ZMB 91791, adult female, collected 12 km N of Condé on rocky ground at the edge of forest, Cuanza-Sul Province, –10.743744°, 14.631923°, 1260 m a.s.l., 22 November 2017, by Pedro Vaz Pinto (Figs 5–7, 9–11).

ZMB 91790, adult male, same data as holo­type; BMNH 2021.7534, adult female, collected on a small plant 30 cm high in Congulo Forest, Cuanza-Sul Province, –10.745881°, 14.630576°, 520 m a.s.l., 21 ­October 2016, by Ninda L. Baptista, Pedro Vaz Pinto and William R. Branch; BMNH 2021.7535, FKH-1086, two adult females, collected in an accommodation yard at Quibala, in broken rocky foothills of a large granite massif, Cuanza-­Sul Province, –10.7399°, 14.979755°, 1303 m a.s.l., 21 September 2018, by Luke Verburgt.

Photographic record of lineage B (Fig. 10F). One male and one female (amplectant pair), in Gabela, Cuanza-Sul Province, approx. –10.85°, 14.38°, 1050 m a.s.l., November 2017, by Nguyen Thi Ngan Thanh. Not collected.

We restrict the type series of P. fernandae sp. nov. to specimens from the escarpment forest and adjacent areas (lineage B). The vouchers from Namba were listed and reported herein as additional referred material of P. cf. fernandae (lineage A): ZMB 91785, adult female, collected approximately 12 km W of Missão da Namba, Chinhundo, Cuanza-Sul Province, –11.914685°, 14.740552°, 1740 m a.s.l., 16 October 2020, by Pedro Vaz Pinto (Figs 12–14; Tables 5, 6). BMNH 2021.7536, FKH-0463, two adult females, same data as previous specimen; ZMB 91786, BMNH 2021.7537 two adult males, same collecting locality as previous specimen, collected on 03 November 2016, by Ninda L. Baptista, Pedro Vaz Pinto and William R. Branch.

Poyntonophrynus fernandae sp. nov. sensu lato are medium-sized pygmy toads with females larger than males, sexual dimorphism in skin texture (females rougher than males), and sexual dichromatism (females with complex dorsal patterns vs. males plain or partially bright yellow; females with speckles ventrally vs. males with nearly immaculate venter). No tarsal fold. Subarticular tubercles at the base of fingers and toes mostly single, and those at the joint between phalanges usually double. Tympanum not visible. Conspicuous parotoid glands, with curved outer edge. Usually two phalanges of toes III and V free of web, webbing between toes III and IV vestigial, not serrated. One or two enlarged palmar tubercles: one large, rounded to triangular-shaped very well developed outer metacarpal tubercle, and one smaller rounded inner metacarpal tubercle sometimes absent. Females with typical Poyntonophrynus arrangement of dorsal coloration pattern: pale single occipital, mid-dorsal, sacral and above arm insertion blotches.

Specimens of P. cf. fernandae (lineage A) differ from those of P. fernandae sp. nov. (lineage B) in shape of parotoid glands (elevated vs. flattened), conspicuousness of dorsal glandular warts (evident vs. discrete), dorsal coloration (dull vs. complex, colorful, in different shades of orange, green and black), ventral coloration (dark thick speckles vs. pale thin speckles), relative width of dark cross-bands on the limbs (similar width than pale ones and dull vs. much wider than pale ones, and orange on the outer surface of the limbs), inner metacarpal tubercle (conspicuous and always present vs. discrete and sometimes absent), overall sturdiness (sturdy vs. slender), relative limb size (short vs. long). Males differ from males of P. fernandae sp. nov. (lineage B) in dorsal coloration (dull coloration with similar pattern than females, but partially with yellow on flanks and thighs vs. plain bright yellow).

