A new earless species of Poyntonophrynus (Anura, Bufonidae) from the Serra da Neve Inselberg, Namibe Province, Angola Author Ceriaco, Luis M. P. Author Marques, Mariana P. Author Bandeira, Suzana Author Agarwal, Ishan Author Stanley, Edward L. Author Bauer, Aaron M. Author Heinicke, Matthew P. Author Blackburn, David C. text ZooKeys 2018 780 109 136 http://dx.doi.org/10.3897/zookeys.780.25859 journal article http://dx.doi.org/10.3897/zookeys.780.25859 1313-2970-780-109 A088FFEB15D74E1DAECD09623AE57066 A088FFEB15D74E1DAECD09623AE57066 Poyntonophrynus pachnodes sp. n. Figs 4, 5, 6, 7, Table 3 Holotype. A female, UF 184184 (field number AMB 10208), collected on Serra da Neve (- 13.77704 S , 13.25905 E ; datum WGS 84; 1488 m a.s.l.), 18 November 2016, by Luis M. P. Ceriaco , Suzana Bandeira and Ishan Agarwal. Paratypes. A male UF 184183 (field number AMB 10191), a juvenile CAS 262729 (field number AMB 10207), a female CAS 262730 (field number AMB 10210), all with the same collecting data as the holotype, a male INBAC/AMB 10209, and male MUHNAC/MB04-000999 (field number AMB 10219) with same locality and collector data as the remaining type series but collected 19 November 2016. Diagnosis. Poyntonophrynus pachnodes sp. n. is a small-bodied bufonid that lacks tarsal folds, a character that distinguishes it from bufonids in Angola except Mertensophryne and Poyntonophrynus . It differs from all Mertensophryne in having inconspicuous parotoid glands, compared to pronounced parotoid glands that form a shelf in the scapular region of Mertensophryne , and in lacking reduction of the phalanges (Grandison, 1981). The newly described species differs from all other members of the genus Poyntonophrynus in lacking a tympanum and columella. Description of the holotype. Small (SVL 31.4 mm), robust, stout and gravid female, with moderately robust limbs (Figure 4-5; All measurements in Table 3); head triangular, wider than long; snout projecting slightly beyond upper jaw; rostral tip straight in dorsal, ventral and lateral views; eyes projecting laterally just beyond eyelids and approximately flush with margins of head in dorsal view; eye projecting about 50% above dorsal mar gin of head in lateral view; interorbital distance 1.1 times eye diameter; pupil large and ellipsoid in life and preservative; loreal region concave; naris small, triangular, directed dorsolaterally; canthus rostralis short, eye diameter 1.2 times eye-narial distance; eye diameter 1.6 times naris to rostral tip; internarial region flat with rounded lateral margins; interorbital distance approximately 1.7 times internarial distance; tympanum and middle ear structures (tympanum, columella) absent. Marginal and vomerine teeth absent. Figure 4. Ventral and dorsal view of the holotype (UF 184184) of Poyntonophrynus pachnodes sp. n. Figure 5. Ventral view of the right hand and left foot of the holotype (UF 184184) of Poyntonophrynus pachnodes sp. n. Table 3. Measurements of the type series of P. pachnodes sp. n. For measurement abbreviations see Materials and methods section. All measurements in mm.

Catalog number	UF 184184	UF 184183	CAS 262730	CAS 262729	MUHNAC/MB04-000999	INBAC/AMB 10209
HW
SVL
TL
IOD
HL
ED
IND
EN
FL
THL
SL
HAL
FLL
UEW
NS

Skin of venter with evenly scattered miniscule asperities; skin of gular region smooth; skin of limbs, dorsal and dorsolateral surface of head and body with scattered tubercles, being larger on dorsum; inconspicuous and flattened parotoid glands, elliptical, and weakly elevated, placed dorsolaterally and extending from posterior corner of mouth to level of axilla. Limbs and digits well-developed; digits of both manus and pes stout; tarsal fold not present; relative length of fingers: III> I> IV = II; finger tips not expanded, but with rounded tips; fingers with rounded, prominent double subarticular tubercles; two palmar tubercles distinct and widely separated from one another, one at ventromedial surface of first finger and other at proximal plantar surface, latter being about 4 times larger than first; webbing between manual digits absent; relative length of toes IV> III> II> I = V; toe tips slightly expanded; toes with prominent, single, and subarticular tubercles; webbing between toes vestigial, not reaching first joint of first phalanx; prominent and globular inner metatarsal tubercle, length 50% of first toe length; tarsal tubercle prominent and projecting, near medial edge and positioned at distal fourth of tarsus. Coloration. In life, dorsal ground color dark brown with coppery to brown mottling and dark brown blotches especially in anterior regions (Figure 6); whitish chevron extending between eyes (scapular patch), directed posteriorly (faded but visible in preservative); iris dark green with dark brown pupil (dark grey and pale gray in preservative, respectively); snout similar in color and pattern to dorsum; dorsal surface of forelimbs and fingers III and IV whitish with dark brown blotches; dorsal surface of fingers I and II white; posterior to head, distinctive white blotch at midbody; dorsum and lateral surface dark brown speckled with coppery markings, becoming faint towards venter; dorsal surface of hind limbs (thighs and crus) greyish white with distinctive dark brown markings; three dark blotches on thigh, crus and feet touching when legs flexed; base color of dorsal foot grayish, with scattered dark brown blotches, extending across toes; region surrounding cloaca cream colored. Lateralmost margin of upper jaw white interrupted by brown markings posterior and anterior to eye; throat immaculate white; ventral surface of forearm and arm whitish; ventral surface of hand and fingers white; venter unpigmented and whitish; anterior part of thighs and ventral crus cream colored; ventral side of legs and plantar surface