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  • 1. Zootaxa 3636 (3): 421–438 ISSN 1175-5326 (print edition) www.mapress.com / zootaxa /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:FE435AD6-66A4-46D8-95D3-7BFFD2ED28F1A new species of porcupine, genus Coendou (Rodentia: Erethizontidae)from the Atlantic forest of northeastern BrazilANTONIO 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. LEITE31 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.com2 Universidade Federal Rural de Pernambuco, Departamento de Morfologia e Fisiologia Animal, Laboratório de Anatomia. Rua DomManoel de Medeiros, S/N, Dois Irmãos, CEP: 52.171-900, Recife, PE, Brazil3 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, BrazilAbstractWe 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 Atlanticcoast of northeastern Brazil north of the São Francisco river, one of the most important known biodiversity hotspots. Thegeographic 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 speratusis a small-bodied, long-tailed species that appears to be completely spiny because it lacks long dorsal fur. The dorsal quillshave conspicuously brownish red tips that contrast with the blackish dorsal background color. The new species is overallsimilar to C. nycthemera, but the dorsal body quills are typically tricolored in the former and bicolored in the latter. Thenew species is externally very distinct from C. insidiosus, especially because the latter has bicolored dorsal quills that arealmost completely hidden beneath longer and homogeneous pale or dark hairs.Key words: Biodiversity hotspot, Coendou, Mammalia, Neotropics, taxonomyIntroductionThe Brazilian Atlantic forest is considered a world priority for biodiversity conservation and the fourth mostimportant hotspot of the planet (Myers et al. 2000). The Pernambuco Endemism Centre, a biogeographical unit ofthe Brazilian Atlantic forest that lies north of the São Francisco river, harbors several endemic species, whichsuggests that it is a hotspot within a hotspot (Punde et al. 2008; Carnaval et al. 2009). Unfortunately, this region hasso 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 onimmature seeds, green or ripe fruit, palm nuts, bark and leaves (Emmons & Feer 1997; Lima et al. 2010; Passamani2010). 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 spinyCoendou (e.g., Husson 1978; Eisenberg & Redford 1999; Nowak 1999; Bonvicino et al. 2002; Woods & Kilpatrick2005), 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 meaningfullydiagnosed as separate taxa. Here we follow this latter view. The number of species of Neotropical erethizontids isstill 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 describedAccepted by M. Weksler: 18 Feb. 2013; published: 5 Apr. 2013 421
  • 2. below had not been detected before in any of several sites recently surveyed in the region, and we found noreference to this small porcupine in historical accounts, such as Marcgrave’s (1648), who carefully described thisregional fauna, including the larger porcupine, Coendou prehensilis Linnaeus (Leite et al. 2011).Material and methodsStudy area and specimens. The biodiversity inventories that resulted in the discovery here reported have beencarried out since January 2002, through the line transect method (Buckland et al. 1993), adapted from MendesPontes (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 Atlanticforest 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 bypeople and domestic animals. The description of the new porcupine, locally known as coandu-mirim, is based on the analysis of fiveindividuals deposited at Museu Nacional, Universidade Federal do Rio de Janeiro, Brazil (MN), the mammalcollection of the Universidade Federal de Pernambuco, Recife, Brazil (UFPE), and the mammal collection of theUniversidade 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. MelloLeitão, 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 zygomaticbreadth; 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 incisiveforamen; BIF = breadth of incisive foramen; BIT = breadth of incisor tips; DI = depth of incisor; MTR = maxillarytoothrow length; LM = length of molars; BP4 = breadth of fourth pre-molar; BM1 = breadth of first molar. Threeadditional cranial measurements were taken: LAB = length of auditory bulla (measured from the suture betweenthe auditory bulla and the basioccipital to the anterior extremity of the bulla); LDE = length of dentary (measuredfrom the posterior extremity of the angular process to the anterior margin of the incisor alveolus); HDE = height ofdentary (measured from the ventral margin of the angular process to the posterior extremity of the condyloidprocess). The new species was compared to specimens of Coendou nycthemera (Olfers) and C. insidiosus (Olfers) housedat MBML, MN, MZUSP and UFES (Appendix 1); measurements of both taxa (including holotypes) were provided byVoss & 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. Morphometricdata were analyzed using Statistica 7.0 for Windows (Statsoft Inc. 2004). Two-tailed t-tests were used to evaluate thesignificance of mean differences between species. Discriminant function analysis (DFA) was applied to cranialmorphometric 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 in95% 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 themitochondrial cytochrome b (cyt-b) gene. PCR products were purified using ExoSAP enzymes (GE HealthcareLife 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 withBigDye Terminator 3.1 (Applied Biosystems, Inc.). Sequences were read on an ABI-310 capillary automatedsequencer (Applied Biosystems, Inc.) and aligned using Clustal W (Thompson et al. 1994) as implemented inMEGA version 4.0 (Tamura et al. 2007). Genetic distances were estimated in MEGA, using Kimuras 2-parametermodel (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 asoutgroups (Table 1).422 · Zootaxa 3636 (3) © 2013 Magnolia Press PONTES ET AL.
