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The Western Amazonian Dwarf Porcupine, Coendou ichillus Voss & Da Silva, 2001, is poorly known in South America. The species is known only from eight localities in the Amazon and Orinoquia regions of Brazil, Colombia, Ecuador, and Peru. In Colombia, it is known from one locality in the Orinoquía region, based on a specimen collected in 1956. We present a new distribution record which adds information about the presence of C. ichillus in the Napo Moist Forests ecoregion, in the Department of Putumayo, Amazon region of Colombia. We suggest that C. ichillus is more widely distributed in the Amazon region of the country. However, based on the number of known records to date, C. ichillus is the rarest species of porcupine.

The Brazilian Porcupine, Coendou prehensilis (Linnaeus, 1758), is the most widespread species of Coendou Lacépède, 1799 in South America, but little is known on its natural history, ecology and distribution. In Colombia, it has been reported in the lowlands and inter-Andean valleys of 18 continental departments, but there are still gaps on its distribution, natural history and ecology. We present new distribution records and add information of the presence of C. prehensilis in 10 additional departments of Colombia located at the Amazonia, Orinoco, Peri-Caribbean Arid Belt, and North Andean Biogeographic provinces. We suggest that C. prehensilis is the most widespread species in the lowlands of Colombia, although it does not occur in the Biogeographic Chocó and in the inter-Andean medium and high valleys of the Cauca and Magdalena rivers.

The Western Amazonian Dwarf Porcupine, Coendou ichillus Voss & Da Silva, 2001, is poorly known in South America. The species is known only from eight localities in the Amazon and Orinoquia regions of Brazil, Colombia, Ecuador, and Peru. In Colombia, it is known from one locality in the Orinoquía region, based on a specimen collected in 1956. We present a new distribution record which adds information about the presence of C. ichillus in the Napo Moist Forests ecoregion, in the Department of Putumayo, Amazon region of Colombia. We suggest that C. ichillus is more widely distributed in the Amazon region of the country. However, based on the number of known records to date, C. ichillus is the rarest species of porcupine.

Coendou prehensilis (Linnaeus, 1758) Brazilian Porcupine, Puercoespín Examined material. BOLIVIA; Province of Ñuflo de Chávez of Santa Cruz Department; Centro “Chiquitos”; 16.3695°S, 062.0052°W; first capture on 12 May 2017; secondary forest patch in pastureland. Identification. This is a medium-sized, arboreal rodent (2–5 kg). Thick spines, 2–6 cm long, cover the entire body except for the muzzle. The general color appears gray or yellowish from a distance, but each spine is whitish at its base and tip and with a dark-brown band in the middle. This is the only porcupine species known in the area., Published as part of Jansen, Martin, Engler, Marc, Blumer, Luka Moritz, Rumiz, Damián I., Aramayo, José Luis & Krone, Oliver, 2020, A camera trapping survey of mammals in the mixed landscape of Bolivia's Chiquitano region with a special focus on the Jaguar, pp. 323-335 in Check List 16 (2) on page 331, DOI: 10.15560/16.2.323

Coendou spinosus (F. Cuvier, 1823) Figure 29 Records. Record was in May 22, 2016 camera trap 12 (25°02.55ʹ S, 049°57.41ʹ W) and by opportunistic observations in the same monitoring site. Identification. Coendou spinosus was identified by coat and body shape. The coat is formed by a mixture of rigid, aculeiform (cylindrical ‘spines’) hairs and finer hairs. The former are longer than the latter ones. Coloration varies from yellowish to dark brown at the back., Published as part of Pereira, Alan Deivid, Bazilio, Sergio & Orsi, Mário Luís, 2018, Checklist of medium-sized to large mammals of Campos Gerais National Park, Paraná, Brazil, pp. 785-799 in Check List 14 (5) on pages 796-797, DOI: 10.15560/14.5.785

Coendou mexicanus (Kerr, 1792) Material examined. Table 2; Figure 11. Coendou mexicanus is the only erethizontid in southern Mexico (Hall 1981). The presence of spines on most of the body make this species unique in the region (Reid 2009). We classify this species as Coendou mexicanus and not Sphiggurus mexicanus following Voss et al. (2013) and Ramirez-Pulido et al. (2014)., Published as part of Hidalgo-Mihart, Mircea G., Contreras-Moreno, Fernando M., Jesús-de la Cruz, Alejandro, Juárez-López, Rugieri, Bravata de la Cruz, Yaribeth, Pérez-Solano, Luz A., Hernández-Lara, Carolina, Friedeberg, Diana, Thornton, Dan & Koller-González, Juan M., 2017, Inventory of medium-sized and large mammals in the wetlands of Laguna de Terminos and Pantanos de Centla, Mexico, pp. 711-726 in Check List 13 (6) on page 718, DOI: 10.15560/13.6.711, {"references":["Reid F (2009) A field guide to the mammals of Central America and Southeast Mexico. 2 nd ed. Oxford University Press, New York, 384 pp.","Voss RS, Hubbard C, Jansa SA (2013) Phylogenetic relationships of New World porcupines (Rodentia, Erethizontidae): implications for taxonomy, morphological evolution, and biogeography. American Museum Novitates 3769: 1 - 36. https: // doi. org / 10.1206 / 3769.2","Ramirez-Pulido J, Gonzalez-Ruiz N, Gardner AL, Arroyo-Cabrales J (2014) List of Recent land mammals of Mexico. Special Publications of the Museum of Texas Tech University 63: 1 - 69."]}

6. Quichua Porcupine Coendou quichua French: Coendou des Andes / German: Rothschild-Greifstachler / Spanish: Puercoespin quichua Other common names: Andean Porcupine, Rothschild’s Porcupine Taxonomy. Coendou quichua Thomas, 1899, “Puembo, Upper Guallabamba River, Province of Pichincha, Ecuador; altitude about 2500 metres.” In the past, C. quichua has been recognized as a subspecies of C. bicolor, but a set of conspicuous differences in phenotype warrants its recognition as a separate species. Furthermore, specimens from Panama were separately recognized as C. rothschildi but are phenotypically similar. Thus, it appears that C. quichuais a widespread and geographically variable species. Phylogenetic analysis of cytochrome-b sequence data places C. quichua as the sister species to C. rufescens and as part of a clade that includes C. mexicanus. Monotypic. Distribution. E Panama, trans-Andean Colombia, and NW Ecuador. Descriptive notes. Head—body 332-438 mm, tail 260-413 mm, ear 23-26 mm, hindfoot 61-78 mm; weight 2-3 kg. The Quichua Porcupine is medium-sized and salt-andpepper colored; it appears spiny because of a lack of emergent fur. Dorsum is blackish and uniformly speckled as a result of bicolored (whitish basally and black distally) and longer tricolored (whitish at base, black medially, with whitish tips) spines. Frontal sinuses are not or are only slightly inflated, and roof of external auditory meatus usually has well-developed transverse bony ridge. Tail lengths are 55-90% of head—body lengths. Tail is black, or speckled black, and white, with white on sides and appearing black ventrally. Spines of young Quichua Porcupines are covered by soft hair. There is geographical variation in appearance. Most Quichua Porcupines in Panama are more strongly speckled with white due to white-tipped dorsal spines, but some Panamanian specimens have so few white-tipped quills that they appear black. Quichua Porcupines from eastern Panama have brown-tipped dorsal quills. Panamanian specimens also have more inflated nasal sinuses, wider nasal apertures, and relatively smaller premolars. A few Quichua Porcupines from Ecuador are mostly blackish dorsally, lacking many pale-tipped quills, but most Ecuadorian specimens have cream, buffy, or brown-tipped dorsal quills. Quichua Porcupines from Ecuador also have much shorter tails (averaging c¢.55% of head-body length) than Panamanian specimens (c.85-90% head-body length), smaller hindfeet, and much narrower nasal apertures. Geographical variation may relate to the fact that most Ecuadorian specimens are from montane localities (above 2000 m) while most Panamanian specimens are from lowlands and foothills. Two Quichua Porcupines from Colombia differ from all others by having smooth roof of external auditory meatus. Habitat..owland evergreen forests, secondary forests, and cultivated areas from sea level to elevations of ¢.3300 m. Food and Feeding. Quichua Porcupines eat fruit and probably leaves. Breeding. There is no information available for this species. Activity patterns. There is no specific information available for this species, but the Quichua Porcupines is probably nocturnal and arboreal. Movements, Home range and Social organization. There is no information available for this species. Status and Conservation. Classified as Data Deficient (as C. quichua) and Least Concern (as C. rothschildi) on The IUCN Red List. Bibliography. Emmons (1997a), Voss (2011, 2015), Voss et al. (2013)., Published as part of Don E. Wilson, Thomas E. Lacher, Jr & Russell A. Mittermeier, 2016, Erethizontidae, pp. 372-397 in Handbook of the Mammals of the World – Volume 6 Lagomorphs and Rodents I, Barcelona :Lynx Edicions on page 377, DOI: 10.5281/zenodo.6603219

