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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.

Torres-Martínez, María M, Aya-Cuero, Carlos A, Noguera-Urbano, Elkin A, Passos, Fernando C, Ramírez-Chaves, Héctor E (2021): Coendou vestitus (Rodentia: Erethizontidae). Mammalian Species 53 (1003): 43-50, DOI: 10.1093/mspecies/seab005, URL: http://dx.doi.org/10.1093/mspecies/seab005

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

Don E. Wilson, Thomas E. Lacher, Jr, Russell A. Mittermeier (2016): Erethizontidae. In: Handbook of the Mammals of the World – Volume 6 Lagomorphs and Rodents I. Barcelona: Lynx Edicions: 372-397, ISBN: 978-84-941892-3-4, DOI: http://doi.org/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

Don E. Wilson, Thomas E. Lacher, Jr, Russell A. Mittermeier (2016): Erethizontidae. In: Handbook of the Mammals of the World – Volume 6 Lagomorphs and Rodents I. Barcelona: Lynx Edicions: 372-397, ISBN: 978-84-941892-3-4, DOI: http://doi.org/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.