Jr. &
Leite 2012). Apparently, the range of C. speratus does not overlap with that of C. nycthemera, an Amazonian
species that ranges from the lower Amazon and lower Tocantins in the Brazilian state of Amazonas to the
northwest part of the state of Maranhão (Handley & Pine 1992; Voss & Angermann 1997; Oliveira et al. 2007).
Habitat. We sighted the new species in systematic census only at Usina Trapiche, where submontane remnants
of Dense Tropical Rainforests are preserved on hilltops that are not suitable for agriculture. These remnants are
highly threatened by selective cutting, intentional fires, presence of domestic animals, and most importantly,
hunting. Patrolling of the remnants is highly limited and we regularly encountered hunters, heard shots and found
hunting devices and suspended hunting platforms in the trails and within the forest. We believe the habitat of the
porcupine may include other forest types along the Atlantic coast of northeastern Brazil, such as Open Canopy
Tropical Rainforests and Deciduous Seasonal Forests, but most of these forest types have been destroyed in this
region.
Description. Head. Tricolored quills whitish yellow basally, black in the middle, brownish red distally (Figure
6G); bristles yellowish white basally, black in the middle, light brownish red distally; long sparse black mystacial
vibrissae, some extending beyond shoulder line (ca. 90 mm); pinkish bulbous muzzle covered with black to
brownish short hairs (Figure 2); rounded flat ears with yellow-tipped black hairs on the inner side; shorter
submental vibrissae present.
Body. Most dorsal quills tricolor (bright yellow basally, black in the middle, and brownish red distally) from
head to mid-back (Figure 6A–B); dorsal quills have a long black band and short brownish red tip in some
specimens (e.g., UFPE 1708), but a short black band and long brownish red tip in others (e.g., MN 72045). Less
conspicuous, black tipped, light yellow based bicolor quills also present in all specimens (Figure 6D). Some

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specimens (e.g., MN 72045) also have yellow based, brownish red tipped bicolor quills from head to mid-back
(Figure 6C). Quills on rump bicolor, shorter than those from head to mid-back, with larger yellow basal band, and
black tip. Bicolor quills on rump and thighs varying from dark yellow (e.g., MN 72045, Figure 6E) to light yellow
(e.g., MN 72046, Figure 6F). Specimens with bicolor, red-brownish quills also have tricolor quills with short black
band on mid-dorsum and dark yellow-based bicolor quills on the rump (e.g., MN 72045), giving an overall
brownish red dorsal coloration (Figure 3). In general, dorsal quills densely grouped in very sharp duals or triads,
some with barbed tips. Thin, sparse, short grayish dorsal hairs are hidden by the quills. Ventral surface sparsely
covered with soft, 14 mm-long, grayish to totally whitish hairs; spinous hairs at the transition to the dorsal surface,
whitish to yellowish basally, black in the middle, and from reddish to beige distally.
Limbs. Inner fore- and hind-limbs covered with brownish to grayish dense hairs; outer fore- and hind-limbs
with predominately spinous hairs, whitish to yellowish basally, blackish to brownish in the middle, and beige to
yellowish distally, measuring ca. 10 mm on forelimbs and 14 mm on hindlimbs. Anconeal, medial antebrachial,
and ulnar carpal vibrissae present, measuring ca. 45 mm; long, curved claws.
Tail. Prehensile; dorsal surface of the proximal half covered with ca. 26 mm-long bicolored quills, similar to
those on the rump and thighs, gradually shorter towards the distal half; quills light yellow and bristles brownish
(MN 72046, UFPE 1708, UFES 1184) or quills brownish yellow and bristles orange to reddish brown (MN 72045,
UFPE 1709); yellow tipped black spinous hairs at the transition between the dorsal and ventral surfaces and lateral
grayish soft hairs at the base of the tail; distal half of the tail covered with ca. 13 mm-long dark brown (MN 72046,
UFPE 1708, UFES 1184) to reddish brown bristles (MN 72045, UFPE 1709), including the ventral surface (Figure
5), except for the naked prehensile tip.
Skull. Dorsal cranial profile flat over the nasals and frontals, contrasting with slightly convex surface of the
parietals in lateral view; post-orbital ridges reach the protruded lambdoidal ridge; nasal bones very long, tapering
posteriorly and anteriorly to pointed ends; zygomatic arches widest posteriorly, converging toward the rostrum
with a slight secondary widening at the orbits; jugal only slightly expanded; right and left incisive foramina
separated by a complete median septum and bordered posteriorly by maxillary bones; posterior diastema and
palatal bridge between cheek teeth narrowly constricted with a median keel; anterior margin of mesopterygoid
fossa ?-shaped, extending to the level of the second molars; roof of mesopterygoid fossa varying from completely
ossified (e.g., UFPE 1709) to perforated by small to large sphenopalatine vacuities (e.g., MN 72045, Figure 4);
auditory bullae large, bean-shaped, and constricted posteriorly to contact paroccipital processes; roof of the
external auditory meatus varies from weakly keeled in some specimens (e.g., UFPE 1709) to smooth in others (e.g.,
MN 72046).
Dentition. Anterior surfaces of incisors orangish and remaining surfaces whitish; maxillary toothrows
subparallel; maxillary teeth pentalophodont, resembling those of other erethizontids (except Chaetomys) in
occlusal morphology; Permanent fourth upper premolar larger than first molar, and second molar smaller than first
molar and larger than third molar.
Etymology. The specific epithet speratus, or ‘hope’, acknowledges our hope that this new species calls the
attention of the world to the critical need to save the highly-threatened region where it occurs. To date, 50% of all
trees (Silva & Tabarelli 2000; Oliveira et al. 2004; Santos et al. 2008), 50% of all medium-sized mammals, and
100% of all large mammals have gone extinct in the Pernambuco Endemism Centre (Mendes Pontes et al. 2005;
Silva Jr. & Mendes Pontes 2008; Mendes Pontes 2009), where even small mammal populations are far below levels
that are sustainable in the long run (Asfora & Mendes Pontes 2009). We also hope to call the attention of funding
agencies and decision-makers to the need for intensive inventories in this hotspot’s hotspot in order to establish the
very first step towards conservation, which is to know the true biological value of the forest remnants, and most
importantly, the need for financial investment in this sector of the Atlantic forest of Brazil.
Phylogenetic relationships and genetic divergences. The phylogenetic position of Coendou speratus is
uncertain, although cyt-b sequences indicate that it is closely related to C. nycthemera, C. spinosus, and C.
insidiosus (Figure 7). The Bayesian and parsimony trees based on the cyt-b gene were almost identical, with only
one difference: the former shows C. nycthemera sister to C. speratus, but with no support (Figure 7), while the
latter shows C. nycthemera sister to C. spinosus, also with no nodal support. Specimens identified as “C. spinosus”
did not form a monophyletic group in the cyt-b tree (Figure 7), but this is not the focus of the present paper and will
be discussed elsewhere.