3. Brazilian Porcupine Coendou prehensilis French: Coendou du Brésil / German: Eigentlicher Greifstachler / Spanish: Puercoespin arboricola Other common names: Prehensile-tailed Porcupine Taxonomy. Hystrix prehensilis Linnaeus, 1758, “Habitat in Asia, America.” Restricted by Y. L. R. Leite and colleagues in 2011 to “Mata Xangua, Usina Trapiche, municipality of Sirthaém, state of Pernambuco, Brazil, 8°38’50” S, 35°10°15” W, elevation 100 m” by neotype selection. Coendou prehensilis appears to be most closely related to a clade of trans-Andean and Andean species including C. mexicanus, C. quichua, and C. rufescens. Several publications on molecular phylogenetics used sequence data from a C. prehensilis specimen misidentified as a C. bicolor, calling earlier results into question. Coendou prehensilis is monotypic, has a very large geographical distribution, and although there is very little genetic variation among specimens from northern South America, Amazonia, and the “cerrado” (bush savanna) biome, genetic divergence was apparent in samples from Pernambuco. Further molecular data are required to help discern whether or not the newly discovered C. baturitensis actually belongs to C. prehensilis, given that phenotypes overlap. Monotypic. Distribution. From most of Colombia (except W), Venezuela, and the Guianas, S throughout most of the forested cis-Andean lowlands of Ecuador, Peru, and Brazil to E Bolivia, N & E Paraguay, and NW Argentina, also on Trinidad [; it may also be found in Uruguay. Descriptive notes. Head-body 444-560 mm, tail 330-578 mm, ear 20-29 mm, hindfoot 80-95 mm; weight 3.2-5.3 kg. The Brazilian Porcupine is large,salt-and-pepper in appearance, and has a long tail. It lacks emergent fur and thus appears spiny. Dorsum, including rump, is covered in bicolored and tricolored barbed defensive spines. Longest dorsal spines (60-110 mm) are tricolored white or pale yellow at bases and tips, with black or dark brown centers, and colors are of about equal length on each spine. Head is round, and face usually white. Pink nose and lips are large, soft, and bulbous; nasal aperture is very wide. Small ears are inconspicuous among sharp, short spines on head. Mystacial vibrissae are stout and long, reaching shoulders. Eyes are small and black, with faint dull red-eye shines. Frontal sinuses of the Brazilian Porcupine are highly inflated, external auditory meatus is weakly keeled, and upperincisors are distinctly procumbent. Tail is long; tail length is 100% of head-body length. Robust tail is dorsally prehensile, distal one-third of dorsalside is naked, remainderis covered in short spines, and it is whitish at the base. Tail is often held curled around branches in an up-curling coil. Venter is covered with short, pale gray-brown or whitish soft spines. Short spines cover legs. Feet are broad with large pads opposing four long, strong claws and appear gray-brown above. Young Brazilian Porcupines are born with long, soft red-brown or brown hair that partially covers spines. Individuals from the Peruvian Basin may appear darker, with small white tips on their spines. Habitat. Old second growth or particularly vine-covered areas of lowland rainforest or tropical dry forests and riparian woodlands in savanna landscapes. Brazilian Porcupines are usually found in midto upper canopies. Food and Feeding. The Brazilian Porcupine eats fruit, immature seeds, buds, bark, green or ripe fruits, and some leaves, butit is less folivorous than smaller New World porcupines. Breeding. In captivity, female Brazilian Porcupines give birth aseasonally to a single precocial young after 195-200 days of gestation. Periods ofrainfall can cause peaksin births. Young weigh 360-450 g at birth and are nutritionally independent at c.15 weeks after birth. During the day, young are left alone in a sheltered place but are nursed at least once per day. Age offirst reproduction of females is ¢.19 months, and individuals can reproduce for more than twelve years. Activity patterns. Brazilian Porcupines are arboreal and nocturnal. They are often immobile and difficult to spot. They rest diurnally in retreats in hollow trees or in shady locations in the forest subcanopy. They are surprisingly agile and climb quickly. Movements, Home range and Social organization. Brazilian Porcupines occur at relatively high densities as indicated by large numbers rescued during construction of a hydroelectric dam. In the Venezuelan Llanos, individuals have been reported to range over 15-20 ha. In one study, home ranges were 8 ha and 38 ha for two radio-tracked males and 10 ha for one female. Home ranges of males overlapped with female’s home range. Brazilian Porcupines used different sleeping sites each day. Distance between trees required some nocturnal travel on the ground. Brazilian Porcupines are solitary but socially tolerant and communicate vocally and by scent marking. Social calls include whistles and a long, low-amplitude moaning cry. Males and females rub, and males mark females with urine. Diurnal restsites include large hollow tree trunks or other cavities, and females are occasionally found with their young. Status and Conservation. Classified as Least Concern (as C. prehensilis) and as Data Deficient (as C. sanctamartae) on The IUCN Red List. Feral dogs regularly eat Brazilian Porcupines in summerin Brazil. Presence of azole-resistant yeast fungus, Candida albicans, could indicate exposure to agricultural azole antifungals in the wild or in captivity, or exposure to environmental contaminants or chemical compounds. Bibliography. Arita et al. (1990), Bonvicino, Penna-Firme & Braggio (2002), Campos et al. (2007), Castelo-Branco et al. (2013), Dobson & Yu Jinping (1993), Eisenberg & Redford (1999), Emmons (1997a), Feijé6 & Langguth (2013), Leite et al. (2011), Miles et al. (1981), Montgomery & Lubin (1978), Vilela et al. (2009), Voss (2011, 2015)., Published as part of Don E. Wilson, Thomas E. Lacher, Jr & Russell A. Mittermeier, 2016, Erethizontidae, pp. 372-397 in Handbook of the Mammals of the World – Volume 6 Lagomorphs and Rodents I, Barcelona :Lynx Edicions on pages 375-376, DOI: 10.5281/zenodo.6603219

16. Roosmalen’s Porcupine Coendou roosmalenorum French: Coendou des Roosmalen / German: Roosmalen-Greifstachler / Spanish: Puercoespin de Roosmalen Other common names: Roosmalen’s Dwarf Porcupine Taxonomy. Coendou roosmalenorum Voss & da Silva, 2001, “Novo Jerusalem near the left bank of the middle Rio Madeira in the Brazilian state of Amazonas,” Brazil. As with C. ichillus, C. pruinosus, and C. vestitus, C. roosmalenorum is placed in the “ C. vestitus group” based on presence of soft dorsal fur, bristle-quills, and defensive quills. Monotypic. Distribution. W Amazonian Brazil, known from both banks of the Rio Madeira in Amazonas and Rondonia states between 5° S and 9° S and likely distributed widely S of the Amazon River. Descriptive notes. Head—body 290 mm,tail 260 mm, hindfoot 53 mm; weight ¢.600 g. Roosmalen’s Porcupine is one of the smallest erethizontids. Like other members of the C. vestitus group,it has soft fur covering dorsal spines and bristle-quills. Long dorsal fur is medium-to-dark brownish or grayish. Bicolored spines are yellowish at bases, with dark-brown tips, and cover entire dorsum. Interspersed over dorsum are 40-70 mm, primarily tricolored bristle-quills that are pale yellowish or white at bases and tips, with bands of dark brown in middles. Head is also covered with quills and has fine mystacial vibrissae. Frontal sinuses may or may not be inflated, and dorsal roof of external auditory meatus may or may not show a weak keel. Tail is long, with tail length ¢.90% of head-body length. Base of tail has woolly fur and bicolored quills on its dorsum. Middle section of tail is covered with blackish bristles on upper and lower surfaces, and prehensile tail tip is naked and calloused dorsally. Venter is densely covered with coarse bicolored and tricolored hairs, rooted in clusters, and fine brown wool. Hands and feet have dense cover of coarse black or dark brown hairs dorsally. Habitat. Lowland rainforest. Food and Feeding. There is no specific information available for this species, but a captive individual consumed a variety of seeds and fruit. Breeding. There is no information available for this species. Activity patterns. Roosmalen’s Porcupine is nocturnal; a semi-captive individual slept in a tree hole during the day. Movements, Home range and Social organization. There is no information available for this species. Status and Conservation. Classified as Data Deficient on The IUCN Red List (as Sphiggurus roosmalenorum). Bibliography. Voss (2015), Voss & da Silva (2001), Voss et al. (2013)., Published as part of Don E. Wilson, Thomas E. Lacher, Jr & Russell A. Mittermeier, 2016, Erethizontidae, pp. 372-397 in Handbook of the Mammals of the World – Volume 6 Lagomorphs and Rodents I, Barcelona :Lynx Edicions on page 380, DOI: 10.5281/zenodo.6603219

1. Broomstraw-spined Porcupine Chaetomys subspinosus French: Coendou subépineux / German: Borstenbaumstachler / Spanish: Puercoespin espinoso Other common names: Bristle-spined Porcupine, Bristle-spined Rat, Thin-spined Porcupine Taxonomy. Hystrix subspinosa Olfers, 1818, “Brazil.” Restricted by R. S. Voss and R. Angermann in 1997 to Salvador, Bahia, Brazil. Placed in the family Echimyidae by B. D. Patterson and A. E. Wood due to alleged retention of deciduous premolars; however, these teeth are not actually retained. Recent molecular evidence, presence of a posterior carotid foramen (shared with all erethizontids), and a prehensile manus and pes structure similar to Coendou support inclusion in the Erethizontidae. Monotypic. Distribution. SE Brazil, from Sergipe to N Rio de Janeiro states. Descriptive notes. Head-body 360-450 mm, tail 260-275 mm; weight 1-2 kg. The Broomstraw-spined Porcupine is medium-sized and pale to dark brown, with long back and short legs. Head is round, with small ear pinnae that are usually difficult to see because they are buried in spines on the head; brown tufts of soft fur emerging above spines reveal pinnae. This is the only soft fur on the body. Medium-length mystacial vibrissae reach to ears. Muzzle is brown and almost naked. Somewhat bulbous nose is not greatly swollen. Skull is heavily built, with cranial sutures that remain evident in adults. Well-developed postorbital process of frontal bone almost touches well-developed postorbital process ofjugal bone. Head and shoulders are evenly and densely covered with short (c.15 mm), kinky spines that stand up and are sharp at tips. Spines extend onto dorsal pelage to rump and on legs and base oftail, and in contrast to spines on head, these are non-defensive, longer (c.50 mm) spines lacking hard, sharp points; they are stiff, slightly wavy dry bristles like straws on a broom. Spines are tricolored with pale yellow bases, dark brown middles, and pale brown tips. Tail lengths are ¢.60-70% of head-body lengths. Tail, thick at the base, is colored like back on proximal one-fifth of dorsal side and rusty colored on ventral side near base, becoming dark brown near tip. Tail tapers, becoming slender distally, and curls dorsally for prehension (although it lacks well-developed prehensile surface seen in some New World porcupines). Other than the base, tail is thinly haired, becoming almost naked nearits tip. Habitat. Patches of native primary and secondary rainforest, gallery forest, and “restinga forest” (low-canopy coastal forest including palms and mangroves) with high vertical complexity. Although Broomstraw-spined Porcupines are found in degraded thickets lacking trees and “cabruca” plantations (cacao orchards with intact native tree canopies), these habitats are not preferred. They may co-occur with but are less common than the smaller Bahia Porcupine (Coendou insidiosus). Preponderance of soft bristles covering the body rather than more sharp quills may limit the Broomstrawspined Porcupine’s ability to exploit some habitat resources used by the Bahian Porcupine and the Brazilian Porcupine (C. prehensilis) due to an increased risk of predation; as a result, the Broomstraw-spined Porcupine appears to have more arboreal and restricted habitat requirements than other Neotropical erethizontids. Food and Feeding. The Broomstraw-spined Porcupine is a somewhat specialized folivore that prefers protein-rich and fibrous leaves from trees in the family Fabaceae —a diet that is also high in plant secondary metabolites. It is likely a hindgut fermenter based on its simple stomach anatomy and its long, large intestine and sacculated cecum. Feeding observations of Broomstraw-spined Porcupines revealed that they eat leaves from locally abundant and widespread pioneer tree species typical of early successional forests, indicating an ability to find food in disturbed forests. Breeding. There is no information available for this species. Activity patterns. Broomstraw-spined Porcupines are nocturnal and arboreal. They are active between 17:30 h and 05:40 h and show bimodal peaks of activity between 19:00 h and 20:00 h and again from c.03:00-04:00 h. During the day, they rest in sites covered by leaves and in tangles oflianas. Individuals observed in the wild spent 74% of their time resting, 14% feeding, 11% traveling, and 2% in other activities. Females met with their offspring for 20-170 minutes each night (21:00-02:00 h) but were otherwise solitary. Movements, Home range and Social organization. Broomstraw-spined Porcupines are excellent climbers; they use their prehensiletail to assist in arboreal movement. They move slowly; their reduced activity levels may be an energy saving strategy to help compensate for a diet high in plant secondary metabolites and fiber. Broomstraw-spined Porcupines move through tree canopies c.10 m off the ground and on small diameter (lesser than 2 cm) substrates, with their tails usually coiled around branches or twigs. They descend to the ground to defecate, and deposit feces and urine in concealed latrines such as bromeliads and tree hollows. Average distance traveled per day for radio-tracked individuals was 277-5 m at a rate of 21-6 m/h in one study and 147 m/night in a second study. Activity, distance traveled, and home range size do not appear to vary with season, although Broomstraw-spined Porcupines spend more time resting and less time feeding on rainy nights. Broomstraw-spined Porcupines have relatively small home ranges of 0-5-9-5 ha, using minimum convex polygon methods; this may be the smallest home range size among Neotropical porcupines. Mean home range of adult males (6-6 ha) is at least three times larger than mean home range of adult females (2 ha). Most (94-99%) of a male’s home range overlaps with the home ranges of neighboring females. In contrast, a female’s home range typically overlaps only 13-45% with that of the neighboring male. Female home ranges overlapped very little with neighboring females, only up to 0-2%. Radio-tracked Broomstraw-spined Porcupines preferred secondary forest and were seldom found in highly modified vegetation types such as rubber plantations. They prefer large trees with moderate-to-high numbers of lianas and growing close to the forest edge and will frequently reuse individual trees in their home ranges. Some individuals prefer intermediate forest strata for diurnal rest sites. Social structure of Broomstraw-spined Porcupines is little studied, but during infrequent occasions in which individuals were seen together, it was usually a mother with her single offspring. Status and Conservation. Classified as Vulnerable on The IUCN Red List. The Broomstraw-spined Porcupine is also listed as vulnerable on the Brazilian red list due to declining population sizes. Its distribution formerly extended from Sergipe to northern Rio de Janeiro, but many local populations are now extinct due to deforestation. Less than 17% of native forest remains within the current distribution of the Broomstrawspined Porcupine. Hunting pressure may also be contributing to their decline. Bibliography. Bezerra et al. (2015), Castilho et al. (2013), Costa et al. (2005), Emmons (1997a), Falcdo et al. (2012), Faria et al. (2006), Fernandez, Barbanti et al. (2012), Fernandez, de Barros et al. (2015), Oliveira, P. A. et al. (2012), Patterson & Wood (1982), Ribeiro et al. (2009), Voss (2015), Voss & Angermann (1997), Wilson & Reeder (2005), Zortéa & de Brito (2010)., Published as part of Don E. Wilson, Thomas E. Lacher, Jr & Russell A. Mittermeier, 2016, Erethizontidae, pp. 372-397 in Handbook of the Mammals of the World – Volume 6 Lagomorphs and Rodents I, Barcelona :Lynx Edicions on page 374, DOI: 10.5281/zenodo.6603219

