Coendou speratus Pontes, Gadelha, Melo, Sá, Loss, Junior, Costa & Leite, 2013, sp. nov.
Creators: name: Pontes, Antonio Rossano Mendes
name: Gadelha, José Ramon
name: Melo, Éverton R. A.
name: Sá, Fabrício Bezerra De
name: Loss, Ana Carolina
name: Junior, Vilacio Caldara
name: Costa, Leonora Pires
name: Leite, Yuri L. R.
Publisher: name: Zenodo
Year: 2013
Subjects: Biodiversity
Taxonomy
Animalia
Chordata
Mammalia
Rodentia
Erethizontidae
Coendou
Coendou speratus

description: 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 Tauá, Usina Trapiche, municipality of Sirinhaém, state of Pernambuco, Brazil, 8°33'46.13"S 35°10'9.09"W, elevation 85 m. Mata Tauá 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, Sirinhaém, 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 Conservação 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 Jaboatão dos Guararapes, approximately 50 km from Usina Trapiche; this animal was subsequently released in the Mata Xanguá (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 Várzea (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), Várzea, Recife, on 30 September 2009, by ICMBio employees (release # 41 of CETAS).
In addition, a specimen housed at the Museu Nacional, Rio de Janeiro, Brazil, also matches the description of C. speratus. This specimen apparently lacks a catalog number but it has field number SNP1 (for “Serviço 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 Viçosa, 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 São Francisco river), which encompasses the coastal region of the states of Alagoas, Pernambuco, Paraíba and Rio Grande do Norte. However, we did not find the new species during 5-year surveys 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 São Francisco River mouth; Oliver & Santos 1991; Caldara Jr. & Leite 2012). Apparently, the range of C. speratus does not overlap with that of C. nycthemera, an 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 Maranhão (Handley & Pine 1992; Voss & Angermann 1997; Oliveira et al. 2007).
Habitat. We sighted the new species in systematic census only at Usina Trapiche, where submontane 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 Tauá, when two individuals were sighted at 00:20 am, resting together on a 20 meter-high branch; the fourth sighting was also at Mata Tauá, at 11:55 pm, at a 10 meter-high branch; the fifth and the six were at Mata Xanguá, at 7:50 pm and 00:38 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 21st 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 Xanguá (470 ha), C. speratus had the highest density (36.8 ind/km²), while in the smallest fragment, Boca da Mata (94 ha), it had the lowest (5 ind/km²). Coendou speratus was found syntopically with C. prehensilis in all but one forest fragment, Mata Tauá (280 ha), and the biomass contribution of C. speratus was always lower than that of C. prehensilis (Table 3).
Conservation implications. The discovery of this new species of small porcupine in this particular zone of endemism highlights the importance of this hotspot for the conservation of the earth’s biota. It also points to the lack of information on its mammalian fauna, and the need for surveys in order to understand their distribution and status throughout the region (Costa et al. 2005).
Human impact is extremely high in the region where we found this new porcupine (Melo 2009), resulting in the regional extinction of large mammals, such as jaguar (Panthera onca), Brazilian tapir (Tapirus terrestris), giant anteater (Myrmecophaga tridactyla), and white-lipped peccary (Tayassu pecari) (Mendes Pontes 2009). The threat level to other medium-size mammals, like the blond capuchin monkey Sapajus flavius (Schreber) (Mendes Pontes et al. 2006; Alfaro et al. 2012), is currently still very high. Thus, long-term research programs and conservation measures must be implemented to assure the survival of the remaining species and protection of these forests.
description: Published as part of Pontes, Antonio Rossano Mendes, Gadelha, José Ramon, Melo, Éverton R. A., Sá, Fabrício 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