Coendou prehensilis Linnaeus 1758
Creators: name: Leite, Yuri L. R.
name: Júnior, Vilacio Caldara
name: Loss, Ana Carolina
name: Costa, Leonora Pires
name: Melo, Éverton R. A.
name: Gadelha, José Ramon
name: Pontes, Antonio Rossano M.
Publisher: name: Zenodo
Year: 2011
Subjects: Biodiversity
Taxonomy
Animalia
Chordata
Mammalia
Rodentia
Erethizontidae
Coendou
Coendou prehensilis

description: Coendou prehensilis (Linnaeus 1758)   
Figs. 2, 3.  
 Neotype. Flat skin, skull of an adult male housed at Museu Nacional, Universidade Federal do Rio de Janeiro, Brazil (MN 73383), collected by Antonio Rossano Mendes Pontes (field number ARMP 63) on 12 May 2009. Tissue sample fixed in ethanol is housed in the Animal Tissue Collection at Universidade Federal do Espírito Santo, Vitória, Brazil (UFES-CTA 1797).   
 Type locality. Mata Xanguá, Usina Trapiche, municipality of Sirinhaém, state of Pernambuco, Brazil, 8°38'50"S 35°10'15"W, elevation 100 m (Fig. 4). This area is in the Pernambuco Endemism Center (herein CEPE), the biogeographical unit of the Brazilian Atlantic forest located north of the São Francisco River in northeastern Brazil, which encompasses the states of Rio Grande do Norte, Paraíba, Pernambuco and Alagoas. The CEPE comprises an important endemism center in South America, housing numerous endemic species (sensu Prance 1982, 1987; Silva & Casteletti 2003), suggesting it is a hotspot within a hotspot, the Pernambuco refugium (Punde et al. 2008; Carnaval & Moritz 2009), a world priority for biodiversity conservation, and one of the most important hotspot on the planet (Myers et al. 2000). Usina Trapiche mill is one of the various scenarios of the 2% of what was left of the entire CEPE. It comprises an isolated archipelago of 5,202 ha of hundreds of small, irregular, impacted and isolated Atlantic forest fragments surrounded by a sugar-cane matrix. Mata Xanguá is the largest forest fragment (469.76 ha) of Usina Trapiche mill and is highly irregular shaped (shape index = 2.88; calculated following Laurance & Yensen 1991), and formed by lowland evergreen tropical rainforest (Oliveira & Fontes 2000).   
 Distribution. According to Cabrera (1961), the distribution of the nominotypical subspecies is eastern Venezuela, the Guianas, and eastern Brazil. Given that specimens from Surinam described by Husson (1978) match our description of the neotype (see below), we believe this is a good approximation of the geographic range of C. prehensilis. Woods & Kilpatrick (2005) extended the distribution of C. prehensilis westward to central Brazil, northern Argentina, eastern Paraguay, and Bolivia, but current data do not allow us to establish the western limits of its distribution, especially given that similar species probably occur in west-central Brazil (see below). Only a comprehensive taxonomic revision of the genus will shed light on the geographic range of C. prehensilis and congeners.   
  Description of the neotype. External  —Body measurements: HB = 460 mm, TL = 400 mm, HF = 76 mm, EL = 28 mm. Weight = 2,720 grams. Tricolored quills on the top of head yellowish white basally, black in the middle, and white or whitish yellow distally. Thin, black and sparse hairs concealed under 5–20 mm long quills. Quills short (2–12 mm) around the eyes and muzzle, slightly longer (19 mm) on cheeks, whitish basally and distally, and black in the middle (Fig. 3 C). Coarse hairs around nose short and brownish; slightly longer pelage on the chin. Mystacial vibrissae long and black, some extending well beyond the shoulder (120 mm). Subocular and supraorbital vibrissae 48–95 mm; genal vibrissae 44–82 mm; shorter submental vibrissae present. Bulbous muzzle very large (ca. 50 x 50 x 30 mm), soft and pinkish in the live animal. Conspicuously rounded ears nearly naked, covered with minute hairs; very short (10 mm) quills (either completely white, or black basally and white distally) around the tragus.   