15. Western Amazonian Dwarf Porcupine Coendow ichillus French: Coendou a longue queue / German: Streifen-Greifstachler / Spanish: Puercoespin manchado Other common names: Streaked Dwarf Porcupine Taxonomy. Coendou ichillus Voss & da Silva, 2001, “Rio Pastaza,” Pastaza, Ecuador. Presence of soft fur in dorsal pelage, bristle-quills, and defensive quills places C. ichillus in the “ C. vestitus group” along with C. pruinosus, C. roosmalenorum, and C. vestitus. Monotypic. Distribution. Lowlands of E Ecuador and NE Peru but may occur throughout most of NW Amazonia N of the Amazon River and W of the Rio Negro. Descriptive notes. Head-body ¢.260-290 mm, tail 210-250 mm; no specific data available for body weight. The Western Amazonian Dwarf Porcupine is small, blackish-yellow, and spiny. Head carries some tricolored spines with ivory-white bases, dark middle bands, and whitish tips. Fine mystacial vibrissae are similar to those of the Blackish Hairy Dwarf Porcupine (C. vestitus). Emergent fur does not occur on body, but under close examination, short blackish woolly hairs are apparent, sparsely distributed among spines and bristles. Defensive quills are 30-40 mm long and yellowish at bases, with terminal one-half to onethird dark brown or blackish. There are bristle-quills, up to 80 mm long and densely scattered among defensive spines. Bristle-quills are yellowish at bases and tips and with dark-brown or blackish middle bands, resulting in streaked effect on dorsum. Tail of the Western Amazonian Dwarf Porcupine is relatively long, with tail length ¢.85% of head-body length. Dorsal base oftail is densely covered with short, bicolored quills, similar to those found on rump. Rump is also covered in a few sparse woolly hairs. Tricolored bristles extend along lateral and caudal surfaces of tail and converge on dorsal side to form an indistinct light colored (white or yellow) chevron in middle of tail. Prehensile tail tip is calloused and naked on dorsal side, with remainder densely covered with blackish bristles that are stiffer and denser under base oftail than elsewhere. Venter is covered in very short coarse bior tricolored hairs rooted most often in triples or,less often, in groups of 2—4 hairs. Hands and feet are covered dorsally with coarse blackish hairs. Habitat. Both paratype specimens collected in dense rainforest catchments less than 500 m above sea level. Western Amazonian Dwarf Porcupines have been observed at La SelvaJungle Lodge at 300 m above sea level on the north bank of the Rio Napo—an area oftall forest on high ground and swamp in valley bottoms. Food and Feeding. There is no information available for this species. Breeding. There is no information available for this species. Activity patterns. The Western Amazonian Dwarf Porcupine is nocturnal and arboreal. Movements, Home range and Social organization. There is no specific information available for this species, but the Western Amazonian Dwarf Porcupine can inhabit holes in hollow palm trees that are being used as building supports. Status and Conservation. Classified as Data Deficient on The IUCN Red List (as Sphiggurus ichillus). Bibliography. Voss (2015), Voss & da Silva (2001), Voss et al. (2013), Published as part of Don E. Wilson, Thomas E. Lacher, Jr & Russell A. Mittermeier, 2016, Erethizontidae, pp. 372-397 in Handbook of the Mammals of the World – Volume 6 Lagomorphs and Rodents I, Barcelona :Lynx Edicions on pages 379-380, DOI: 10.5281/zenodo.6603219

17. Black-tailed Porcupine Coendou melanurus French: Coendou a queue noire / German: Schwarzschwanz-Greifstachler / Spanish: Puercoespin de cola negra Other common names: Black-tailed Hairy Dwarf Porcupine Taxonomy. Cercolabes melanurus Wagner, 1842, “Rio Negro,” Amazonas, Brazil. Phylogenetic analysis using cytochrome-b sequence data indicates that C. melanurus is the sister taxon to the “ C. vestitus group” including C. vestitus, C. pruinosus, C. ichillus, and likely C. roosmalenorum. Coendou melanurus is sympatric with the larger C. prehensilis. Monotypic. Distribution. NE Amazonian lowlands N of the Amazon River and E of Orinoco and Rio Negro rivers in E Venezuela, the Guianas, and N Brazil. Descriptive notes. Head-body 280-380 mm, tail 220-363 mm, ear 7-24 mm, hindfoot 52-75 mm; weight 1.5-2.4 kg. The Black-tailed Porcupine is small and blackish, streaked with yellow. Long emergent dorsal fur is blackish and streaked with long, yellowish-tipped guard hairs that cover bicolored defensive spines. Fur is pale and is thickest on shoulders. Head of the Black-tailed Porcupine is finely grizzled; mystacial vibrissae are slender and fine. Frontal sinuses are seldom inflated, and roof of external auditory meatus has well-developed transverse bony ridge. Tail is relatively long and pitch-black; tail lengths are 80-95% of head-body lengths. Rump is covered with thick yellow defensive spines that may be hidden by fur. Venter is pale gray-brown and frosted with white, and legs and feet are grizzled dark gray-brown or blackish. Habitat. L.owland rainforest. Food and Feeding. There is no information available for this species. Breeding. There is no information available for this species. Activity patterns. The Black-tailed Porcupine is arboreal and nocturnal. It likely uses high forest canopy at least 15-20 m above ground whereit is difficult to detect. Movements, Home range and Social organization. The Black-tailed Porcupine may be locally abundant as indicated by large numbers rescued during construction of a hydroelectric dam. Status and Conservation. Classified as Least Concern on 7he IUCN Red List (as Sphiggurus melanurus). Bibliography. Eisenberg & Redford (1999), Emmons (1997a), Voss (2015), Voss et al. (2013)., Published as part of Don E. Wilson, Thomas E. Lacher, Jr & Russell A. Mittermeier, 2016, Erethizontidae, pp. 372-397 in Handbook of the Mammals of the World – Volume 6 Lagomorphs and Rodents I, Barcelona :Lynx Edicions on page 380, DOI: 10.5281/zenodo.6603219

12. Baturite Porcupine Coendou baturitensis French: Coendou de Baturité / German: Ceara-Greifstachler / Spanish: Puercoespin de Baturité Taxonomy. Coendou (Coendow) baturitensis Feljo & Langguth, 2013, “Community Sitio Barreiros, municipality of Aratuba, Baturite Range, Ceara, Brazil (4° 23’ 30.5” S, 39° 00’ 45.4" W).” This species is monotypic. Distribution. E Brazil, only known from Baturité Mts, Ceara State. Descriptive notes. Head-body 500 mm, tail 460 mm, ear 20 mm, hindfoot 80 mm; weight 3-5 kg. The Baturite Porcupine is medium-sized, densely covered with tricolor spines and lacking emergentfur, resulting in a somewhat salt-and-pepper spiny appearance. Body spines vary in length and thickness from snoutto tail. Long tricolored spines on dorsum are whitish at bases, followed by long intermediate brown bands and dirty white tips. On sides of body, spines are shorter and distinctly darker, due to short basal and distal bands of dirty white color and a long intermediate brown band. Venter and inner regions of limbs are covered with thin, flexible grayish-brown quills. Hard quills cover outer sides of limbs. Hands and feet are covered with stiff hairs rather than quills. Large nose is bulbous and soft. Quills conceal short ears, and short quills are located around eyes and muzzle, with longer quills on cheeks. Mystacial vibrissae are long and black, extending to shoulders. Broad rostrum of the Baturite Porcupine includes strong pneumatization in naso-frontal region, extending to anterior one-half of nasal bones and resulting in uniformly convex dorsal profile. As with other species of Coendou,tail is dorsally prehensile. Tail length is 92% of head-body length (only one individual measured). Upper side of proximate one-half of tail has the same covering of tricolored spines as dorsum. Distally, tail spines become gradually thinner and shorter, disappearing entirely toward tail tip. Upperside oftail is differentiated into thin, transverse bands of calloused skin. Proximal one-half of ventral surface oftail carries short, thin, and flexible quills that decrease in number distally. Habitat. Caatinga Moist Forest ecoregion. Food and Feeding. There is no information available for this species. Breeding. There is no information available for this species. Activity patterns. There is no information available for this species. Movements, Home range and Social organization. There is no information available for this species. Status and Conservation. The Baturite Porcupine has not been assessed on The IUCN Red List, but it appears to have a very restricted distribution in the Baturite Range. Its limited distribution coupled with habitat loss means it is probably at high conservation risk. Bibliography. Feij6 & Langguth (2013), Hance (2013), Nascimento & dos Santos (2014)., Published as part of Don E. Wilson, Thomas E. Lacher, Jr & Russell A. Mittermeier, 2016, Erethizontidae, pp. 372-397 in Handbook of the Mammals of the World – Volume 6 Lagomorphs and Rodents I, Barcelona :Lynx Edicions on page 379, DOI: 10.5281/zenodo.6603219