Poyntonophrynus fernandae sp. nov. sensu lato differs from all Poyntonophrynus species except P. pachnodes in lacking a tympanum and a columella. It differs from P. pachnodes in having a better-developed neopalatine. Males differ from P. pachnodes and P. nambensis sp. nov. in dorsal coloration (partially or plain bright yellow vs. complex dull patterns in P. pachnodes, and P. nambensis sp. nov.). It differs from P. nambensis sp. nov. in being smaller, less sturdy, and having relatively longer fore and hindlimbs, in ventral coloration (speckles vs. nearly immaculate). It differs from P. beiranus in parotoid glands conspicuousness (conspicuous, with clearly demarcated margins vs. inconspicuous). It differs from P. damaranus in ventral patterning (speckles vs. immaculate). It differs from P. dombensis in tympanum (not visible vs. conspicuous), and ventral patterning (speckles vs. immaculate). It differs from P. fenoulheti in parotoid glands width (thinner or around same width of eye diameter vs. wider than eye diameter). It differs from P. grandisonae in tympanum (not visible vs. conspicuous) and skin texture (rough vs. leathery). It differs from P. grindleyi in pale occipital and sacral patches (present vs. absent) and dorsal spines (small vs. large). It differs from P. hoeschi in ventral patterning (speckles vs. immaculate). It differs from P. jordani in shape of parotoid glands (kidney-shaped vs. a cluster of glands) and pale occipital patch (present vs. absent). It differs from P. lughensis in conspicuousness of parotoid glands (conspicuous vs. inconspicuous) and tympanum (not visible vs. conspicuous). It differs from P. kavangensis in foot webbing (non-serrated vs. serrated), dorsal patterning (absent vertebral line vs. present), and ventral patterning (speckles vs. immaculate). It differs from P. parkeri in ventral patterning (speckles vs. immaculate). It differs from P. vertebralis in dorsal patterning (vertebral line absent vs. present), and conspicuousness of parotoid glands (conspicuous vs. inconspicuous).

External morphology. Small (SVL 29.4 mm), slender, gravid female (Figs 9, 10, all measurements in Table 6). SVL approximately 2.9 times head width, 3.6 times head length, 2.4 times thigh length, 2.5 times tibiofibula length, and 4.2 times forearm length. Head rounded in dorsal view. Head length approximately 0.8 head width. Rostral tip rounded in dorsal and lateral views, truncate in ventral view. Eyes projecting laterally just beyond eyelids and not beyond margins of head in dorsal view. Eye not projecting above dorsal margin of head in lateral view. Interorbital distance approximately 1.2 times eye diameter, and approximately 1.7 times internarial distance. Eye diameter approximately 1.2 times eye-nostril distance, and 2.2 times naris to rostral tip. Tympanum not visible. Naris small, oval, directed dorsolaterally. Canthus rostralis sharp. Loreal region concave. Limbs and digits slender and well-developed. Tarsal fold absent. Digits of manus and pes elongated. Finger III length approximately 0.5 times hand length. Relative length of fingers: III > IV > II > I. Finger tips rounded, not expanded to discs. Fingers with prominent subarticular tubercles that are always double or bilobate, except under finger IV, which is single, on the right hand; and double or bilobate at the base of fingers I, II, and under finger III, and single at the base of fingers III, IV, and under finger IV, on the left hand. Plants of hands beset with supernumerary tubercles. Digits with subdigital tubercles, often double. Oval outer metacarpal tubercle well developed, larger and much more conspicuous and protuberant than inner metacarpal tubercle (Fig. 9C). Webbing between manual digits absent. Toe IV length approximately 0.6 times foot length. Relative length of toes IV > III = V > II > I. Toe tips rounded, not expanded to discs. Toes with prominent subarticular tubercles that are always single at toe bases, except for toe I on right foot, which is double, toe I on left foot, which is not visible, and those at the joint between phalanges being all double and elongated, except under finger V on right foot, which is single. Toes without a margin of web. Webbing between toes vestigial, webbing margin not serrated. Inner metatarsal tubercle prominent, nearly diamond shaped, pointy, approximately half the length of toe I, and less than twice longer than outer. Outer metatarsal tubercle prominent, oval. Mid-tarsal tubercle discrete, same size of the remaining tarsal tubercles, near medial edge and positioned at around the middle of the tarsus.

Figure 9. 

Holotype of Poyntonophrynus fernandae sp. nov. (ZMB 91791, female) in preservative. A Dorsal view. B Ventral view. C Palm of right hand. D Palm of right foot. Photos by F. Tillack. Short scale bars represent 1 mm (A, B), black scale bars represent 5 mm (C, D).

Figure 10. 

Pictures of live Poyntonophrynus fernandae sp. nov. A Female holotype (ZMB 91791). B, C Male paratype (ZMB 91790) with yellow color when collected, and with fading coloration after three days. D Female paratype (BMNH 2021.7534), darker individual collected in forest. E Female paratype (FKH-1086) collected on red gravel, typical of Cuanza-Sul Province ferralitic soils. F Pair in axillary amplexus from Gabela (not collected). Photos by L. Verburgt (E) and N.T. Ngan Thanh (F).