Dorsal body surface densely covered with long quills (37–83 mm), gradually longer from shoulder to rump; with thin and sparse black hairs concealed underneath. Three patterns of quill bands on the shoulders: bicolored quills (white or whitish yellow base and black tip); tricolored quills (white or whitish yellow base, blackish middle, and whitish tip); and four-banded quills (same as previous, with an additional brownish tip). Quills on mid-dorsum bright yellow basally, black in the middle, and white to pale yellow distally, with different color proportions on each quill. Quills on rump tend to be dirty yellow distally. Quills gradually shorter and thinner from mid-dorsum towards the ventral surface. Basal bright yellow quill band gradually paler and finally white toward the venter; black middle band also gradually wider toward the venter. Transition between dorsal and ventral surface with three-banded bristles: white basally, brownish black in the middle, and white distally. Dense, 17 mm-long spinous hairs on ventral surface, white basally, brownish black in the middle, and white distally. Soft, long (ca. 20 mm) hairs on the lower venter, anterior to the hindlimbs, white basally and brownish distally. Conspicuous white line from mid-venter to the genitals (Fig. 3 A). Sparse, 50 mm black-and-white vibrissae on the belly.  
Fore- and hindlimbs densely covered with coarse hairs, black or brown basally and whitish distally; hairs become progressively stiffer toward the dorsal surface. Small (<20 mm) quills along the outer margin of the forelimbs become shorter and whitish basally toward the feet. Hindlimb quills concentrated near the rump; quills bright yellow basally, brownish black in the middle, and white distally. Postcranial vibrissae present, predominantly black basally and whitish distally, 20–32 mm long on the forelimbs and 30–52 mm on the hindlimbs. Plantar surface yellowish brown in the live animal; claws long, curved inward.  
Tail dorsally prehensile and slightly shorter than head-and-body. Caudal quills bright yellow basally, black in the middle, and white to pale yellow distally; distal band wider than others. Proximal one-third of tail covered dorsally with longer quills (ca. 50 mm), covered laterally with bicolored bristles (brownish black basally and white distally, stiffer and darker toward the ventral surface), and covered ventrally with brownish bristles. Midde of tail covered dorsally with shorter quills (ca. 35 mm), laterally with bristles darker than those on the proximal third. Distal third of tail naked and calloused dorsally, covered laterally and ventrally with relatively sparse black bristles (Fig. 3 B).  
 Skull —Frontal and nasal sinuses inflated dorsally (Fig. 1), from mid-nasals to fronto-parietal suture; parietals slightly inflated. Postorbital ridge well developed and slightly inflated; left and right temporalis scars are closely approximated for about 16 mm along the dorsal midline of the posterior braincase, but do not form a definitive sagittal crest; supraoccipital ridge well developed. Rostrum short and tall; nasal aperture heart-shaped in frontal view. Nasals short, anterior margins convex and longer medially. Naso-frontal suture U-shaped and anterior to the postorbital ridge. Gnathic process well developed; zygomatic arches curved and widest across the jugals in dorsal view. Lacrimal-sphenoid suture nearly touches sphenofrontal foramen. Jugal tall, tapering posteriorly in lateral view. Incisive foramina oval, contained completely by the premaxillae and adjacent to premaxillary suture. Longitudinal maxillary ridge extending from anteromedian edge of upper premolar to premaxillary suture. Anterior margin of mesopterygoid fossa extends to the middle of the third molars; bony roof of the fossa perforated by sphenopalatine fenestrae. Auditory bullae large and bean-shaped, constricted posteriorly and contacting paroccipital processes. Basoccipital width subequal to auditory bulla width. Distinct, inward-extending ridge on roof of external auditory meatus. Depression between occipital condyle and paroccipital process U-shaped. Cranial measurements in millimeters: CIL = 86.27; LD = 23.31; LIF = 5.11; BIF = 3.11; MTR = 19.11; LM = 14.50; BP4 = 4.86; BM1 = 5.09; APB = 6.46; PPB = 8.82; PZB = 49.78; HIF = 12.92; ZL = 31.27; LN = 28.53; BNA = 21.98; BB = 35.29; DI = 4.18; BIT = 7.42.   