10. Bahian Hairy Dwarf Porcupine Coendou insidiosus French: Coendou de Bahia / German: Bahia-Greifstachler / Spanish: Puercoespin de Bahia Other common names: Bahia Porcupine, Bahia Hairy Dwarf Porcupine Taxonomy. Hystrix insidiosa Olfers, 1818, “Brazil.” Restricted by R. S. Voss and R. Angermann in 1997 to Salvador, northern Bahia, Brazil. Previously considered Sphiggurus insidiosus, but lack of diagnosable characters to reliably differentiate Sphiggurus from Coendou led it to be placed within Coendou. Coendou insidiosus has been hypothesized to be the sister species to C. spinosus. It is likely sympatric with C. spinosus (thus non-vouchered specimens may not be reliable in areas of co-occurrence) and Chaetomys subspinosus. Monotypic. Distribution. E Brazil; distribution poorly known but likely stretches from S Sergipe to extreme E Minas Gerais and N Rio de Janeiro. Descriptive notes. Head-body 290-350 mm, tail 180-222 mm; no specific data available for body weight. The Bahian Hairy Dwarf Porcupine is medium-sized and smoky brown, entirely covered with thick, soft fur and thus appearing spineless. Head 1s less furred, with some white showing at bases of spines; face is dark brown with black whiskers. Frontal sinuses are not inflated, and roof of external auditory meatus is smooth. Sharp, thick, bicolored defensive quills are hidden beneath fur and are pale to whitish basally, with distal one-half dark brown or occasionally tricolored with tips faintly orange. Dorsal fur is unicolored to base. Prehensile tail is short to intermediate in length, ranging from 50-75% of head-body lengths. Tail is colored like body for basal one-third, dark brown to black for middle one-third, and sparsely haired to tip on last one-third. Feet and venter are dark-to-medium brown, although often venter is sparsely covered with a mix of black and yellow hairs. The Bahian Hairy Dwarf Porcupine can be distinguished from the similar Paraguayan Hairy Dwarf Porcupine (C. spinosus) by presence of unicolored, dark vibrissae, and caudal bristles. Habitat. Primary and mature secondary rainforest, gallery forest, restinga forest (lowcanopy coastal forest of palms and mangroves), “macega scrub” (degraded thickets without large trees), “cabruca” plantations (cacao orchards with an intact canopy of native trees), and “capoeira” (dense young secondary growth). Food and Feeding. The Bahian Hairy Dwarf Porcupine eats fruit, ant pupae, and cultivated vegetables and roots. Breeding. There is no information available for this species. Activity patterns. The Bahian Hairy Dwarf Porcupine is nocturnal and arboreal. Movements, Home range and Social organization. There is no information available for this species. Status and Conservation. Classified as Least Concern on The IUCN Red List (as Sphiggurus insidiosus). The Bahian Hairy Dwarf Porcupine is hunted for meat and its quills, which are said to have medicinal properties. It is considered somewhat rare because it has a restricted distribution, but it can occur at high densities. Bibliography. Arita et al. (1990), Caldara & Leite (2012), Dobson & Yu Jinping (1993), Eisenberg & Redford (1999), Emmons (1997a), Voss (2015), Voss & Angermann (1997)., Published as part of Don E. Wilson, Thomas E. Lacher, Jr & Russell A. Mittermeier, 2016, Erethizontidae, pp. 372-397 in Handbook of the Mammals of the World – Volume 6 Lagomorphs and Rodents I, Barcelona :Lynx Edicions on page 378, DOI: 10.5281/zenodo.6603219

15. Western Amazonian Dwarf Porcupine Coendow ichillus French: Coendou a longue queue / German: Streifen-Greifstachler / Spanish: Puercoespin manchado Other common names: Streaked Dwarf Porcupine Taxonomy. Coendou ichillus Voss & da Silva, 2001, “Rio Pastaza,” Pastaza, Ecuador. Presence of soft fur in dorsal pelage, bristle-quills, and defensive quills places C. ichillus in the “ C. vestitus group” along with C. pruinosus, C. roosmalenorum, and C. vestitus. Monotypic. Distribution. Lowlands of E Ecuador and NE Peru but may occur throughout most of NW Amazonia N of the Amazon River and W of the Rio Negro. Descriptive notes. Head-body ¢.260-290 mm, tail 210-250 mm; no specific data available for body weight. The Western Amazonian Dwarf Porcupine is small, blackish-yellow, and spiny. Head carries some tricolored spines with ivory-white bases, dark middle bands, and whitish tips. Fine mystacial vibrissae are similar to those of the Blackish Hairy Dwarf Porcupine (C. vestitus). Emergent fur does not occur on body, but under close examination, short blackish woolly hairs are apparent, sparsely distributed among spines and bristles. Defensive quills are 30-40 mm long and yellowish at bases, with terminal one-half to onethird dark brown or blackish. There are bristle-quills, up to 80 mm long and densely scattered among defensive spines. Bristle-quills are yellowish at bases and tips and with dark-brown or blackish middle bands, resulting in streaked effect on dorsum. Tail of the Western Amazonian Dwarf Porcupine is relatively long, with tail length ¢.85% of head-body length. Dorsal base oftail is densely covered with short, bicolored quills, similar to those found on rump. Rump is also covered in a few sparse woolly hairs. Tricolored bristles extend along lateral and caudal surfaces of tail and converge on dorsal side to form an indistinct light colored (white or yellow) chevron in middle of tail. Prehensile tail tip is calloused and naked on dorsal side, with remainder densely covered with blackish bristles that are stiffer and denser under base oftail than elsewhere. Venter is covered in very short coarse bior tricolored hairs rooted most often in triples or,less often, in groups of 2—4 hairs. Hands and feet are covered dorsally with coarse blackish hairs. Habitat. Both paratype specimens collected in dense rainforest catchments less than 500 m above sea level. Western Amazonian Dwarf Porcupines have been observed at La SelvaJungle Lodge at 300 m above sea level on the north bank of the Rio Napo—an area oftall forest on high ground and swamp in valley bottoms. Food and Feeding. There is no information available for this species. Breeding. There is no information available for this species. Activity patterns. The Western Amazonian Dwarf Porcupine is nocturnal and arboreal. Movements, Home range and Social organization. There is no specific information available for this species, but the Western Amazonian Dwarf Porcupine can inhabit holes in hollow palm trees that are being used as building supports. Status and Conservation. Classified as Data Deficient on The IUCN Red List (as Sphiggurus ichillus). Bibliography. Voss (2015), Voss & da Silva (2001), Voss et al. (2013)

13. Blackish Hairy Dwarf Porcupine Coendou vestitus French: Coendou brun / German: Schwarzbrauner Greifstachler / Spanish: Puercoespin marrén Other common names: Brown Hairy Dwarf Porcupine Taxonomy. Coendou vestitus Thomas, 1899, “Colombia.” Phylogenetic analysis of cytochrome-b sequence data places C. vestitus in a clade with C. ichillus and C. pruinosus. These three species, together with C. roosmalenorum make up the “ C. vestitus group” and share presence of bristle-quills and soft fur in dorsal pelage. Monotypic. Distribution. C Colombia, currently known from only two specimens from the W foothills of E Andes Range in Cundinamarca Department but possibly also in the lower E Andean slopes at 1300-2600 m. Descriptive notes. Head—body 290-370 mm, tail 170-195 mm; no specific data are available for body weight. The Blackish Hairy Dwarf Porcupine is small and medium brown as a result of soft brown hair that covers spines. Two types of spines are present on dorsal pelage. Ordinary defensive quills are up to 37 mm long and pale yellow or whitish basally and, for most of their length, with short brown tips. Bristle-quills emerge through long dorsal fur and are bicolored, pale at their bases, and brown distally, blending almost invisibly with dorsal fur. Emergent fur is lacking on head and face, revealing white-tipped spines that are thick, sharp, and robust. Mystacial vibrissae are fine. Tail is short, with tail length ¢.50% of head-body length. Venter is a warm brown that is somewhat paler than dorsum and is thickly haired and lacks spines. Feet are colored like dorsum. Habitat. There is no information available for this species. Food and Feeding. There is no information available for this species. Breeding. There is no information available for this species. Activity patterns. There is no information available for this species. Movements, Home range and Social organization. There is no information available for this species. Status and Conservation. Classified as Data Deficient on The IUCN Red List (as Sphiggurus vestitus). The Blackish Hairy Dwarf Porcupine is extremely rare and known from only a few individual specimens. The region where specimens were originally collected has been almost entirely deforested. Bibliography. Emmons (1997a), Voss (2011, 2015), Voss & da Silva (2001), Voss et al. (2013)., Published as part of Don E. Wilson, Thomas E. Lacher, Jr & Russell A. Mittermeier, 2016, Erethizontidae, pp. 372-397 in Handbook of the Mammals of the World – Volume 6 Lagomorphs and Rodents I, Barcelona :Lynx Edicions on page 379, DOI: 10.5281/zenodo.6603219