  • 3. TABLE 1. Specimens used in the phylogenetic analyses of cytochrome b, specimen number, GenBank accessionnumber, collecting locality and reference data.Species Specimen number GenBank Locality Reference accessionC. speratus MN 72045* KC261595 Usina Trapiche, Pernambuco this studyC. speratus MN 72046 KC261594 Usina Trapiche, Pernambuco this studyC. speratus UFPE 1708 KC261592 Usina Trapiche, Pernambuco this studyC. speratus UFPE 1709 KC261593 Usina Trapiche, Pernambuco this studyC. speratus UFES 1184 KC261596 Usina Trapiche, Pernambuco this studyC. nycthemera UFES 2079 KC261597 UHE Estreito, Tocantins this studyC. insidiosus UFES 136 KC261591 Nova Viçosa, Bahia this studyC. melanurus UFPB 3001 AF411583 São João da Baliza, Roraima Bonvicino et al. 2002C. prehensilis MN 73383** HM 462243 Usina Trapiche, Pernambuco Leite et al. 2011C. prehensilis Manso 138 AF411582 UHE Manso, Mato Grosso Bonvicino et al. 2002C. prehensilis Manso 212 AF411581 UHE Manso, Mato Grosso Bonvicino et al. 2002C. prehensilis Manso 849 AF411584 UHE Manso, Mato Grosso Bonvicino et al. 2002C. prehensilis MNFS 1016 U34851 Fazenda Santa Fé, Acre Lara et al. 1996C. prehensilis MNFS 439 U34852 Eirunepé, Amazonas Lara et al. 1996C. spinosus CIT 1326 EU544661 UHE Rosal, Espírito Santo Vilela et al. 2009C. spinosus MN 46937 AF411580 Sumidouro, Rio de Janeiro Bonvicino et al. 2002C. spinosus MN 46938 AF407277 Rio das Ostras, Rio de Janeiro Bonvicino et al. 2002C. spinosus CIT 2113 EU544662 Biritiba Mirim, São Paulo Vilela et al. 2009E. dorsatum – FJ357428 – Vilela et al. 2009Ch. subspinosus – EU544660 Salvador, Bahia Vilela et al. 2009* holotype; ** neotype. We examined hierarchical relationships among cyt-b haplotypes using maximum parsimony and Bayesianinference. Maximum parsimony analysis with heuristic search was performed in PAUP* 4.0 beta 10 (Swofford2003) with stepwise addition and a limit of 100 rearrangements of TBR (tree-bisection-reconnection) used in thebranch-swapping algorithm. Statistical support was estimated using the bootstrap with 500 pseudo-replicates, andwe considered robust only nodes with bootstrap support above 80%. Bayesian inference was performed inMrBayes 3.1.2 (Huelsenbeck & Roquist 2001) running for 106 generations and sampling one tree every 103generations, resulting in 103 trees. We eliminated the first 250 trees as burn-in and estimated a 50% majority-ruleconsensus from the remaining trees. Only nodes with Bayesian posterior probability (BPP) greater than 95% wereconsidered robust. The program jModelTest0.1.1 (Posada 2008) was used to establish the best evolutionary modelaccording to the data, using the Bayesian Information Criterion (BIC). The Hasegawa et al. (1985) model wasselected, 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 discussionA