426 · Zootaxa 3636 (3) © 2013 Magnolia Press PONTES ET AL.

FIGURE 2. Live specimen of Coendou speratus in captivity (MN 72046, paratype).

The average pairwise genetic distance of 7.7% between C. speratus and C. nycthemera and 7.2% between
them and C. spinosus + C. insidiosus (Figure 7) are consistent with other cyt-b divergences among species of
Coendou, although smaller when compared to C. melanurus and C. prehensilis (10.2% and 11.9%, respectively).
Comparisons. Coendou speratus is overall similar to C. nycthemera, but there are clear morphological
differences between them: the dorsal body quills are typically bicolored (white base, black tip) in C. nycthemera
and tricolored (yellow base, black middle, brownish red tip) in C. speratus, although most C. nycthemera show at
least some tricolored (white- or pale-brown-tipped) dorsal quills. Moreover, the mass effect is black speckled with
white or pale brown in C. nycthemera, but brownish red in C. speratus. The tail is mostly covered with short black
hairs in C. nycthemera, but with dark brown to reddish bristles in C. speratus. The bulbous muzzle is relatively
larger in C. speratus than in C. nycthemera, where it is only slightly swollen. Coendou speratus has long mystacial
vibrissae, some extending beyond the shoulder line, but in C. nycthemera these hairs extend just beyond the ears.
When compared to C. nycthemera, the new porcupine species is heavier, has longer body, longer nasals, longer
zygomatic arch, longer infraorbital foramen, longer diastema, longer dentary, wider braincase and posterior palate
(Table 4).
Coendou speratus is externally very distinct from C. insidiosus, especially because the latter has bicolored
dorsal quills (light yellow based and black tipped) that are almost completely hidden beneath longer and
homogeneous pale or dark hairs. Some specimens of C. insidiosus have tricolored dorsal quills, present only on the
head and shoulders (Caldara Jr. & Leite 2012). C. speratus has a longer body, tail, skull, and auditory bullae, and
higher infraorbital foramen than C. insidiosus. In addition, C. speratus has a narrower anterior palate and smaller
incisive foramina than C. insidiosus (Table 4). The discriminant function analysis also supported species
distinction based on cranial measurements, confirming that C. speratus is morphometrically different from both C.
insidiosus and C. nycthemera (Figure 8).

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FIGURE 3. Variation in the dorsal pelage of Coendou speratus: A) holotype (MN 72045); B, C) two paratypes (MN 72046,
UFES 1184).

428 · Zootaxa 3636 (3) © 2013 Magnolia Press PONTES ET AL.