2. North American Porcupine Erethizon dorsatum French: Porc-épic dAmérique / German: Nordamerikanischer Baumstachler / Spanish: Puercoespin de Norteameérica Other common names: Quiller, Quill Pig Taxonomy. Hystrix dorsata Linnaeus, 1758, “Habitat in Canada.” Seven subspecies are recognized. Subspecies and Distribution. E.d.dorsatumLinnaeus,1758—CanadaEofYukonTerritoryandBritishColumbiaProvince(exceptLabradorandNewfoundlandProvince)andNC&NEUSA. E.d.bruneriSwenk,1916—CUSA(GreatPlainsEoftheRockyMts). E.d.couesiMearns,1897—SWUSA(CArizona,NewMexico,andWTexasstates)andNMexico(Sonora,Chihuahua,Coahuila,NuevoLeon,Tamaulipas,NSinaloa,andNDurangostates). E.d.epixanthumBrandt,1835—WUSA(SWashingtonStateEtoIdaho,Montana.Wyoming,Nevada,Utah,WColorado,andStoCaliforniastates). E.d.myopsMerriam,1900—AlaskaandtheYukonTerritoryinNWCanada. E.d.nagriscensJ.A.Allen,1903—WCanada(BritishColumbiaProvince)andNWUSA(NWashingtonState). E. d. picinum Bangs, 1900 — NE Canada (Newfoundland and Labrador Province). Descriptive notes. Head—body 600-1300 mm, tail 160-250 mm, hindfoot 75-110 mm; weight 5-12 kg. The North American Porcupine is the largest member of the family Erethizontidae. Robust with round bodies and relatively short legs, North American Porcupines generally appear blackish, reddish, brownish, or yellowish at a distance; thick underfur and long guard hairs conceal many of defensive quills. Tail is short, with tail length ¢.31% of head-body length. Head and body are well protected by ¢.30,000 defensive quills that are distributed everywhere except on venter, inner limbs, face, and ears and are ¢.20-100 mm long. Quills are bicolored, creamy white or yellowish at bases with black tips, resulting in aposematic coloration when quills rest against the body. Quills are composed of keratin proteins fabricated during quill growth into a cylindrical outer shell with an interior foam core, and tips are barbed,as is the case for most defensive quills in erethizontids. Head of the Northern American Porcupine is small and round with a squarish muzzle. Mouth has two chambers. Yellow incisors are located in the outer chamber and cheek teeth in the inner chamber; lips meet across diastema, behind incisors, allowing an individual to gnaw on bark without holding its mouth open, which may reduce heat loss in cold weather. Thick and relatively shorttail is solidly muscular; dorsal surface is covered with defensive quills. Underside oftail is covered with stiff bristles that, when combined with muscular action of the tail, aid in climbing. Unlike other species of erethizontids,tail of the North American Porcupine lacks prehensile surface. Dorsal region at base oftail, called the rosette, is distinguished by absence of fur or guard hairs; naked skin in this area appears greasy due to secretions from sebaceous glands, and quills are somewhat shorter and more barbed, functioning as osmotrichia to aid in propagating the porcupine’s distinctive warning odor. When a porcupine erects its quills to enter a defensive posture, lack of hair in the rosette amplifies the “pin cushion” appearance. North American Porcupines are plantigrade; soles of feet are naked and textured. Feet have long claws (four on forefeet and five on hindfeet) that can close tightly around the footpad and enable grasping. These adaptations make North American Porcupines excellent climbers but somewhat awkward and slow walkers. Habitat. Variety of habitats across a large geographical distribution. In the northern part of its distribution, the North American Porcupine can be found in tundra and boreal forest. In the eastern part ofits distribution,it is found in deciduous, mixed, and coniferous forests. In the western and south-western parts of its distribution, it is found in hemlock (7T5uga, Pinaceae) forest, mixed coniferous forest, oak (Quercus, Fagaceae) forest, and scrub desert habitat. Individual habitat selection is sometimes a function of perceived predation risk; in the Great Basin, sleeping adults andjuveniles generally avoided open areas and moved into higherrisk foraging areas more often as body condition deteriorated. Food and Feeding. Diets of North American Porcupines change with season and location. During spring, they feed on emerging leaf buds and will descend to the ground to eat fresh stems of grasses, wildflowers, and sedges. It will also eat leaves, forbs, and herbs on the ground and in trees during spring and summer. During autumn, diets may include agricultural crops, such as corn and apples, and acorns. Winter diet of the North American Porcupine consists of inner bark and cambium of a variety of species of deciduous tree and foliage of select conifers. In Alaska, porcupines alternate between eating needles of white spruce (Picea glauca, Pinaceae) and inner cambium of paper birch (Betula papyrifera, Betulaceae). In eastern North America, important deciduous food trees include sugar maple (Acer saccharum, Sapindaceae), red maple (A. rubrum), American beech (Fagus grandifolia, Fagaceae), and yellow birch (Betula alleghaniensis) and important coniferous food trees include the highly preferred eastern hemlock (75uga canadensis), white pine (Pinus strobus), and balsam fir (Abies balsamea). In western parts of its distribution, preferred food trees include western hemlock (7. heterophylla), Douglas fir (Pseudotsuga menziesii), and ponderosa pine (PF. ponderosa); other pines, spruces, and firs are also eaten. Winter feeding on bark of trees can result in tree death and is one reason that North American porcupines are considered as pests throughout much of their distribution, although there is little evidence that their feeding results in significant economic damage. Selective feeding on dominant species in the forest may in fact relieve competition on subdominant tree species, increasing tree diversity. Diets of North American Porcupines are typically low in nitrogen and high in fiber and contain a variety of plant secondary chemical compounds including tannins and terpenes. Mechanisms for detoxifying plant secondary chemicals are not yet known. Nevertheless, the North American Porcupineis a hindgut fermenter with long intestine and enlarged, fermenting cecum. Distal colon is four times longer than that of the similarly sized North American Beaver (Castor canadensis), and food is retained for a relatively long time in the digestive system, about 1-5 times longer than whatis typical of an 11kg hindgut fermenter. In Alaska, North American Porcupines alternate between feeding on more nutritious but secondary chemical-laden white spruce and less nutritious but low in secondary chemical paper birch; they may switch to paper birch when detoxification pathways used to process spruce needles are saturated. Breeding. Female North American Porcupines become receptive to mating in autumn and experience an 8-12hour estrus during which mating takes place. Vaginal and perineal secretions carry olfactory signals that help male porcupines determine when a female is reaching peak receptivity. Mating system is female-defense polygyny, in which males compete for access to females. Male home ranges become larger in autumn when they seek mating opportunities; males will fight with one another to gain access to a receptive female. These fights can be vicious; porcupines are just as susceptible as nonporcupine aggressors to injury from quills. Reproductive success in male porcupinesis variable and appears to be correlated with body size. A male will guard a female for a few days surrounding estrus. He may climb into a tree and sit in branches below the female, occasionally showering her with urine; chemicals in the male’s urine may advance estrus. When a female porcupine has accepted a male as a mate, she raises her hindquarters to him and arches hertail above her rosette; doing so presents a relatively quillfree and hence safe approach for the male. Copulation takes place repeatedly over several hours. Vaginal plug forms an hour or two after copulations end. Purpose of the plug has not been studied, but it likely helps to block copulation by other males, prevents semen loss, and may also reduce chemical signaling by the female toward other males. Reproductive success is much less variable for females than males; typically, 95-99% of females in a population breed every year. Gestation of the North American Porcupine is long at c.210 days (compare with an average of 280 days for humans). Lengthy gestation may be required due to a porcupine’s inability to maintain a positive energy balance in winter; females lose weight (as a result of burning fat stores) during the same period in which they are developing a fetus. North American Porcupines give birth to a single offspring called a “porcupette.” Porcupettes are precocial, born with fur and quills. Soft quills harden within a few hours of birth. Although porcupettes begin consuming herbaceous vegetation within a few weeks of birth, they typically nurse for ¢.127 days. Thus, an adult female North American Porcupine typically spends 337 days/year pregnant or nursing. Because female porcupines need to feed in trees, they will leave their young in brush piles and return to them periodically through the night to nurse. Female porcupines typically have four nipples: one pair near armpits and the other pair on abdomen. Young apparently takes turn nursing from different nipples. When young are c¢.3 months old, they spend nights away from its mother. Activity patterns. North American Porcupines are arboreal and nocturnal and remain active year-round, enduring low winter temperatures (e.g. —39°C) while consuming woody plants low in nitrogen and high in fiber and plant secondary metabolites. To survive harsh winter conditions, porcupines rely on fat stores accumulated from late spring to late summer. During winter, fat mass declines from an average of 50% to 27%. Fat losses are minimized by lowering rates of energy expenditure, not body temperature. Adult males begin to lose mass during early breeding period in autumn. Movements, Home range and Social organization. Given the very large distribution of the North American Porcupine, space use varies depending on habitat and season and appears to be related to den-site and food availabilities, predation risk, and, in winter, snow depth. In British Columbia, North American Porcupines appear to prioritize den location over food availability, first choosing larger diameter Sitka spruce (Picea sitchensis, Pinaceae) for winter dens and then selecting forage trees (with a preference for western hemlock) based on proximity to a winter den. In the Great Basin Desert of Nevada and mixed deciduous and spruce-fir forests of Quebec, juvenile porcupines, at high risk of predation, select habitats that provide good cover. Body condition can also influence habitat selection; individuals will move into high-risk foraging areas when body condition declines. Snow depth appears to limit movement and thus home rangesize. During years of heavy snow in the Catskill Mountains of New York, winter home range averaged 7-4 ha. In contrast, during a year of light snow, winter home range averaged 64-9 ha, which did not differ statistically from the average non-winter home range size. In the desert-shrub habitat of southern Idaho, non-winter home range size averaged 23-1 ha for females and was 61-7 ha for one male; in winter, home range size for two animals followed by telemetry averaged 0-07 ha. Winter home ranges in Alaska average 171 ha, are larger than in more temperate zones, and do not differ between sexes or correlate with body mass. Larger winter home ranges in Alaska, despite presence of snow, may be explained by patchily distributed forage trees and low availability of den sites, coupled with potentially low predation risk and larger body size compared with those at lower latitudes. Although North American Porcupines come together to breed, and females are occasionally seen with presumed young, they are, on the whole,solitary animals. Finding food and avoiding predation are the focus of daily activities. Daily movementdistances do not appear to differ by sex or age. In deciduous forests of the Catskill Mountains, female North American Porcupines are strongly territoriality. Female territories do not overlap, although male territories overlap those of females and other males. In an unusual pattern for mammals, males are philopatric and females disperse. Juvenile females disperse from their mother’s territories in first autumn. Young males, in contrast, set up home ranges within or adjacent to territories of their mothers. Porcupines are sometimes observed to den together in either male-female or mother—offspring pairs. Very occasionally, and generally only when they need to escape inclement weather, North American Porcupines can be found denning in larger groups of 3-8 or more individuals. Mating system is female-defense polygyny, with males guarding females during the breeding season, butlittle is known about variation in mate choice from year to year, nor is there any information available on genetic structuring within populations. Porcupines are long-lived, some living beyond 20 years old in the wild. Recent development of microsatellite markers for the North American porcupine will allow exploration of questions about mating system, reproductive success, and possibility of kin selection during, for example, periods of group denning. Populations of North American Porcupines are reported to cycle, sometimes in synchrony with lagomorphs. Data collected between 1995 and 2004 from southern Ontario showed population lows of 0-02 captures/100 trap nights and highs of 0-08 captures/100 trap nights; densities are affected by local density of Fishers (Martes pennanti), the one predator that appears to specialize on porcupines. Status and Conservation. Classified as Least Concern on The IUCN Red List. Humans are the greatest threat to North American Porcupines because they are viewed as pests throughout their distribution and are often killed in response to foraging on valued trees and buildings, or in orchards. Cars frequently hit North American Porcupines because they are attracted to road salt and likely tend to stand their ground and erect their quills in response to threats. Bibliography. Anderson, Hall-Martin & Russell (1985), Baird et al. (2009), Barthelmess & Brooks (2010), Bar thelmess et al. (2013), Berteaux et al. (2005), Bowman et al. (2006), Caro (2009), Chou et al. (2012), Coltrane & Barboza (2010), Coltrane & Sinnott (2013), Coltrane et al. (2011), Cott & Mochnacz (2007), Craig & Keller (1986), Escalante et al. (2013), FirstVoices (2015), Gatica-Colima et al. (2014), Jones & Genoways (1968), Mabille & Ber teaux (2014), Naughton (2012), Pokallus & Pauli (2016), Roze (2009), Roze & lise (2003), Smith (1979), Sweitzer (1996), Tenneson & Oring (1985), Wall (2007), Whitaker & Hamilton (1998), Wilson & Ruff (1999), Wolfe (1990), Woods (1973), Zimmerling (2005), Zimmerling & Croft (2001)., Published as part of Don E. Wilson, Thomas E. Lacher, Jr & Russell A. Mittermeier, 2016, Erethizontidae, pp. 372-397 in Handbook of the Mammals of the World – Volume 6 Lagomorphs and Rodents I, Barcelona :Lynx Edicions on pages 374-375, DOI: 10.5281/zenodo.6603219