new species of small porcupine was discovered during regular field surveys within a privately owned area in thestate of Pernambuco, northeastern Brazil. Although this species had not been sighted in 23 other forest fragmentspreviously surveyed in the same region, it has regularly been sighted at Usina Trapiche, suggesting that the speciesis relatively common in that particular area.A NEW SPECIES OF PORCUPINE FROM BRAZIL Zootaxa 3636 (3) © 2013 Magnolia Press · 423
  • 4. FIGURE 1. The location of the Pernambuco Endemism Centre in northeast Brazil; former and current forest remnants, and thelocation of the study site at Usina Trapiche; exact locations of the sightings and collections of Coendou speratus at UsinaTrapiche, Brazil.Coendou speratus sp. nov.Figure 2Holotype. MN 72045, a pregnant adult female collected by A. R. Mendes Pontes (field number ARMP 61) on 25March 2009. The holotype consists of a flat skin (Figure 3) and skull (Figure 4) accompanied by liver, heart andmuscle samples preserved in 95% ethanol. Type locality. Mata Tauá, Usina Trapiche, municipality of Sirinhaém, state of Pernambuco, Brazil,8°3346.13"S 35°109.09"W, elevation 85 m. Mata Tauá is a 280.33 ha isolated fragment of Submontane TropicalRainforest (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 Trapichearea, Sirinhaém, state of Pernambuco, Brazil. Like the holotype, these specimens consist of flat skins and skullwith liver, heart and muscle tissue preserved in 95% ethanol. Three paratypes were collected at Boca da Mata(8°3156.64"S 35°539.85"W): of these, MN 72046 (=ARMP 60) was a pregnant adult female collected on 15February 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 damagedby 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 deConservação da Biodiversidade (ICMBio) on two different occasions. The first was rescued on 20 September 2009from a house close to a forest fragment in Curado IV (8°354.84"S 34°5955.76"W), city of Jaboatão dosGuararapes, approximately 50 km from Usina Trapiche; this animal was subsequently released in the Mata Xanguá(8°3913.14"S 35°1011.78"W) on 24 September 2009 by ICMBio employees (release # 38 of Centro de Triagemde Animais Silvestres, CETAS). The second animal was rescued on 28 September 2009 in Várzea (8°239.46"S34°5740.10"W), city of Recife, approximately 60 km from Usina Trapiche; this animal was released in an urbanprotected forest fragment, Mata Brennand (8°313.29"S 34°5852.52"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 ofC. speratus. This specimen apparently lacks a catalog number but it has field number SNP1 (for “Serviço Nacionalde Peste” #1) and consists of the skin and skull of an adult female. It was collected during a plague survey in the424 · Zootaxa 3636 (3) © 2013 Magnolia Press PONTES ET AL.