of pes unpigmented and whitish in appearance. Figure 6. Photograph in life of Poyntonophrynus pachnodes sp. n., paratype UF 184183 (AMB 10191). Photo by Ishan Agarwal. Variation. The coloration of the remaining paratypes do not differ in any important details from those of the holotype. Measurements of the remaining type series in Table 3. Osteology. Based on CT scan of UF 184184 (holotype; gravid female; Figure 7). The skull is wider than long, incompletely ossified, and lacks ornamentation on the dermal roofing bones. The jaw joint is anterior to the otic region. The parotic plate is incompletely ossified but synostosed to the frontoparietal. The premaxillae lack teeth, and have both a robust pars dentalis and a robust alary process that is taller than wide and widely separated from the nasals. The maxillae lack teeth and are weakly concave (with apex directed labially). The quadratojugals are thin and elongate with a broad articulation with the maxillae and reaching anterior to the articulation of the maxilla and pterygoid. The pterygoids are slender, with a long medially curved anterior ramus with a broad articulation with the adjacent maxilla, a short posterior ramus approaching, but not articulating with, the cartilages of the jaw joint, and a short medial ramus approaching but not articulating with the prootic. Vomers are large and plate-like, lacking teeth. The neopalatine is a thin flat rod, not articulating with adjacent bones. The curved and triradiate septomaxillae are present at the anterior margin of the nasal capsule. The prominent sphenethmoid is coosified across the midline and visible in dorsal view be tween the nasals and frontoparietals. The parasphenoid narrows anteriorly and exhibits a small bifurcation at its rostral extent. The squamosals are greatly reduced with only the dorsalmost otic region and a small reduced zygomatic ramus present. The prootic is poorly ossified and has a poorly defined fenestra ovalis. Neither a bony operculum nor a columella is present. The posteromedial processes of the hyoid are ossified and slender, and expanded weakly at their articulation with the cartilaginous hyoid plate. Figure 7. High-resolution Computed Tomography reconstructions showing the skeleton of the holotype (UF 184184) of Poyntonophrynus pachnodes sp. n. in dorsal view (A), skull in dorsal (B), right lateral (C) and ventral (D) views, pelvis in right lateral view (E), pectoral girdle in ventral view (F), and right foot (G) and right hand (H) in ventral views. Selected skeletal elements are labeled based on discussion in the text. Panel A also highlights the eggs that are visible within the oviducts. Abbreviations: m - maxilla; np - neopalatine; pt - pterygoid; q - quadrate; sph - sphenethmoid; sq - squamosal; v - vomer. Scale bars in each row equal 2 mm. There are eight distinct, procoelous, non-imbricating presacral vertebrae that are not synostosed. The atlas lacks transverse processes and has widely separated cotyles. The sacrum is procoelous with laterally expanded transverse processes. No sesamoid is observed at the sacroiliac joint. The urostyle is long and thin with a weakly developed dorsal ridge. The pectoral girdle is fermisternal with widely spaced and slender coracoids. The clavicles are slender, nearly reaching one another. The scapulae are stout, directed laterally but strongly curving dorsally at their lateral extent. Neither an ossified sternum nor omosternum is present. The pelvic girdle comprises the ilium, pubis, and ischium, which are not synostosed to one another. The acetabulum is incompletely ossified. The shaft of the ilium is long and slender, lacking a dorsal crest, and with a small prominent dorsal protuberance. The synostosed ischia form a broad posteriorly directed plate. The radioulna is shorter than the humerus. The radiale and ulnare are large and subequal in size, though the other carpals are difficult to distinguish as they are incompletely ossified. The phalangeal formula for the manus is 2 -2-3- 3, and there is both a single ossified prepollex and a small palmar sesamoid. The tips of the terminal manual phalanges are weakly expanded into small knobs. The tibiofibula is slightly shorter than the femur. There are two small distal tarsals. The phalangeal formula for the pes is 2 -2-3-4- 3 and there is a single small ossified prehallux and a plantar sesamoid. The tips of the terminal pedal phalanges are weakly expanded as in the fingers. Distribution and ecology. The species is currently only known from the Serra da Neve Inselberg (Figs 2, 8) in northern Namibe Province. Specimens were found on moist soil under rocks and leaf-litter at dusk in a semi-open miombo forest area. Grandvaux-Barbosa (1970) considered that Serra da Neve is characterized by a "sparse Miombo," dominated by trees of the genera Julbernardia spp. and Brachystegia spp., and shrubs like Combretum spp. or Annona spp., which were observed at the site. The species was found sympatrically with the frogs Sclerophrys gutturalis and Tomopterna tuberculosa , the lizards Agama schacki , Trachylepis sulcata , Chondrodactylus pulitzerae , Hemidactylus benguellensis , Pachydactylus angolensis , and Heliobolus lugubris , and the snake Hemirhagerrhis viperina . Figure 8. Southern view of the base of the Serra da Neve Inselberg (A), and habitat of the type locality at about 1500 m a.s.l. (B). Photos by Luis Ceriaco . Etymology. The specific name pachnodes (Gr.) means "frosty" and is used as an adjective ( Brown 1954 ). This is a reference to both the cool climate at the higher elevation where this species occurs (1488 m) and that Serra da Neve (Port.) translates to "mountain of snow." We suggest "Serra da Neve Pygmy Toad" and "Sapo Pigmeu da Serra da Neve" as the English and Portuguese common names respectively.