14. Frosted Porcupine Coendou pruinosus French: Coendou givré / German: Raureif-Greifstachler / Spanish: Puercoespin escarchado Other common names: Brown Hairy Dwarf Porcupine, Frosted Hairy Dwarf Porcupine Taxonomy. Coendou pruinosus Thomas, 1905, “Montanas de la Pedregosa,” Mé- rida, Venezuela. Recent cytochrome-b sequence data suggests that C. pruinosus may be closely related to C. ichillus, C. roosmalenorum, and C. vestitus. These species have soft emergent fur in dorsal pelage, bristle-quills, and defensive quills. Monotypic. Distribution. N & W Venezuela (Venezuelan Coastal Range and the foothills of the Serrania de Perija, lowlands of W Maracaibo Basin, and Cordillera de Mérida) and NC Colombia (E Andes Range and the adjacent Serrania de la Macarena). Descriptive notes. Head-body 320-380 mm; tail 190 mm; no specific data available for body weight. The Frosted Porcupine is medium-sized and covered in dark graybrown to blackish fur that covers spines and is frosted whitish at tips, especially on sides. Gray-brown head carries tricolored spines that are pale basally, dark in middles, and pale again at tips; a few of these tricolored quills also appear on shoulders. Head is finely ticked with white. Mystacial vibrissae are black and stiff. Frontal sinuses are not inflated, and roof of external auditory meatus is weakly keeled. Defensive quills on dorsum are very short, less than 25 mm, and mostly bicolored, pale yellow at bases with black tips. Long, thick, and white-tipped tricolored bristle-quills are sprinkled throughout dorsal fur. Tail is short to intermediate, with tail lengths 50-70% of head— body lengths. As with other species of Coendou, prehensile tail is naked dorsally but at tip (last 50 mm) only on the Frosted Porcupine. Tail base is pale yellow or whitish, grizzled with fine white spines. Tail tip and ventral surface oftail are black. Venteris thickly furred and lacks spines; fur is dark brown basally and heavily frosted with silver. Feet are also brown and grizzled with white. Habitat. Rainforest and cloud forest in mountains, foothills, and lowlands at elevations of 54-2600 m. Food and Feeding. There is no information available for this species. Breeding. There is no information available for this species. Activity patterns. There is no information available for this species. Movements, Home range and Social organization. The Frosted Porcupine nests in hollows of rotten tree trunks and perhaps in branches of trees. Four individuals were found together in a nest. Status and Conservation. Classified as Least Concern The IUCN Red List (as Sphiggurus pruinosus). The Frosted Porcupineis likely threatened in parts or all of its distribution by deforestation. Bibliography. Delgado & Gémez-Laverde (2008), Emmons (1997a), Voss (2011, 2015), Voss & da Silva (2001)., Published as part of Don E. Wilson, Thomas E. Lacher, Jr & Russell A. Mittermeier, 2016, Erethizontidae, pp. 372-397 in Handbook of the Mammals of the World – Volume 6 Lagomorphs and Rodents I, Barcelona :Lynx Edicions on page 379, DOI: 10.5281/zenodo.6603219

8. Eastern Amazonian Dwarf Porcupine Coendou nycthemera French: Coendou noir / German: Koopman-Greifstachler / Spanish: Puercoespin negro Other common names: Black Dwarf Porcupine, Koopman's Porcupine Taxonomy. Hystrix nycthemera Olfers, 1818, “Brazil.” Restricted by R. S. Voss and R. Angermann in 1997 to “eastern Amazonia (below Obidos) south of the main channel of the Rio Amazonas.” Recent phylogenetic analysis of cytochrome-b sequence data indicates that C. nycthemera is likely the sister species to the larger C. bicolor of the western Amazon. Coendou nycthemera probably occurs sympatrically with C. prehensilis, and its distribution may overlap with C. roosmalenorum, the other erethizontid inhabiting the eastern bank of the Rio Madeira. Monotypic. Distribution. Amazonian Brazil, in Amazonas and Para states E of the Rio Madeira and along the S bank of the Amazon River, including at least part of Marajo I. Distribution may extend into adjacent Atlantic coastal watershed of Maranhao State. Descriptive notes. Head-body 290-380 mm, tail 280-370 mm, hindfoot 59-75 mm; weight 0-95 kg. The Eastern Amazonian Dwarf Porcupine is sometimes called the “Black Dwarf Porcupine” due to its overall black color, and it is sometimes confused with the larger Bicolorspined Porcupine (C. bicolor) of the western Amazonian Basin. The Eastern Amazonian Dwarf Porcupine is spiny in appearance due to absence of emergent dorsal fur. Most spines are bicolored, white or yellow on basal one-third and black for remaining two-thirds of distal part. As a result, an individual appears yellow or white in regions where bases of spines are visible, such as head and forequarters. Many individuals have some tricolored defensive quills, with small white or pale redbrown tips, resulting in speckled appearance. Head is round, and ear pinnae are naked dorsally. Frontal sinuses are not inflated, and roof of external auditory meatus is smooth. Spines on rump above base oftail are short and upright, with sharp tips worn off resulting in black appearance of rump. Length of black prehensile tail is ¢.90% of head-body length. Ventral surface is dark brown or gray-brown and is covered with soft, hair-like bristles. Feet are black. Habitat. Lowland rainforest. Food and Feeding. There is no information available for this species. Breeding. There is no information available for this species. Activity patterns. There is no specific information available for this species, but short, worn spines at base oftail suggest that the Eastern Amazonian Dwarf Porcupine rests with its rump wedged against a surface. Movements, Home range and Social organization. There is no information available for this species. Status and Conservation. Classified as Data Deficient on The IUCN Red List. Less than 20% of the likely habitat of the Eastern Amazonian Dwarf Porcupine is protected. Bibliography. Azevedo-Ramos et al. (2006), Emmons (1997a), Voss (2011, 2015), Voss & Angermann (1997), Wilson & Reeder (2005)., Published as part of Don E. Wilson, Thomas E. Lacher, Jr & Russell A. Mittermeier, 2016, Erethizontidae, pp. 372-397 in Handbook of the Mammals of the World – Volume 6 Lagomorphs and Rodents I, Barcelona :Lynx Edicions on pages 377-378, DOI: 10.5281/zenodo.6603219

Injuries to humans caused by porcupines are rare. However, they may occur due to the proximity of urban areas and the animal’s habitat in areas such as the Floresta da Tijuca in Rio de Janeiro. Outdoor sports and leisure activities in areas close to forests or in the rain forest are also relevant for incidents of this kind and a better knowledge of the local forest fauna would prevent such undesirable accidents. Porcupine quills have microscopic barbs at their tips which facilitate skin penetration, but hampering their removal. Once the spines are lodged in tissue, the microscopic backward-facing deployable barbs at the tips cause trauma if anyone tries to remove them. Local haemorrhage and an inflammatory response to the contaminated foreign body occur. Depending on the time lapse in removing the spines either septic or sterile foreign body reactions may occur. There is also the risk of migration of the spines, where fatal cases have been reported in human and veterinary medicine. Herein we report two unusual cases of accidents involving humans and the South American porcupine. The Sphiggurus villosus spines removed from scalp skin were also documented through Scanning Electron Microscopy.

A new species of the murid rodent genus Thomasomys is described from the Cordillera Oriental of eastern Ecuador, a major range of the northern Andes that remains almost completely unexplored mammalogically. In order to provide a baseline for future faunal inventory work in these mountains, this report summarizes specimen data from the vicinity of Papallacta (0°22′S, 78°08′W), a small village located near tree line on the eastern (Amazonian) slope. In addition to the new Thomasomys, the local fauna includes 32 other species of nonvolant mammals, of which several (Akodon latebricola, Thomasomys cinnameus, and T. erro) have not been reported since their original descriptions in the 1920s. Taxonomic research based on this material indicates that at least six taxa currently treated as synonyms or subspecies should be recognized as valid species, including Thomasomys popayanus (possibly including T. nicefori) and T. praetor (both formerly synonymized with T. aureus), T. cinnameus and T. hudsoni (both f...