  • 5. vicinity of Viçosa, state of Alagoas, northeast Brazil, just 150 km southwest of the type locality of C. speratus. Thecommon name coandu-mirim is written on the skin tag, and the collecting date is unknown, but it probably tookplace 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 completelyspiny because it lacks long fur (Figures 3, 5). The dorsal quills have conspicuously brownish red tips (Figure 6) thatcontrast with the blackish dorsal background color. The sigmoid notch of the mandible is shallow; the condylar andcoronoid 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. Thesphenopalatine 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 340Tail length (TAL) 320 310 NA 286 314Ear length (EAR) 22 16 NA 27 24Hind foot length (HFL) 56 58 NA 50 51Weight (W) 1650 1050 NA 1600 1450* pregnant; NA = not available Geographic distribution. The new species is probably endemic to the Pernambuco Endemism Centre orPernambuco refugium (Punde et al. 2008; Carnaval et al. 2009) in the northeast sector of the Brazilian Atlanticforest (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 in23 other areas, including the largest and best-preserved forest fragment, the Mata Coimbra (3,642 ha), not far fromthe type locality at Usina Trapiche. Further surveys along the Atlantic coast of northeastern Brazil, are needed inorder 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 Amazonianspecies that ranges from the lower Amazon and lower Tocantins in the Brazilian state of Amazonas to thenorthwest 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 remnantsof Dense Tropical Rainforests are preserved on hilltops that are not suitable for agriculture. These remnants arehighly 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 foundhunting devices and suspended hunting platforms in the trails and within the forest. We believe the habitat of theporcupine may include other forest types along the Atlantic coast of northeastern Brazil, such as Open CanopyTropical Rainforests and Deciduous Seasonal Forests, but most of these forest types have been destroyed in thisregion. Description. Head. Tricolored quills whitish yellow basally, black in the middle, brownish red distally (Figure6G); bristles yellowish white basally, black in the middle, light brownish red distally; long sparse black mystacialvibrissae, some extending beyond shoulder line (ca. 90 mm); pinkish bulbous muzzle covered with black tobrownish short hairs (Figure 2); rounded flat ears with yellow-tipped black hairs on the inner side; shortersubmental vibrissae present. Body. Most dorsal quills tricolor (bright yellow basally, black in the middle, and brownish red distally) fromhead to mid-back (Figure 6A–B); dorsal quills have a long black band and short brownish red tip in somespecimens (e.g., UFPE 1708), but a short black band and long brownish red tip in others (e.g., MN 72045). Lessconspicuous, black tipped, light yellow based bicolor quills also present in all specimens (Figure 6D). SomeA NEW SPECIES OF PORCUPINE FROM BRAZIL Zootaxa 3636 (3) © 2013 Magnolia Press · 425
  • 6. 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, andblack 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 blackband on mid-dorsum and dark yellow-based bicolor quills on the rump (e.g., MN 72045), giving an overallbrownish 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 sparselycovered 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-limbswith predominately spinous hairs, whitish to yellowish basally, blackish to brownish in the middle, and beige toyellowish 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 tothose 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 lateralgrayish 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 (Figure5), except for the naked prehensile tip. Skull. Dorsal cranial profile flat over the nasals and frontals, contrasting with slightly convex surface of theparietals in lateral view; post-orbital ridges reach the protruded lambdoidal ridge; nasal bones very long, taperingposteriorly and anteriorly to pointed ends; zygomatic arches widest posteriorly, converging toward the rostrumwith a slight secondary widening at the orbits; jugal only slightly expanded; right and left incisive foraminaseparated by a complete median septum and bordered posteriorly by maxillary bones; posterior diastema andpalatal bridge between cheek teeth narrowly constricted with a median keel; anterior margin of mesopterygoidfossa -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 theexternal 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 toothrowssubparallel; maxillary teeth pentalophodont, resembling those of other erethizontids (except Chaetomys) inocclusal morphology; Permanent fourth upper premolar larger than first molar, and second molar smaller than firstmolar and larger than third molar. Etymology. The specific epithet speratus, or ‘hope’, acknowledges our hope that this new species calls theattention of the world to the critical need to save the highly-threatened region where it occurs. To date, 50% of alltrees (Silva & Tabarelli 2000; Oliveira et al. 2004; Santos et al. 2008), 50% of all medium-sized mammals, and100% 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 levelsthat are sustainable in the long run (Asfora & Mendes Pontes 2009). We also hope to call the attention of fundingagencies and decision-makers to the need for intensive inventories in this hotspot’s hotspot in order to establish thevery first step towards conservation, which is to know the true biological value of the forest remnants, and mostimportantly, 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 isuncertain, 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 onlyone difference: the former shows C. nycthemera sister to C. speratus, but with no support (Figure 7), while thelatter 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 willbe discussed elsewhere.426 · Zootaxa 3636 (3) © 2013 Magnolia Press PONTES ET AL.