 Dentition —Dental formula: I 1 /1, C 0/0, P 1/1, M 3/3. Anterior surface of incisors orange-yellow and remaining surfaces whitish. Maxillary toothrows nearly parallel and showing typical erethizontid occlusal surface (Fig. 1). Ancestral pentalophodont morphology (Vucetich & Verzi 1994) on maxillary teeth, where anteroloph, protoloph, mesoloph and metaloph link to posteroloph.   
 Comparisons. There is no other large porcupine with tricolored quills within the range of C. prehensilis. This taxon and C. bicolor Tschudi are the two largest species of Neotropical porcupines, measuring from about 380 to 560 mm of head-and-body length and weighting from to 3 to 5 kg (Emmons & Feer 1997). The neotype of C. prehensilis is on the lower range of these values, measuring 460 mm (HB) and weighing 2.7 kg. Despite its overall small size, this specimen has heavily worn molars, indicative of an old adult individual according to Voss & Angermann (1997). Coendou bicolor has typically bicolored spines, yellow-white basally and black distally (Emmons & Feer 1997).   
The pelage of the neotype matches Marcgrave’s overall description (Appendix 1): the dorsal surface is covered with yellowish-based quills that are blackish medially and whitish distally. The tail is slightly longer than headand-body in Marcgrave’s description, but the neotype has slightly longer head-and-body. Among the specimens from Surinam, housed at the Rijksmuseum van Natuurlijke Historie, Leiden (RMNH), and described by Husson (1978), one had longer head-and-body (RMNH 18299: HB = 487 mm, TL = 471 mm) and another had longer tail (RMNH 21902: HB = 460 mm, TL = 480 mm). In general, the pelage of the neotype agrees with Husson’s (1978) description of Surinamese material, albeit the dorsal quills are shorter (20 mm on the head and 83 mm on the rump) on the former than the latter (40 mm on the head and 100 mm on the rump). The detailed description of the tail given by Husson (1978) matches very well the pattern we found on the neotype (Fig. 3 B).  
The cranial morphology of the neotype is also very similar to Husson’s (1978) description of Surinamese material. The two main distinctions are: the posterior margin of the incisive foramen is anterior to the suture between the premaxillary and the maxillary in the neotype, and at this suture on Surinamese specimens; anterior margin of mesopterygoid fossa reaches mid-M3 on the neotype and between M2 and M3 on Surinamese specimens.  
 Phylogenetic analyses of DNA sequences. The strict consensus of the six most-parsimonious trees recovered by our analyses (length = 361 steps) showed a topology very similar to the ML tree (-Ln = 2654.6), but higher bootstrap support for nodes in general (Fig. 5). Only MP confirmed the monophyly of the three “ C. prehensilis ” sequences from Bonvicino et al. (2002), but with no bootstrap support. These sequences grouped with specimens identified as “ C. bicolor” in Genbank, diverging only 0.6% from them (Fig. 5). Together, they formed a well supported clade (100% MP and 90% ML bootstrap) that probably represents a single species, considering that up to 2,000 km separate some of them (Fig. 4). The sequence from the neotype of C. prehensilis diverged 4.9% on average from a sister-group comprising haplotypes from specimens identified as “ C. bicolor ” and “ C. prehensilis ” by Bonvicino et al. (2002). Coendou and Sphiggurus formed two reciprocally monophyletic groups, well supported in MP analysis (99% and 92%, respectively), showing 10.9% genetic divergence between them (Fig. 5). The reciprocal monophyly of Coendou and Sphiggurus had low support in the ML analysis (76% and 66%, respectively).
description: Published as part of Leite, Yuri L. R., Júnior, Vilacio Caldara, Loss, Ana Carolina, Costa, Leonora Pires, Melo, Éverton R. A., Gadelha, José Ramon & Pontes, Antonio Rossano M., 2011, Designation of a neotype for the Brazilian porcupine, Coendou prehensilis (Linnaeus, 1758), pp. 30-40 in Zootaxa 2791 on pages 33-36, DOI: 10.5281/zenodo.201871