Coendou Lacépède, 1799

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), pp. 43-50 : 43-45

Coendou Lacépède, 1799

Coendou Lacépède, 1799:11 . Type species Coendou prehensilis (= Hystrix prehensilis Linnaeus, 1758 ), by monotypy.

Coendus E. Geoffroy St. -Hilaire, 1803:157. Incorrect subsequent spelling of Coendou Lacépède, 1799 .

Coandu G. Fischer, 1814:102 . Incorrect subsequent spelling of Coendou Lacépède, 1799 .

Sinethere F. Cuvier, 1823a:427 . Type species Hystrix prehensilis Linnaeus, 1758 , by original designation.

Sphiggure F. Cuvier, 1823a:427. Type species Sphiggure spinosa F. Cuvier, 1823a, by original designation.

Sinoetherus F. Cuvier, 1823b:256 . Incorrect subsequent spelling of Sinethere F. Cuvier, 1823a .

Sphiggurus F. Cuvier, 1823b:256 . Incorrect subsequent spelling of Sphiggure F. Cuvier, 1823b (but to be maintained due to prevailing subsequent usage; Voss 2011:6).

Coendu Lesson, 1827:290 . Incorrect subsequent spelling of Coendou Lacépède, 1799 .

Synethere Lesson, 1827:291 . Incorrect subsequent spelling of Sinethere F. Cuvier, 1823a .

Sinetheres J. B. Fischer, 1829:369 . Incorrect subsequent spelling of Sinethere F. Cuvier, 1823a .

Cercolabes Brandt, 1835:391 . Type species Hystrix prehensilis Linnaeus, 1758 (proposed as a replacement name for Coendou Lacépède, 1799 ).

Synoetheres Lund, 1839 a:227 . Incorrect subsequent spelling of Sinethere F. Cuvier, 1823a .

Sphingurus Tschudi, 1844:185 . Incorrect subsequent spelling of Sphiggure F. Cuvier, 1823b.

Echinoprocta Gray, 1865:321. Type species Erethizon (Echinoprocta) rufescens Gray, 1865 by monotypy.

Cryptosphingurus Miranda-Ribeiro, 1936:975. Type species Cryptosphingurus villosus (= Sphiggure villosa F. Cuvier, 1823b), by original designation.

CONTEXT AND CONTENT. Order Rodentia , suborder Hystricomorpha , superfamily Erethizontoidea , family Erethizontidae , subfamily Erethizontinae (Woods and Kilpatrick 2005; Voss 2015). The genus Coendou includes 14 species: the bicolor-spined porcupine ( C. bicolor ), the western Amazonian dwarf porcupine ( C. ichillus ), the Bahian hairy dwarf porcupine ( C. insidiosus ), the black-tailed porcupine ( C. melanurus ), the Mexican hairy porcupine ( C. mexicanus ), the eastern Amazonian dwarf porcupine ( C. nycthemera ), the Brazilian porcupine ( C. prehensilis ), the frosted porcupine ( C. pruinosus ), the Quichuan porcupine ( C. quichua ), the Roosmalens’s porcupine ( C. roosmalenorum ), the stump-tailed porcupine ( C. rufescens ), the Pernambuco dwarf porcupine ( C. speratus ), the Paraguayan hairy dwarf porcupine ( C. spinosus ), and the brown hairy dwarf porcupine ( C. vestitus — Voss 2015). C. prehensilis and C. quichua might represent species complexes based on molecular and morphological evidence ( Leite et al. 2011; Ramírez-Chaves et al. 2016). An identification key including all recognized species (except C. mexicanus ) has been recently provided by Voss (2015). The following key for the 14 currently recognized species of Coendou is adapted from Voss (2015) and Álvarez-Castañeda et al. (2017):

1. Size small (adult head–body length <400 mm; adult condyloincisive length of skull <70 mm); dorsal pelage includes defensive quills (with hard, sharp, microscopically barbed points) intermingled with bristle–quills (with long, flexible, wiry tips), except on head, rump, and base of tail (which have only defensive quills); tail always shorter than head–body length; frontal sinuses not inflated ...... 2

Size small or large; dorsal pelage does not include bristle–quills intermingled with defensive quills; tail longer or shorter than head–body length; frontal sinuses inflated or not ................ 5

2. Tail very short (about 50% of head–body length); dorsal pelage includes long blackish fur that partially or completely conceals defensive quills; bristle–quills bicolored (whitish or yellowish basally and dark brown or blackish distally); buccinator–masticatory foramen and foramen ovale confluent; mandible lacks a distinct coronoid process ............................................................ C. vestitus

Tail> 50% of head–body length (except, perhaps, in highmontane populations of C. pruinosus ); dorsal pelage with or without long fur; bristle–quills tricolored (with pale tips); buccinator–masticatory foramen and foramen ovale separate; mandible with distinct coronoid process ........ 3