  • 7. 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% betweenthem and C. spinosus + C. insidiosus (Figure 7) are consistent with other cyt-b divergences among species ofCoendou, 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 morphologicaldifferences between them: the dorsal body quills are typically bicolored (white base, black tip) in C. nycthemeraand tricolored (yellow base, black middle, brownish red tip) in C. speratus, although most C. nycthemera show atleast some tricolored (white- or pale-brown-tipped) dorsal quills. Moreover, the mass effect is black speckled withwhite or pale brown in C. nycthemera, but brownish red in C. speratus. The tail is mostly covered with short blackhairs in C. nycthemera, but with dark brown to reddish bristles in C. speratus. The bulbous muzzle is relativelylarger in C. speratus than in C. nycthemera, where it is only slightly swollen. Coendou speratus has long mystacialvibrissae, 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, longerzygomatic 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 bicoloreddorsal quills (light yellow based and black tipped) that are almost completely hidden beneath longer andhomogeneous pale or dark hairs. Some specimens of C. insidiosus have tricolored dorsal quills, present only on thehead and shoulders (Caldara Jr. & Leite 2012). C. speratus has a longer body, tail, skull, and auditory bullae, andhigher infraorbital foramen than C. insidiosus. In addition, C. speratus has a narrower anterior palate and smallerincisive foramina than C. insidiosus (Table 4). The discriminant function analysis also supported speciesdistinction based on cranial measurements, confirming that C. speratus is morphometrically different from both C.insidiosus and C. nycthemera (Figure 8).A NEW SPECIES OF PORCUPINE FROM BRAZIL Zootaxa 3636 (3) © 2013 Magnolia Press · 427
  • 8. 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.
  • 9. FIGURE 4. Skull of the holotype of Coendou speratus sp. nov. (MN 72045) in dorsal, lateral and ventral views (A, B, D), anddentary in lateral and dorsal views (C, E).A NEW SPECIES OF PORCUPINE FROM BRAZIL Zootaxa 3636 (3) © 2013 Magnolia Press · 429
  • 10. FIGURE 5. Dorsal (A) and ventral (B) surfaces of body and ventral surface of the prehensile tail (C) of Coendou speratus (MN72046, paratype).430 · Zootaxa 3636 (3) © 2013 Magnolia Press PONTES ET AL.
  • 11. FIGURE 6. Variation in Coendou speratus quills. A, B) Mid-dorsal tricolor quills (UFPE 1708, MN 72045); C, D) Mid-dorsalbicolor quills (MN 72045, MN 72046); E, F) Bicolor quills from the rump (MN 72045, MN 72046); G) tricolor quills from thehead (MN 72045).TABLE 3. Density (ind/km²) and biomass (kg/km²) of Coendou speratus and C. prehensilis in each forest fragment at UsinaTrapiche, 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 biomassCoendou speratus 4.98 6.73 6.2 8.44 25 33.75 28.1 37.93Coendou prehensilis 6.83 31.57 4.62 21.35 – – 4.42 20.42A NEW SPECIES OF PORCUPINE FROM BRAZIL Zootaxa 3636 (3) © 2013 Magnolia Press · 431
  • 12. 432 · Zootaxa 3636 (3) © 2013 Magnolia Press PONTES ET AL.