Dorsal skin rough, texture resembling sandpaper, with pale-tipped conical spines on dorsum, arms, legs, and lateral surface of snout. Skin of top of head smooth. Ventral and gular skin, and skin of ventral surface of limbs covered in minuscule spines, less pronounced and pointy than dorsal ones. Skin of venter and gular region granular. Parotoid glands elongated, flattened, conspicuous, with clearly discernible margins, kidney-shaped, with a curved outer margin, placed dorsolaterally and extending from behind the eye to slightly beyond forearm insertion. Poorly developed glandular warts around mid-dorsal pale patch.

Color. In life, dorsal coloration consists of a set of black and paler (mostly beige) blotches, forming a symmetrical pattern (Fig. 10A). From tip of head to sacrum, black blotches are either single or paired, located along the vertebral region. Single black blotches consist of one elongated blotch on top of snout; one continuous interorbital thin chevron directed posteriorly; and small occipital, mid-dorsal, and sacral blotches. Paired black blotches are located anteriorly and posteriorly to the triangular beige mid-dorsal blotch, and on the sides of the sacral beige blotch. From tip of head to sacrum, a series of beige blotches, which have brick orange portions on dorsal part of head and snout. Posterior to that, a triangular mid-dorsal beige blotch pointing forward, followed by an elongated beige blotch resembling a thick half vertebral stripe, outlined in coral green. Side of head cream with black and coral green blotches anterior and posterior to eye. Region below eye cream-colored. Black blotch from anterior corner of eyes to tip of snout. Region on top of lip beige with small black spots. Conspicuous cream roundish blotch above arm insertion. Parotoid glands mostly brick orange, with beige on the medial part, and black outer margins. Eyelid brick orange. Iris golden. Pupil black ellipsoid. Flanks with a reticulate of bright coral green, black, and cream markings. Dorsal surface of forelimbs with thick black cross-bands alternate with much thinner cream inter-spaces. Dorsal surface of fingers I and II cream, and of fingers III and IV cream with black cross-bands. Dorsal surface of hindlimbs (thighs and crus) with three thick black cross-bands, with thin paler inter-spaces that are cream on the inner surface, and bright orange on the outer surface of the limb (Fig. 10A). Black cross-bands on thigh, crus and feet touching when legs flexed. Ventral skin whitish and semi-transparent allowing to see interior of belly, full of eggs. Small grey speckles scattered on chest and belly, as well as on throat, where they are slightly smaller and less densely distributed. Ventral thighs and crus whitish. Ventral tarsus whitish with a grey bar. Plantar surface of pes pale grey, with white tubercles. Ventral surfaces of arm, forearm, hand and fingers whitish. Preserved specimen after 5 years in ethanol, with beige and orange areas now pale grey. Black and coral green areas turned black (Fig. 9). Soles of hands and feet became whitish, tubercles now white.

Osteology (Fig. 11). Skull wider than long, lacking ornamentation on dermal roofing bones. Jaw joint anterior to otic region. Parotic plate incompletely ossified but synostosed to frontoparietal. Premaxillae without teeth, with a robust pars dentalis and a robust alary process taller than wide and widely separated from nasals. Maxillae and mandibles curved and without teeth. Quadratojugals thin and elongate with broad articulation with maxillae. Pterygoids slender, with a long medially curved anterior ramus with broad articulation with adjacent maxilla, short posterior ramus approaching jaw joint, and short medial ramus approaching prootic. Vomers large and plate-like, without teeth, with short pointy mediolateral processes. Neopalatines synostosed (left) or approaching (right) anterior ramus of pterygoid, and synostosed (right) or approaching (left) edge of sphenethmoid. Septomaxillae present at anterior margin of nasal capsule. Prominent sphenethmoid co-ossified across midline and visible in dorsal view between nasals and frontoparietals. Parasphenoid narrows anteriorly and ends in a curve on rostral extent. Squamosals reduced, with dorsalmost otic ramus in contact with frontoparietal dorsally, very reduced zygomatic ramus present, and very poorly developed ventral ramus. Poorly ossified prootic. Columella absent. Posteromedial processes of hyoid ossified and slender.

Figure 11. 