3. Tail usually <70% of head–body length (never much longer); dorsal pelage always includes long fur that partially or completely conceals defensive quills; auditory bullae small, uninflated, not contacting paraoccipital processes .. ....................................................................... C. pruinosus

Tail usually> 75% of head–body length (never much shorter); dorsal pelage with or without long fur; auditory bullae large and inflated, contacting paraoccipital processes ... 4

4. Dorsal pelage appears completely spiny (long fur absent); at least some quills on head tricolored (with pale tips); terminal one-third to one-half of postcranial defensive quills black; ventral pelage includes clusters of soft spines mixed with sparse wool hairs .......................... C. ichillus

Dorsal pelage includes long fur that partially or completely conceals defensive quills; no pale-tipped quills on head; postcranial defensive quills mostly pale (only tips dark brown or blackish); ventral pelage consists of guard hairs and sparse wool hairs ......................... C. roosmalenorum

5. Tail very short (usually <50% of head–body length); dorsal body pelage without long fur; postcranial quills with hard, sharp, microscopically barbed points only on rump (remaining quills non-defensive, with soft points) ........................... C. rufescens

Tail consistently longer (> 50% of head–body length except in highland populations of C. quichua ); dorsal body pelage with or without long fur; all postcranial quills with hard, sharp, microscopically barbed points ................... 6

6. Large (adult head–body length> 400 mm; condylo-incisive length of adult skull> 77 mm; adult maxillary toothrow> 18 mm); dorsal body pelage with or without long fur; frontal sinuses inflated in most fully adult specimens ... 7

Usually smaller (adult head–body length usually <400 mm; condylo-incisive length of adult skull usually <77 mm; adult maxillary toothrow usually <18 mm); dorsal body pelage with or without long fur; frontal sinuses seldom inflated ........................................................................... 9 7. Dorsal body pelage appears completely spiny (long fur absent). Distributed in South America .............................. 8

Dorsal pelage appears covered with blackish to dark brown long fur, combined with yellowish spines with black tips. Frontal region of skull almost 30% wider than the rostrum width. Distributed from south Mexico to north Panama.. ................................................................... C. mexicanus

8. Quills uniformly bicolored and abruptly shorter on lower back and rump than on shoulders and upper back (resulting in a visually conspicuous cape or mantle of long quills); mesopterygoid fossa usually extends anteriorly between second upper molars ................. C. bicolor

Long tricolored quills (with white tips) intermingled with shorter bicolored quills over most of the dorsal surface, including rump (except in specimens from the southern Maracaibo Basin in Venezuela, which have uniformly bicolored quills); dorsal quills decrease gradually in length from upper back to rump, without abrupt transition; mesopterygoid fossa usually extends anteriorly between third molars .................................................... C. prehensilis

9. Dorsal pelage does not include long fur (appearing completely spiny) ............................................................... 10

Dorsal pelage usually includes long fur that partially or completely conceals the quills .................................... 12

10. Occurs in northern and western Colombia and western Ecuador; roof of external auditory meatus usually strongly keeled .............................................. C. quichua

Occurs in Brazil; roof of external auditory meatus smooth or weakly keeled ............................................................. 11

11. Occurs along south bank of the Amazon east of the Rio Madeira to Maranhão in Brazil (on the Atlantic coast); dorsal body quills mostly bicolored (whitish or yellowish basally with black tips), but at least a few lateral quills with pale (usually whitish but sometimes brownish) tips; ventral pelage consists of soft spines; dorsal surface of tail covered with blackish bristles ............. C. nycthemera

Occurs in extreme eastern Brazil (Pernambuco and Alagoas); most dorsal body quills tricolored, with long reddish tips; ventral pelage consists of soft fur; dorsal surface of tail covered with brownish or reddish bristles ...................................................................... C. speratus

12. Occurs north of Amazon and east of the Rio Negro– Casiquiare–Orinoco; tail about as long as head–body length; dorsal fur blackish streaked with long yellowish guard hairs; all quills bicolored; roof of external auditory meatus smooth .......................................... C. melanurus

Occurs in southeastern Brazil; tail substantially shorter than head–body length; dorsal fur dark or pale but not streaked with long yellowish guard hairs; quills all bicolored or some quills tricolored; roof of external auditory meatus smooth ............................................................................ 13

13. Long dorsal fur (when present) dark basally with pale hair tips (except in some northern specimens which have pale gray fur); at least some of the longer quills tricolored (with yellowish, orange, or reddish tips); vibrissae and caudal bristles bicolored (dark basally with yellowish or orange tips); adult maxillary toothrow> 15 mm ....... C. spinosus

Long dorsal fur uniformly pale (white, yellowish, or grayish), or pale basally with darker (brownish) hair tips; all vibrissae and caudal bristles dark (without pale tips); adult maxillary toothrow <15 mm ............... ................................................................... C. insidiosus