  • 13. FIGURE 7. Bayesian inference tree based on cytochrome b sequences from Coendou species. Numbers above branchesindicate 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 andChaetomys 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 thereforeresembles C. speratus. However, there are several differences among them, especially the bicolored spines on therump partly hidden by hair in C. roberti, but not in C. speratus. In addition, C. roberti is currently considered aphenotypic variant of C. spinosus because they share several characters and there is clear intergradation amongdistinct phenotypes (Caldara Jr. & Leite 2012). Voss (2011) appropriately designated a neotype for C. spinosus andconsidered C. roberti as a junior synonym. Natural history and ecology. Coendou speratus is sympatric with its larger congener C. prehensilis, but theformer appears to prefer the lower forest strata. Coendou speratus dens in hollow trees and has a strong andcharacteristic pungent odor, which is much stronger than the odor of sympatric C. prehensilis. Seven C. speratusindividuals were sighted in four forest fragments. The first sighting occurred at Boca da Mata, when an individualwas spotted at 11:29 pm, resting on a 12 meter-high horizontal branch; the second was at Mata Xanguazinho, alsoA NEW SPECIES OF PORCUPINE FROM BRAZIL Zootaxa 3636 (3) © 2013 Magnolia Press · 433
  • 14. of a single individual resting at 10:27 pm, on a 10 meter-high branch; the third was at Mata Tauá, when twoindividuals were sighted at 00:20 am, resting together on a 20 meter-high branch; the fourth sighting was also atMata 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 00:38am, 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 Coendouinsidiosus (Cins), C. nycthemera (Cnyc), and C. speratus (Cspe). Coendou speratus seems to be solitary, although two individuals were sighted together once, probably a maleand a female, one of which vocalized at the observer’s presence. At all times when C. speratus was sighted, itbecame either motionless or moved very slowly and then became motionless, whereas Coendou prehensilisescaped quickly at all times. Coendou speratus was once sighted 3 meters from a collared anteater, Tamanduatetradactyla (Linnaeus). According to locals, the new species feeds on the fruits of the exotic African oil palmElaeis guineensis Jacq. (locally known as dendezeiro). Previous long-term line transect surveys carried out between 2000 and 2008 in the Pernambuco EndemismCenter (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-sizedmammal 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 thefragment or degree of isolation, which implied that the occurrence of a mammal species in a given forest patchvaried unpredictably, and that in this ongoing process of mass extinction the 21st century medium-sized remainingmammalian fauna will soon be extinct if strict conservation measures are not implemented (Gadelha 2009; Melo2009).434 · Zootaxa 3636 (3) © 2013 Magnolia Press PONTES ET AL.
  • 15. The new subsequent line transect surveys that resulted in the discovery of this new species were carried out inthe highly impacted 4,000 ha forest fragments at Usina Trapiche, between 2008 and 2011 (Gadelha 2009; Melo2009; Leite et al. 2011; Freitas 2012), with a sample effort of 302.5 km walked (diurnal and nocturnal) in nineforest fragments, totaling 68 surveys in 610 h. Reinforcing our previous findings that in this region the species areexpected to occur in the forest fragments in an unpredictable fashion, density and biomass derived from 143 km ofnocturnal surveys, revealed that Coendou speratus had comparatively high local densities in the four fragmentswhere it was encountered. In the largest fragment, Mata Xanguá (470 ha), C. speratus had the highest density (36.8ind/km²), while in the smallest fragment, Boca da Mata (94 ha), it had the lowest (5 ind/km²). Coendou speratuswas found syntopically with C. prehensilis in all but one forest fragment, Mata Tauá (280 ha), and the biomasscontribution 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 ofendemism highlights the importance of this hotspot for the conservation of the earth’s biota. It also points to thelack of information on its mammalian fauna, and the need for surveys in order to understand their distribution andstatus throughout the region (Costa et al. 2005). Human impact is extremely high in the region where we found this new porcupine (Melo 2009), resulting inthe regional extinction of large mammals, such as jaguar (Panthera onca), Brazilian tapir (Tapirus terrestris), giantanteater (Myrmecophaga tridactyla), and white-lipped