FIGURE 4. Skull of the holotype of Coendou speratus sp. nov. (MN 72045) in dorsal, lateral and ventral views (A, B, D), and
dentary in lateral and dorsal views (C, E).

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FIGURE 5. Dorsal (A) and ventral (B) surfaces of body and ventral surface of the prehensile tail (C) of Coendou speratus (MN
72046, paratype).

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FIGURE 6. Variation in Coendou speratus quills. A, B) Mid-dorsal tricolor quills (UFPE 1708, MN 72045); C, D) Mid-dorsal
bicolor quills (MN 72045, MN 72046); E, F) Bicolor quills from the rump (MN 72045, MN 72046); G) tricolor quills from the
head (MN 72045).

TABLE 3. Density (ind/km²) and biomass (kg/km²) of Coendou speratus and C. prehensilis in each forest fragment at Usina
Trapiche, Pernambuco, Brazil.
Forest fragment (size)
Species Boca da Mata Xanguazinho Mata Tauá Mata Xanguá
(94 ha) (100 ha) (280 ha) (470 ha)
density biomass density biomass density biomass density biomass
Coendou speratus 4.98 6.73 6.2 8.44 25 33.75 28.1 37.93
Coendou prehensilis 6.83 31.57 4.62 21.35 – – 4.42 20.42

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FIGURE 7. Bayesian inference tree based on cytochrome b sequences from Coendou species. Numbers above branches
indicate Bayesian posterior probabilities (BPP) ≥ 0.95 and parsimony bootstrap support values ≥ 80%. Numbers below
branches indicate average pairwise genetic distances (Kimura 2-parameter) among specimens. Erethizon dorsatum and
Chaetomys subspinosus were used as outgroups.

Thomas (1902) described Coendou roberti from southern Brazil as a spiny, short-haired species allied to C.
spinosus, but showing tricolored, orange-tipped spines, not hidden by a long clothing of fur, which therefore
resembles C. speratus. However, there are several differences among them, especially the bicolored spines on the
rump partly hidden by hair in C. roberti, but not in C. speratus. In addition, C. roberti is currently considered a
phenotypic variant of C. spinosus because they share several characters and there is clear intergradation among
distinct phenotypes (Caldara Jr. & Leite 2012). Voss (2011) appropriately designated a neotype for C. spinosus and
considered C. roberti as a junior synonym.
Natural history and ecology. Coendou speratus is sympatric with its larger congener C. prehensilis, but the
former appears to prefer the lower forest strata. Coendou speratus dens in hollow trees and has a strong and
characteristic pungent odor, which is much stronger than the odor of sympatric C. prehensilis. Seven C. speratus
individuals were sighted in four forest fragments. The first sighting occurred at Boca da Mata, when an individual
was spotted at 11:29 pm, resting on a 12 meter-high horizontal branch; the second was at Mata Xanguazinho, also

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of a single individual resting at 10:27 pm, on a 10 meter-high branch; the third was at Mata Tauá, when two
individuals were sighted at 00:20 am, resting together on a 20 meter-high branch; the fourth sighting was also at
Mata Tauá, at 11:55 pm, at a 10 meter-high branch; the fifth and the six were at Mata Xanguá, at 7:50 pm and 00:38
am, respectively, both on a 20 meter-high branch.

FIGURE 8. Bivariate plot of scores for two discriminant functions (DF1 and DF2) based on skull measurements of Coendou
insidiosus (Cins), C. nycthemera (Cnyc), and C. speratus (Cspe).

Coendou speratus seems to be solitary, although two individuals were sighted together once, probably a male
and a female, one of which vocalized at the observer’s presence. At all times when C. speratus was sighted, it
became either motionless or moved very slowly and then became motionless, whereas Coendou prehensilis
escaped quickly at all times. Coendou speratus was once sighted 3 meters from a collared anteater, Tamandua
tetradactyla (Linnaeus). According to locals, the new species feeds on the fruits of the exotic African oil palm
Elaeis guineensis Jacq. (locally known as dendezeiro).
Previous long-term line transect surveys carried out between 2000 and 2008 in the Pernambuco Endemism
Center (Mendes Pontes et al. 2005; Melo 2009; Mendes Pontes 2009; Gadelha 2009) detected the collapse of ~50%
of the entire regional mammalian fauna and did not register the new species. Out of 38 medium- and large-sized
mammal species formerly occurring in the study area, only 53.8% (n=21) were sighted or camera-trapped.
According to them, no fragment hosted the entire remaining mammal community, and only four species (19%)
occurred in very small fragments (≤10 ha, which sums ~70% of what remains); the mammalian community was
highly simplified, with all large mammals being regionally extinct. There was no nestedness regarding area of the
fragment or degree of isolation, which implied that the occurrence of a mammal species in a given forest patch
varied unpredictably, and that in this ongoing process of mass extinction the 21st century medium-sized remaining
mammalian fauna will soon be extinct if strict conservation measures are not implemented (Gadelha 2009; Melo
2009).