CT-scan of Poyntonophrynus fernandae sp. nov. female holotype (ZMB 91791). A Skeleton in dorsal view. B Lateral, dorsal and ventral views of skull (left to right). C Pectoral girdle in ventral view. D Ventral views of right hand and right foot (left to right). Blue indicates the parotic plate. Scale bars represent 5 mm.

Eight distinct, procoelous, non-imbricating and not synostosed presacral vertebrae. Atlas without transverse processes, with widely separated cotyles. Sacrum procoelous with laterally expanded transverse processes, bearing expanded diapophyses. Urostyle long and thin, with weakly developed dorsal ridge on proximal half, and bicondylar articulation with sacrum. Firmisternal pectoral girdle, with widely spaced and slender coracoids. Clavicles slender, nearly reaching one another. Scapulae stout, directed laterally but strongly curving dorsally at their lateral extent. No visible ossified sternum or omosternum. Pelvic girdle comprising ilium, pubis, and ischium. Shaft of ilium long and slender, without dorsal crest. Radioulna shorter than humerus. Humerus bearing ventral crest on proximal half, and without medial and lateral crests. Phalangeal formula for manus 2–2–3–3. A single ossified prepollex. Tips of terminal manual phalanges weakly expanded into small knobs. Tibiofibula longer than femur. Phalangeal formula for pes 2–2–3–4–3. A single ossified prehallux. Tips of terminal pedal phalanges weakly expanded as in fingers.

Male (SVL 25.8 mm) smaller than females (28.5–29.9 mm). Male with nuptial pads consisting of a dense cover of minute dark asperities on upper and inner surfaces of finger I and to a lesser extent on the upper medial surface of finger II and inner metacarpal tubercle, absent on females. Male without spines on dorsum or venter, present on females. Male bright plain yellow, different from elaborate dorsal pattern on females.

Measurements of the type series are presented in Table 6, and variation of selected body ratios is summarized in Figure 4. Relative length of toes sometimes IV > III > V > II > I. Subarticular tubercles at the base of fingers: always double on finger I; always double or bilobate on finger II, except for BMNH 2021.7535 on right hand, which is single; always single on fingers III and IV. Subarticular tubercles at the joint between finger phalanges: always double on finger III; always single on finger IV, except for BMNH 2021.7535 on right hand, which is double. Subarticular tubercles at the base of toes: always double on toe I, except for BMNH 2021.7534 on left foot, which is single; always single on toe II, except for ZMB 91790 left foot, which is double; always single on toes III–V, except for BMNH 2021.7534 and BMNH 2021.7535, not visible. Subarticular tubercles at the joint between toe phalanges: when visible, always double on toe III; proximal always double on toe IV, except for ZMB 91790 left foot, which is single; distal always double on toe IV. When single, tubercles are usually rounded, and when double, usually thin and elongated, sometimes extremely pointy. Metatarsal tubercles oval, inner ranging from slightly longer to twice the length of outer. Toes without margin of web. Webbing between base of toes absent or vestigial, not serrated. All female paratypes have spines on top of head. Density of dorsal spines variable (BMNH 2021.7535 > BMNH 2021.7534 > ZMB 91791). Spine tips coloration varies between white, beige and brown. Male (ZMB 91790) with no spines on dorsum or venter, but a few spines on fore and hindlimbs. A few small glandular warts located latero-dorsally, behind parotoid glands.