peccary (Tayassu pecari) (Mendes Pontes 2009). The threatlevel to other medium-size mammals, like the blond capuchin monkey Sapajus flavius (Schreber) (Mendes Ponteset al. 2006; Alfaro et al. 2012), is currently still very high. Thus, long-term research programs and conservationmeasures must be implemented to assure the survival of the remaining species and protection of these forests.AcknowledgementsWe are grateful to Salgado and Trapiche mills, within which the forest fragments stretch, for their cooperation, forhaving granted some seed money essential for the accomplishment of this study, and also for having provided thenecessary infrastructure for the project. We are especially indebted to Mrs. Evânia Freire e Dr. Mário Jorge SeixasAguiar. Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) granted an undergraduatescholarship to JRG and a productivity fellowship to YLRL. In 2008 we received the recognition and financialsupport from Conservation International in Brazil, which has sponsored this study, both field inventories andmolecular analysis through the Critical Ecosystems Partnership Fund (CEPF). Fundação de Amparo à Pesquisa doEspírito Santo (FAPES) provided additional grant support to this project. ACL has a graduate scholarship fromCoordenação 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. CelsoAzevedo (UFES) provided helpful comments on taxonomic and nomenclatural issues, including the new scientificname proposed here. Carlos Eduardo D. Cintra (Systema Naturae Consultoria Ambiental Ltda.) kindly provided avaluable specimen of C. nycthemera for comparison and molecular analysis. Marcelo Weksler and two anonymousreviewers provided valuable comments that improved the quality of this manuscript. Instituto Chico Mendes deConservação 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 deAnimais Silvestres (ICMBio, Recife), kindly granted us the privilege to join the reintroduction to the wild of aspecimen of Coendou speratus sp. nov.ReferencesAlfaro, J.W.L., Silva Jr., J.S., & Rylands, A.B. (2012) How different are robust and gracile capuchin monkeys? An argument for the use of Sapajus and Cebus. American Journal of Primatology, 74, 273–286. http://dx.doi.org/10.1002/ajp.22007Asfora, P.H. & Mendes Pontes, A.R. (2009) The small mammals of the highly impacted north-eastern Atlantic forest of Brazil, Pernambuco Endemism Center. Biota Neotropica, 9, 31–35. http://dx.doi.org/10.1590/S1676-06032009000100004Bonvicino, C.R., Penna-Firme, V. & Braggio, E. (2002) Molecular and karyologic evidence of the taxonomic status of CoendouA NEW SPECIES OF PORCUPINE FROM BRAZIL Zootaxa 3636 (3) © 2013 Magnolia Press · 435
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  • 18. Vilela, R.V., Machado, T., Ventura, K., Fagundes, V., Silva, M.J.J. & Yonenaga-Yassuda, Y. (2009) The taxonomic status of the endangered thin-spined porcupine, Chaetomys subspinosus (Olfers, 1818), based on molecular and karyologic data. BMC Evolutionary Biology, 9, e29. http://dx.doi.org/10.1186/1471-2148-9-29Voss, R.S. (2011) Revisionary notes on Neotropical porcupines (Rodentia: Erethizontidae). 3. An annotated checklist of the species of Coendou Lacépède, 1799. American Museum Novitates, 3720, 1–36. http://dx.doi.org/10.1206/3720.2Voss, R.S. & Angermann, R. (1997) Revisionary notes on Neotropical porcupines (Rodentia: Erethizontidae). 1. Type material described by Olfers (1818) and Kuhl (1820) in the Berlin Zoological Museum. American Museum Novitates, 3214, 1–42.Voss, R.S. & Silva, M.N.F. (2001) Revisionary notes on Neotropical porcupines (Rodentia: Erethizontidae). 2. A review of the Coendou vestitus group with descriptions of two new species from Amazonia. American Museum Novitates, 3351, 1–36. http://dx.doi.org/10.1206/0003-0082(2001)351%3C0001:RNONPR%3E2.0.CO;2Woods, C.A. & Kilpatrick, C.W. (2005) Infraorder Hystricognathi Brandt, 1855. In: Wilson, D.E. & Reeder, D.M. (Eds), Mammal Species of the World: A Taxonomic and Geographic Reference. Vol. 2. 3rded., Johns Hopkins University Press, Baltimore, pp. 1538–1600.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 daVitória, Fazenda Pirataquissé (MN 11211); Porto Seguro (MN 55527). ESPÍRITO SANTO: São Mateus, Guriri (MBML2347); Linhares, Rio São José (MN 8277);C. nycthemera: PARÁ: Belém, Sítio Velho (MZUSP 13519). Cametá (MZUSP 5031, 5034, 5035 5036, 5037, 5038). Curralinho (MN4913, 4914, 4915, 4916, 4919, 4920, 4921, 4638, 4639, 4678, 4681, 4683, 4695). Santarém, Mata Virgem, Terra Firme (MN6530, 7652, 7653, 7654). TOCANTINS: Darcinópolis, Usina Hidrelétrica Estreito (UFES 2079).438 · Zootaxa 3636 (3) © 2013 Magnolia Press PONTES ET AL.