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 Tau��, Usina Trapiche, municipality of Sirinha��m, state of Pernambuco, Brazil, 8 �� 33 ' 46.13 "S 35 �� 10 ' 9.09 "W, elevation 85 m. Mata Tau�� 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, Sirinha��m, 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 Conserva����o 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 Jaboat��o dos Guararapes, approximately 50 km from Usina Trapiche; this animal was subsequently released in the Mata Xangu�� (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 V��rzea (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), V��rzea, 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 SNP 1 (for ���Servi��o Nacional de Peste��� # 1) and consists of the skin and skull of an adult female. It was collected during a plague survey in the vicinity of Vi��osa, 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 kidneyshaped auditory bullae are inflated anteriorly, and each one has a small external auditory meatus. The sphenopalatine vacuities are present in most specimens examined. 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 S��o Francisco river), which encompasses the coastal region of the states of Alagoas, Pernambuco, Para��ba 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 S��o 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 Maranh��o (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 6 G); 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 6 A���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 6 D). Some specimens (e.g., MN 72045) also have yellow based, brownish red tipped bicolor quills from head to mid-back (Figure 6 C). 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 6 E) to light yellow (e.g., MN 72046, Figure 6 F). 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. 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). 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 of a single individual resting at 10: 27 pm, on a 10 meter-high branch; the third was at Mata Tau��, when two individuals were sighted at 00: 20 am, resting together on a 20 meter-high branch; the fourth sighting was also at Mata Tau��, at 11: 55 pm, at a 10 meter-high branch; the fifth and the six were at Mata Xangu��, at 7: 50 pm and 00: 38 am, respectively, both on a 20 meter-high branch. 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 21 st century medium-sized remaining mammalian fauna will soon be extinct if strict conservation measures are not implemented (Gadelha 2009; Melo 2009). 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 Xangu�� (470 ha), C. speratus had the highest density (36.8 ind/km ��), while in the smallest fragment, Boca da Mata (94 ha), it had the lowest (5 ind/km ��). Coendou speratus was found syntopically with C. prehensilis in all but one forest fragment, Mata Tau�� (280 ha), an, Published as part of Pontes, Antonio Rossano Mendes, Gadelha, Jos�� Ramon, Melo, ��verton R. A., S��, Fabr��cio Bezerra De, Loss, Ana Carolina, Junior, Vilacio Caldara, Costa, Leonora Pires & Leite, Yuri L. R., 2013, A new species of porcupine, genus Coendou (Rodentia: Erethizontidae) from the Atlantic forest of northeastern Brazil, pp. 421-438 in Zootaxa 3636 (3) on pages 424-435, DOI: 10.11646/zootaxa.3636.3.2, http://zenodo.org/record/220373

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 Esp��rito Santo, Vit��ria, Brazil (UFES-CTA 1797). Type locality. Mata Xangu��, Usina Trapiche, municipality of Sirinha��m, 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 S��o Francisco River in northeastern Brazil, which encompasses the states of Rio Grande do Norte, Para��ba, 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 Xangu�� 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 (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; BP 4 = 4.86; BM 1 = 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-M 3 on the neotype and between M 2 and M 3 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)., Published as part of Leite, Yuri L. R., J��nior, Vilacio Caldara, Loss, Ana Carolina, Costa, Leonora Pires, Melo, ��verton R. A., Gadelha, Jos�� 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, {"references":["Linnaeus, C. (1758) Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Laurentii Salvii, Holmiae [= Stockholm], ii + 824 pp. Available from: http: // gdz. sub. uni-goettingen. de / dms / load / img /? IDDOC = 265100 (accessed on 10 April 2010).","Prance, G. T. (1982) Forest refuges: evidences from woody angiosperms. In: Whitmore, T. C. & Prance, G. T. (Eds), Biological diversification in the tropics. Columbia University Press, New York, pp. 137 - 158.","Prance, G. T. (1987) Biogeography of neotropical plants. In: Prance, G. T. (Ed), Biogeography and quaternary history in tropical America. Oxford Monographs on Biogeography. Clarendon Press, Oxford, pp. 175 - 196.","Silva, J. M. C. & Casteletti, C. H. M. (2003) Status of the biodiversity of the Atlantic forest of Brazil. In: Galindo-Leal, C. & Camara, I. G. (Eds), The Atlantic forest of south America: biodiversity status, threats, and outlook. CABS and Island Press, Washington, DC, pp. 43 - 59.","Punde, S., Godbole, A. & Sarnaik, J. (2008) Searching for hotspots within a hotspot - using global prioritization schemes at a regional level. An example from the Northern Western Ghats. Paper presented at the annual meeting of the International Congress for Conservation Biology, Convention Center, Chattanooga, TN, July 10, 2008. Available from http: // www. allacademic. com / meta / p 237444 _ index. html (accessed 6 June 2010).","Carnaval, A. C. & Moritz, C. (2009) Predicting diversity within hotspots to enhance conservation. Science, 323, 785 - 789.","Myers, N., Mittermeier, R. A., Mittermeier, C. G., Fonseca, G. A. B. & Kent, J. (2000) Biodiversity hotspots for conservation priorities. Nature, 403, 853 - 858.","Laurance, W. F. & Yensen, E. (1991) Predicting the impacts of edge effects in fragmented habitats. Biological Conservation, 55, 77 - 92.","Oliveira, A. T. & Fontes, M. A. L. (2000) Patterns of floristic differentiation among Atlantic forests in southeastern Brazil and the influence of climate. Biotropica, 32, 793 - 810.","Cabrera, A. (1961) Catalogo de los mamiferos de America del Sur. Revista del Museo Argentino de Ciencias Naturales Bernardino Rivadavia, 4, 309 - 732.","Husson, A. M. (1978) The mammals of Suriname. E. J. Brill, Leiden, 569 pp., 160 pls.","Woods, C. A. & Kilpatrick, C. W. (2005) Suborder Hystricognathi. In: Wilson, D. E. & Reeder, D. M. (Eds), Mammal species of the world: a taxonomic and geographic reference. 3 rd ed., Johns Hopkins University Press, Baltimore, pp. 1538 - 1600.","Vucetich, M. G. & Verzi, D. H. (1994) Las homologias en los disenos oclusales de los roedores Caviomorpha: un modelo alternativo. Mastozoologia Neotropical, 1 (1), 61 - 72.","Emmons, L. H. & Feer, F. (1997) Neotropical rainforest mammals: a field guide. 2 nd ed., University of Chicago Press, Chicago.","Voss, 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.","Bonvicino, C. R., Penna-Firme, V. & Braggio, E. (2002) Molecular and karyotypic evidence of the taxonomic status of Coendou and Sphiggurus (Rodentia: Hystricognathi). Journal of Mammalogy, 83 (4), 1071 - 1076."]}

Coendou vestitus Thomas, 1899 Coendou vestitus Thomas, 1899: 284 (original description). Coendou (Sphiggurus) vestitus: Tate, 1935: 307 (name combination). Coendou (Sphiggurus) vestitus vestitus: Cabrera, 1961: 603 (name combination). Sphiggurus vestitus: Honacki et al., 1982: 572 (name combination). DISTRIBUTION: I have examined specimens of Coendou vestitus from only two definitely identifiable localities, both of which are in the western foothills of the eastern Andean cordillera in the Colombian department of Cundinamarca. However, Alberico et al. (1999) report a specimen from Villavicencio, on the eastern side of the Andes in the department of Meta. REMARKS: This species may be closely related to Coendou ichillus, C. pruinosus, and C. roosmalenorum, all of which also have bristle-quills in addition to ordinary defensive quills and soft fur in their dorsal pelage (Voss and da Silva, 2001). Diagnostic comparisons with those species were provided by Voss and da Silva (2001), whose text should also be consulted for a detailed description of the skin and skull of C. vestitus. SPECIMENS EXAMINED: See Voss and da Silva (2001)., Published as part of Voss, Robert S., 2011, Revisionary Notes on Neotropical Porcupines (Rodentia: Erethizontidae) 3. An Annotated Checklist of the Species of Coendou Lacépède, 1799, pp. 1 in American Museum Novitates 2011 (3720) on page 1, DOI: 10.1206/3720.2, http://zenodo.org/record/5359512, {"references":["Honacki, J. H., K. E. Kinman, and J. W. Koeppl. 1982. Mammal species of the world, a taxonomic and geographic reference. Lawrence, KS: Allen Press."]}

Voss, Robert S. (2011): Revisionary Notes on Neotropical Porcupines (Rodentia: Erethizontidae) 3. An Annotated Checklist of the Species of Coendou Lacépède, 1799. American Museum Novitates 2011 (3720): 1, DOI: 10.1206/3720.2, URL: http://www.bioone.org/doi/abs/10.1206/3720.2

Coendou Lacépède, 1799 Coendou Lacépède, 1799: 11; type species Hystrix prehensilis Linnaeus, 1758, by monotypy. Coendus: E. Geoffroy St. -Hilaire, 1803: 157 (misspelling of Coendou Lacépède, 1799). Coandu: G. Fischer, 1814: 102 (misspelling of Coendou Lacépède, 1799). Sinethere F. Cuvier, 1823: 427; type species S. prehensilis (Linnaeus, 1758) by original designation. Sphiggure F. Cuvier, 1823: 427; type species S. spinosa F. Cuvier, 1823, by original designation. Sinoetherus: F. Cuvier, 1825: 256 (misspelling of Sinethere F. Cuvier, 1823). Sphiggurus: F. Cuvier, 1825: 256 (misspelling of Sphiggure F. Cuvier, 1823; but see Remarks, below). Coendu: Lesson, 1827: 290 (misspelling of Coendou Lacépède, 1799). Synethere: Lesson, 1827: 291 (misspelling of Sinethere F. Cuvier, 1823). Sinetheres: J. Fischer, 1829: 369 (misspelling of Sinethere F. Cuvier, 1823). Cercolabes Brandt, 1835: 391; type species Hystrix prehensilis Linnaeus, 1758 (proposed as a replacement name for Coendou Lacépède, 1799)., Published as part of Voss, Robert S., 2011, Revisionary Notes on Neotropical Porcupines (Rodentia: Erethizontidae) 3. An Annotated Checklist of the Species of Coendou Lacépède, 1799, pp. 1 in American Museum Novitates 2011 (3720) on page 1, DOI: 10.1206/3720.2, http://zenodo.org/record/5359512, {"references":["Linnaeus, C. 1758. Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. 10 th ed., vol. 1. Holmiae [Stockholm]: Laurentii Salvii.","Lesson, R. - P. 1827. Manuel de mammalogie ou histoire naturelle des mammiferes. Paris: Roret."]}

Coendou roosmalenorum Voss and da Silva, 2001 Coendou roosmalenorum Voss and da Silva, 2001: 24; type locality “ Novo Jerusalem near the left bank of the middle Rio Madeira in the Brazilian state of Amazonas.” Sphiggurus roosmalenorum: Woods and Kilpatrick, 2005: 1549 (name combination). DISTRIBUTION: Coendou roosmalenorum occurs on both banks of the Rio Madeira in the Brazilian states of Amazonas and Rondônia. Although specimens have only been collected at three localities (Voss and da Silva, 2001), photographs of a small porcupine closely resembling this species were taken in 1992 on the south bank of the Amazon near Alvarães (13°13′ S, 64°48′W), about 500 km NW of the type locality (Franz Kern, personal commun., 2011)., Published as part of Voss, Robert S., 2011, Revisionary Notes on Neotropical Porcupines (Rodentia: Erethizontidae) 3. An Annotated Checklist of the Species of Coendou Lacépède, 1799, pp. 1 in American Museum Novitates 2011 (3720) on page 1, DOI: 10.1206/3720.2, http://zenodo.org/record/5359512