Dorsal pale markings in life show some differences in shape, extent, and considerable variation in color intensity between type series and holotype. This seems to be at least to some degree substrate-related, and individual toads can apparently change coloration intensity (to darker, brighter, or paler shades of each tone). The shape of the pale dorsal blotches on the paratype females is similar to the typical Poyntonophrynus arrangement (e.g. Fig. 10E) (Poynton and Broadley 1988), with the mid-dorsal blotch being roundish or horse-shoe shaped, different from the holotype’s, which is triangular. Sections that are bright coral green on the holotype (outlining the black markings) have a paler shade of green in individuals found on reddish gravel ferralitic substrate (BMNH 2021.7535, FKH-1086, Fig. 10E), and very dark (nearly black) green in the specimen collected on a darker substrate in forest (BMNH 2021.7534, Fig. 10D). Pale marking above forearm insertion absent in one female (BMNH 2021.7534), which is generally darker than all others (Fig. 10D). Pale inter-spaces on fore and hindlimbs, that are cream on the holotype, are very bright orange on the specimen found in forest (BMNH 2021.7534, Fig. 10D), and pale beige on individuals found on reddish gravel ferralitic substrate (BMNH 2021.7535, FKH-1086, Fig. 10E). In BMNH 2021.7535 and FKH-1086 all the colors are present (orange, green and beige), but in less intense shades than in holotype, giving a duller aspect that resembles the more common Poyntonophrynus coloration. Parotoid glands on females vary from very bright orange (BMNH 2021.7534) to less intense orange (BMNH 2021.7535, FKH-1086), always with a black outer margin, and are lemon yellow on the male (ZMB 91790). The only collected male (ZMB 91790) was uniform lemon yellow in life, that changed to various degrees of yellow-greenish wash after a few days (Fig. 10B, C), and became grey when preserved, with parotoid glands slightly brownish. This plain bright yellow differed completely from all the females’ elaborate coloration pattern. Ventral patterning on females varied from immaculate in the male (ZMB 91790), to a sparse speckling of small grey spots placed especially in midline of pectoral region (BMNH 2021.7535, FKH-1086), and similar to holotype (BMNH 2021.7534). Soles of hands and feet whitish or pale grey, tubercles white. Some of the ventral patterning lost intensity or disappeared after preservation.

Male’s (ZMB 91790) humerus with medial and lateral distal crests (Fig. 7). Maxillae and mandibles not curved. Neopalatines not synostosed to sphenethmoid, and distant from anterior ramus of pterygoid. Parasphenoid narrowing anteriorly and ending in pointy edge. Ventral rami of squamosals minimally developed, even less than in holotype.

P. cf. fernandae (lineage A) (Figs 12–14; Tables 5, 6). Males (23.9–24.2 mm, n = 2) smaller than females (30.3–31.8 mm, n = 3). Fewer and less conspicuous ventral speckles in males. Parotoid glands on males more flattened and less conspicuous than on females. Males have the same dorsal coloration pattern than females, but have shades of yellow on the flanks (Fig. 13B, D) (different from P. fernandae sp. nov. (lineage B), where males are plain yellow). Measurements of the P. cf. fernandae series are presented in Table 6, and a compilation of selected body ratios in Figure 5. Relative length of fingers sometimes III > IV > II = I. Relative length of toes sometimes IV > III > V > II > I. Subarticular tubercles at the base of finger IV always single. Outer metacarpal tubercle varying between oval and near-triangular, inner metacarpal tubercle always very reduced, sometimes absent. Metatarsal tubercles oval, inner varying between being around the same size to almost two times longer than outer, sometimes being very pointy. Webbing between bases of toes ranging between vestigial to reaching up to the first phalange of toe IV. Conical spines present also on dorsal surface of snout and top of head, with tips being more often white, rarely brown. Parotoid glands elevated, dorsum with glandular warts, located mostly around pale mid-dorsal blotch and dorsolaterally (Fig. 13A). Males sometimes with spines only on fore and hindlimbs, and less conspicuous glandular warts. In life, pale dorsal blotches pale to dark greyish, on a dark brown and dark green ground color (Fig. 13A), distributed along the vertebral region and consisting of a thick conspicuous chevron shaped occipital blotch, extending between the eyes and directed posteriorly, a mid-dorsal horse-shoe shaped blotch, and a small rounded sacral blotch. Glandular warts dark brown. Grey occipital chevron-shaped blotch demarcating dorsal part of head and snout, which are dark brown and dark green. Sides of head grey with dark blotches in front, behind and below the eye. Parotoid glands dark brown. Iris green and golden. Flanks with alternate dark brown, dark green, black and grey thick vermiculation. Dorsal surface of limbs and distal digits grey with dark brown cross-bands with similar width than the pale grey inter-spaces. On females, chest and belly region adjacent to flanks heavily marked with thick black speckles, which sometimes reach throat and belly.

Figure 12. 

Poyntonophrynus cf. fernandae (lineage A) (ZMB 91785, female) in preservative. A Dorsal view. B Ventral view. C Palm of right hand. D Palm of left foot. Short scale bars represent 1 mm (A, B), black scale bars represent 5 mm (C, D).

Figure 13. 

Coloration of live Poyntonophrynus cf. fernandae (lineage A). A Female (ZMB 91785). B–D Male (ZMB 91786) with partial yellow coloration. E Female (BMNH 2021.7536). Photos by W.R. Branch (B–D).

Figure 14. 