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 The new subsequent line transect surveys that resulted in the discovery of this new species were carried out in
the highly impacted 4,000 ha forest fragments at Usina Trapiche, between 2008 and 2011 (Gadelha 2009; Melo
2009; Leite et al. 2011; Freitas 2012), with a sample effort of 302.5 km walked (diurnal and nocturnal) in nine
forest fragments, totaling 68 surveys in 610 h. Reinforcing our previous findings that in this region the species are
expected to occur in the forest fragments in an unpredictable fashion, density and biomass derived from 143 km of
nocturnal surveys, revealed that Coendou speratus had comparatively high local densities in the four fragments
where it was encountered. In the largest fragment, Mata Xanguá (470 ha), C. speratus had the highest density (36.8
ind/km²), while in the smallest fragment, Boca da Mata (94 ha), it had the lowest (5 ind/km²). Coendou speratus
was found syntopically with C. prehensilis in all but one forest fragment, Mata Tauá (280 ha), and the biomass
contribution of C. speratus was always lower than that of C. prehensilis (Table 3).
Conservation implications. The discovery of this new species of small porcupine in this particular zone of
endemism highlights the importance of this hotspot for the conservation of the earth’s biota. It also points to the
lack of information on its mammalian fauna, and the need for surveys in order to understand their distribution and
status throughout the region (Costa et al. 2005).
Human impact is extremely high in the region where we found this new porcupine (Melo 2009), resulting in
the regional extinction of large mammals, such as jaguar (Panthera onca), Brazilian tapir (Tapirus terrestris), giant
anteater (Myrmecophaga tridactyla), and white-lipped peccary (Tayassu pecari) (Mendes Pontes 2009). The threat
level to other medium-size mammals, like the blond capuchin monkey Sapajus flavius (Schreber) (Mendes Pontes
et al. 2006; Alfaro et al. 2012), is currently still very high. Thus, long-term research programs and conservation
measures must be implemented to assure the survival of the remaining species and protection of these forests.

Acknowledgements

We are grateful to Salgado and Trapiche mills, within which the forest fragments stretch, for their cooperation, for
having granted some seed money essential for the accomplishment of this study, and also for having provided the
necessary infrastructure for the project. We are especially indebted to Mrs. Evânia Freire e Dr. Mário Jorge Seixas
Aguiar. Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) granted an undergraduate
scholarship to JRG and a productivity fellowship to YLRL. In 2008 we received the recognition and financial
support from Conservation International in Brazil, which has sponsored this study, both field inventories and
molecular analysis through the Critical Ecosystems Partnership Fund (CEPF). Fundação de Amparo à Pesquisa do
Espírito Santo (FAPES) provided additional grant support to this project. ACL has a graduate scholarship from
Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES). The curators João Alves de Oliveira
(MN) and Diego Astúa (UFPE) kindly received museum specimens and promptly provided catalog numbers. Celso
Azevedo (UFES) provided helpful comments on taxonomic and nomenclatural issues, including the new scientific
name proposed here. Carlos Eduardo D. Cintra (Systema Naturae Consultoria Ambiental Ltda.) kindly provided a
valuable specimen of C. nycthemera for comparison and molecular analysis. Marcelo Weksler and two anonymous
reviewers provided valuable comments that improved the quality of this manuscript. Instituto Chico Mendes de
Conservação da Biodiversidade (ICMBio) granted collecting permits to the authors (954-1, 954-2, 1095-1, 1251-
1). Catarina Cabral, Fernando Rodrigues, Joana Maranhão, and Yuri Valença, from the Centro de Triagem de
Animais Silvestres (ICMBio, Recife), kindly granted us the privilege to join the reintroduction to the wild of a
specimen of Coendou speratus sp. nov.

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APPENDIX 1. Specimens of Coendou insidiosus and C. nycthemera examined.

C. insidiosus:
BAHIA: Nova Viçosa, Fazenda Elma (UFES 136); Caravelas, Fazenda Monte Castelo (UFES 137); Ilhéus, Banco da
Vitória, Fazenda Pirataquissé (MN 11211); Porto Seguro (MN 55527). ESPÍRITO SANTO: São Mateus, Guriri (MBML
2347); Linhares, Rio São José (MN 8277);
C. nycthemera:
PARÁ: Belém, Sítio Velho (MZUSP 13519). Cametá (MZUSP 5031, 5034, 5035 5036, 5037, 5038). Curralinho (MN
4913, 4914, 4915, 4916, 4919, 4920, 4921, 4638, 4639, 4678, 4681, 4683, 4695). Santarém, Mata Virgem, Terra Firme (MN
6530, 7652, 7653, 7654). TOCANTINS: Darcinópolis, Usina Hidrelétrica Estreito (UFES 2079).

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