Coendou melanurus (Wagner, 1842) Cercolabes melanurus Wagner, 1842: 360; type locality “Rio Negro [Barra]” (= Manaus), Amazonas, Brazil. Sphiggurus melanurus Gray, 1842: 262; type locality “ Brazil.” Synetheres (Sphiggurus) melanurus: Trouessart, 1881: 184 (name combination). Coendu melanurus: Trouessart, 1897: 622 (name combination). Coendou (Sphiggurus) melanurus: Tate, 1935: 307 (name combination). Coendou (Sphiggurus) insidiosus melanurus: Cabrera, 1961: 601 (name combination). Sphiggurus melanura: Bonvicino et al., 2002 (epithet spelled with incorrect gender). DISTRIBUTION: Apparently throughout the northeastern Amazonian lowlands (north of the Amazon and east of the Orinoco-Cassiquiare–Rio Negro), including eastern Venezuela, Guyana, Surinam, French Guiana, and northern Brazil (Voss et al., 2001: fig. 66). REMARKS: This species was misidentified as or treated as a synonym or subspecies of Coendou insidiosus by Cabanis (1848), Cabrera (1961), Husson (1978), and Woods (1993), among others. Voss et al. (2001) provided a detailed description of C. melanurus and illustrated diagnostic comparisons with C. insidiosus. Reports of C. melanurus from Ecuador and Colombia (Emmons and Feer, 1997; Alberico et al., 1999; Tirira, 1999) were based on misidentified material of C. ichillus and C. pruinosus, respectively (Voss and da Silva, 2001). SPECIMENS EXAMINED: See Voss et al. (2001: 136)., Published as part of Voss, Robert S., 2011, Revisionary Notes on Neotropical Porcupines (Rodentia: Erethizontidae) 3. An Annotated Checklist of the Species of Coendou Lacépède, 1799, pp. 1 in American Museum Novitates 2011 (3720) on page 1, DOI: 10.1206/3720.2, http://zenodo.org/record/5359512, {"references":["Gray, J. E. 1842. Descriptions of some new genera and fifty unrecorded species of Mammalia. Annals and Magazine of Natural History (1) 10: 255 - 267.","Husson, A. M. 1978. The mammals of Suriname. Leiden: Brill."]}

Sphiggurus spinosus (F. Cuvier) Table 1 Type Locality: Paraguay, along the Paran�� River. Distribution: Northeast Argentina, southeastern Brazil, Paraguay, northeastern Uruguay. New records: CAAZAPA: 17) Estancia Dos Mar��as, 12.2 Km SSO de las casas, Costa del R��o Tebicuary, 26 �� 47.719 ��� S 56 �� 32.979��W (UMMZ 174975). ITAPUA: 18) Estancia San Isidro, 6.6 km NE of houses, 26 �� 31.372 ��� S, 55 �� 51.767��W (GD 252). Figure 1. Taxonomy: Sphiggurus is a taxon in need of revision. Problems range from its distinction from Coendou Lac��p��de to the limits of its species, including S. spinosus. Although morphologically Sphiggurus is similar to Coendou (e.g., Handley & Pine 1992; but see Massoia & Vaccaro 1991), recent phylogenetic analysis of molecular data indicates that both constitute independent lineages (Bonvicino et al. 2002). Comments: To our knowledge, these specimens constitute the first records corroborated by voucher specimens for these departments of the Paraguayan Oriental Region. The specimen from Caazap�� was collected in riparian forest of the Tebicuary River; the specimen from Itap��a was caught by hand in an artificial prairie about 300 m from the closest forest island., Published as part of D'El��a, Guillermo, Mora, Ismael, Myers, Phil & Owen, Robert D., 2008, New and noteworthy records of Rodentia (Erethizontidae, Sciuridae, and Cricetidae) from Paraguay, pp. 39-57 in Zootaxa 1784 on page 52, DOI: 10.5281/zenodo.182407, {"references":["Handley, C. O. Jr. & Pine R. H. (1992) A new species of Coendou Lacepede, from Brazil. Mammalia, 56, 237 - 244.","Massoia, E. & Vaccaro, O. B. (1991) El estado sistematico de los generos Sphiggurus F. Cuvier, 1825 y Coendou Lacepede, 1799 (Mammalia, Rodentia, Histricomorpha, Erethizontidae). Boletin Cientifico, APRONA, 19, 47 - 52.","Bonvicino, C. R., Penna-Firme, V., & Braggio, E. (2002) Molecular and karyologic evidence of the taxonomic status of Coendou and Sphiggurus (Rodentia: Hystricognathi). Journal of Mammalogy, 83, 1071 - 1076."]}

Coendou quichua Thomas SPECIMENS COLLECTED: None. OTHER MATERIAL: I examined three NHRS specimens (A58/2822, A58/2962, A59/2962) collected by Ludovic Söderström in 1911 at ‘‘Tablon above Tumbaco’’ with recorded elevations of 9000– 11,000 ft [2744–3354 m]. TAXONOMY: Coendou quichua is a morphologically distinctive porcupine whose diagnostic characters were accurately described by Thomas (1899). Cabrera (1961), however, treated quichua as a subspecies of C. bicolor Tschudi without providing any justification for doing so. Although Emmons (1990), Albuja (1991), Tirira (1999), and Alberico et al. (1999) have subsequently recognized that quichua is a valid species, some checklists (e.g., Woods, 1993) continue to treat this name as a synonym of bicolor. To date, no rationale has been provided for the zoogeographically incoherent and morphologically divergent collection of taxa that Cabrera (1961) lumped together as Coendou bicolor. Although this name has been applied by authors to a wide range of morphologies, specimens collected in the vicinity of the Peruvian type locality (e.g., AMNH 147500, FMNH 65799) are distinctively large porcupines (ca. 900 mm total length) with tails that are almost as long as the combined length of head­and­body; the visible dorsal pelage consists entirely of bicolored (blacktipped) quills, of which those over the forequarters are conspicuously longer than those over the lower back and rump. By contrast, C. quichua is much smaller (ca. 600 mm or less) with a proportionately much shorter tail (approximately half the length of head­andbody) and tricolored (pale­tipped) dorsal quills that are not conspicuously longer over the forequarters than on the lower back and rump. Cranially, quichua has a proportionately narrower rostrum than bicolor, smaller orbits, less expanded jugals, and less inflated frontal sinuses. Other relevant morphological comparisons will be provided in an upcoming generic revision (Voss, in prep.), but the characters given here together with other traits mentioned by Emmons (1990) and Alberico et al. (1999) are sufficient for unambiguous identifications of these dissimilar taxa. REMARKS: The original specimen tag of NHRS A58/2822 notes that the animal was ‘‘found in the underbrush’’. Lönnberg (1913) originally reported this material as having been collected above Tumbaco, without mentioning the actual collecting site (Tablón). Cuniculus (Stictomys) taczanowskii (Stolzmann) SPECIMENS COLLECTED: None. OTHER MATERIAL: A single specimen that I have not examined (QCAZ 954) was collected in the páramo near Paso de Guamaní by G. Onore in 1993 (D. Tirira, personal commun.). TAXONOMY: The mountain paca is morphologically distinctive (Thomas, 1924) and was formerly distinguished generically (as Stictomys) from the lowland paca (Cuniculus paca). Although only a single species of mountain paca is currently recognized, no critical analysis of morphological or molecular data is currently available to test the hypothesis that C. taczanowskii (from Ecuador and Peru) is actually conspecific with populations from Venezuela and Colombia that were formerly known as C. sierrae (e.g., by Thomas, 1905; Krumbiegel, 1940). The International Commission on Zoological Nomenclature (ICZN, 1998) recently ruled that Cuniculus Brisson, 1762, is the oldest available name for pacas, previously referred by most American authors to Agouti Lacépède, 1799., Published as part of VOSS, ROBERT S., 2003, A New Species of Thomasomys (Rodentia: Muridae) from Eastern Ecuador, with Remarks on Mammalian Diversity and Biogeography in the Cordillera Oriental, pp. 1-48 in American Museum Novitates 3421 on pages 34-35, DOI: 10.1206/0003-0082(2003)4212.0.CO;2, http://zenodo.org/record/4734917, {"references":["Thomas, O. 1899. On new small mammals from South America. Annals and Magazine of Natural History 7 (3): 152 - 155.","Cabrera, A. 1961. Catalogo de los mamiferos de America del Sur. Revista del Museo Argentino de Ciencias Naturales '' Bernardino Rivadavia' ' (Ciencias Zoologicas) 4 (2): 309 - 732.","Emmons, L. H. 1990. Neotropical rainforest mammals: a field guide. Chicago: University of Chicago Press.","Albuja V. L. 1991. Lista de vertebrados del Ecuador: mamiferos. Revista Politecnica 16: 163 - 203.","Tirira S., D. 1999. Mamiferos del Ecuador [vol.] II. Quito: Museo de Zoologia, Pontifica Universidad Catolica del Ecuador.","Alberico, M., V. Rojas-Diaz, and J. G. Moreno. 1999. Aporte sobre la taxonomia y distribucion de los puercoespines (Rodentia: Erethizontidae) en Colombia. Revista de la Academia Colombiana de Ciencias Exactas, Fisicas, y Naturales 23 (supplemento especial): 595 - 612.","Woods, C. A. 1993. Suborder Hystricognathi. In D. E. Wilson and D. M. Reeder (editors), Mammal species of the world, 2 nd ed.: 771 - 806. Washington, DC: Smithsonian Institution Press.","Lonnberg, E. 1913. Mammals from Ecuador and related forms. Arkiv for Zoologi 8 (16): 1 - 36, pl. 1.","Thomas, O. 1924. Some notes on pacas. Annals and Magazine of Natural History 9 (13): 237 - 238.","Thomas, O. 1905. New Neotropical Molossus, Conepatus, Nectomys, Proechimys, and Agouti, with a note on the genus Mesomys. Annals and Magazine of Natural History 7 (15): 584 - 591.","Krumbiegel, V. I. 1940. Die Saugetiere der Sudamerika-Expeditionen Prof. Dr. Kriegs. 7. Pakas. Zoologische Anzeiger 132: 223 - 238.","ICZN. 1998. Opinion 1894 [of the International Commission on Zoological Nomenclature]. Regnum animale..., ed. 2 (M. J. Brisson, 1762): rejected for nomenclatural purposes, with the conservation of the mammalian generic names Philander (Marsupialia), Pteropus (Chiroptera), Glis, Cuniculus, and Hydrochoerus (Rodentia), Meles, Lutra and Hyaena (Carnivora), Tapirus (Perissodactyla), Tragulus and Giraffa (Artiodactyla). Bulletin of Zoological Nomenclature 55: 64 - 71."]}