CT-scan of Poyntonophrynus cf. fernandae (lineage A) (ZMB 91785, female). A Skeleton in dorsal view. B Lateral, dorsal and ventral views of skull (left to right). C Pectoral girdle in ventral view. D Ventral views of right hand and right foot (left to right). Blue indicates the parotic plate. Scale bars represent 5 mm.

All analyzed specimens (ZMB 91785 (Fig. 14) and BMNH 2021.7536, females; and ZMB 91786, male) had vomers with very elongated, thin, and pointy mediolateral processes, and not curved maxillae and mandibles. ZMB 91785 parasphenoid narrowing anteriorly and arrow shaped at rostral extent. Squamosals of ZMB 91785 reduced, with only dorsalmost otic region, and negligible zygomatic and ventral ramus, which are even more reduced in ZMB 91786. ZMB 91785 neopalatines with small ventral process, in contact with the edges of sphenethmoid, and approaching, but not in contact with, maxillae and anterior ramus of pterygoid distally. BMNH 2021.7536 neopalatines approaching (right) and synostosed (left) to sphenethmoid. ZMB 91786 with ventral process only on left neopalatine. Both ZMB 91786 and BMNH 2021.7536 had single ossified prepollices and prehalluxes, and ossified and slender posteromedial processes of hyoid. In ZMB 91785, prepollex and prehallux not visible due to the resolution of the microCT-scan, and there is also no visible ossification on posteromedial processes of hyoid. Male (ZMB 91786) with short medial and lateral distal humeral crests (Fig. 7).

All specimens were collected in the rainy season. Poyntonophrynus fernandae sp. nov. (lineage B) ZMB 91790 and ZMB 91791 (male and female) were found during the day, on November 22^nd 2017, on a site approximately 12 km northwest of the village of Condé, with the male displaying bright lemon yellow coloration. The female was found on the ground at the base of a hill with many large granite boulders, while the male was found higher up on moist ground with moss near a natural water seepage. These two individuals were kept alive for three days, upon which the coloration in the male faded to a darker yellow-greenish color (Fig. 10B, C), a case of dynamic sexual dichromatism. BMNH 2021.7535, FKH-1086 (two females) were collected shortly after a thunderstorm, near the town of Quibala, in degraded savanna habitat with large granitic boulders. The brightly colored female specimen BMNH 2021.7534 was also collected in November, found on the ground but at night, well inside a moist forest block after rains, and not near water. One amplectant pair ascribed to this species was observed during the day near the town of Gabela (Fig. 10F). The species was always found in close proximity to large granite boulders at elevations from 520 m to 1303 m a.s.l. approximately, but the surrounding habitat was diverse, including moist escarpment forest at lower elevation (Congulo), secondary coffee forest (plantation of exotic tree species – Grevillia robusta) near Condé and degraded Miombo savanna at Quibala (Fig. 8), both the latter at higher elevations. Amphibian species found in sympatry included Hyperolius cinnamomeoventris Bocage, 1866, H. parallelus Günther, 1858, H. platyceps (Boulenger, 1900), Kassina senegalensis (Duméril & Bibron, 1841), Leptopelis cf. cynammomeus (Bocage, 1893), L. cf. jordani Parker, 1936, Ptychadena anchietae (Bocage, 1868), Sclerophrys pusilla (Mertens, 1937).

Poyntonophrynus cf. fernandae (lineage A) from Chinhundo, in Namba region, differed in some respects from P. fernandae sp. nov. (lineage B). Two males were collected in November 2016, while active during the day and after heavy rains, showing partially yellow-greenish coloration (Fig. 13B–D), found in syntopy with P. nambensis sp. nov. On the same location, in October 2020 and also following rains, three gravid lineage A females were collected at night, and again in syntopy with P. nambensis sp. nov. All P. cf. fernandae (lineage A) were found on the steep slopes of the same large rock boulder, among moss and water seepages (Fig. 8). The surrounding habitat was montane grasslands with stunted Miombo savanna, and some remnant patches of Afromontane forest in nearby ravines.

P. fernandae sp. nov. sensu stricto. The species is known from the Angolan central escarpment zone in the region of Gabela, and extending eastwards at least to Quibala, in Cuanza-Sul province, at elevations of 520–1303 m a.s.l. (Fig. 1). It appears to be strongly associated with large granite boulders in moist habitats, and may be more widely distributed along the western escarpment and large rock outcrops in central Cuanza-Sul province. Some of the habitats where it has been recorded were highly threatened by deforestation for wood, agriculture, and encroachment by invasive species (Inga vera). From Chinhundo, Namba, at an elevation of 1730 m a.s.l., in southern Cuanza-Sul Province we recorded P. cf. fernandae (lineage A). This site lies on in the Angolan highlands, on the south-western foothills of Namba mountain chain (Fig. 1). In all other surveyed sites around Namba region we failed to find this toad, including additional places where we retrieved P. nambensis sp. nov. Currently the conservation status of P. fernandae sensu lato needs to remain Data Deficient (DD) as per IUCN Red List Guidelines (IUCN 2022).

The specific epithet fernandae (Pt.) is a tribute to Fernanda Lages, a researcher and professor of Genetics based in Lubango, Angola. Her continuous investment in capacity building over the last decades and dedication to various research projects and international collaborations gave opportunities and transformed the professional paths of several young Angolan biologists, and thus of research in Biology in the country. The name, built in the feminine singular genitive, also pays homage to women in science. We suggest “Fernanda’s pygmy toad” and “sapo pigmeu da Fernanda” as English and Portuguese common names, respectively.

ZMB 91787, adult female, collected in granite bedrock 200 m S of old farm paddock, in Fazenda Namba, Cuanza-Sul Province,–11.914167°, 14.820556°, 1840 m a.s.l., 03 November 2016, by Ninda L. Baptista, Pedro Vaz Pinto and William R. Branch (Figs 5–7, 15–17).

CHL 0326, BMNH 2021.7538, BMNH 2021.7539, 3 females, same data as holotype; BMNH 2021.7540, ZMB 91788, two males, same data as holotype; FKH-0378, FKH-0379, FKH-0380, FKH-0381, four males, collected at night on bedrock near forested stream, in Missão da Namba, Cuanza-Sul Province, –11.922078°, 14.835542°, 1740 m a.s.l., 11 February 2020, by Pedro Vaz Pinto and Javier Lóbon-Rovira; FKH-0377 unconfirmed sex, same data as FKH-0378–0381; FKH-0457, FKH-0458, FKH-0462, ZMB 91789, four females, collected approximately 12 km W of Missão da Namba, Chinhundo, Cuanza-Sul Province, –11.914685°, 14.740552°, 1730 m a.s.l, 16 October 2020, by Pedro Vaz Pinto; CHL 0472, eggs, same data as holotype.

A medium-sized pygmy toad with females larger and rougher than males. No tarsal fold. Subarticular tubercles at the base of fingers and toes usually single, remaining ones, located at the joint between phalanges, usually double. Tympanum varies between visible and not visible. Conspicuous protuberant parotoid glands, with curved outer edge. Very conspicuous dorsal glandular warts. Usually 1.75 phalanges of toes III and V free of web, webbing between toes III and IV vestigial, not serrated. Two enlarged well-developed palmar tubercles: large, rounded to triangular-shaped outer metacarpal tubercle, and smaller oval inner metacarpal tubercle. Typical Poyntonophrynus dorsal coloration: one pale occipital blotch, one mid-dorsal blotch, one sacral blotch, and one pale blotch over arm insertion. In most individuals, the occipital and mid-dorsal pale blotches are connected in the vertebral area, resembling an hourglass shape only observed in this species. Dorsal coloration identical in both sexes, generally dull, varying in shades of grey, beige and brown with some traces of brick orange and dark brown. Dark cross-bands on limbs around the same width as pale interspaces.

Poyntonophrynus nambensis sp. nov. differs from P. pachnodes and P. fernandae sp. nov. sensu lato in having a columella. It differs from P. pachnodes in having a better-developed neopalatine. Differs from P. fernandae sp. nov. sensu lato in ventral patterning (few speckles along midline of chest, and dark line in front of arm insertion vs. scattered speckles at least on the chest), and dorsal coloration (dorsal pale hourglass-shaped blotch almost always present vs. absent), and breeding male coloration (dulls, similar to females vs. partially or completely bright yellow). It differs from P. beiranus in parotoid glands conspicuousness (conspicuous, elevated, with clearly demarcated margins vs. inconspicuous), and dorsal patterning (vertebral line absent vs. present). It differs from P. damaranus in ventral patterning (few dark speckles along midline of chest, and dark line in front of arm insertion vs. immaculate). It differs from P. dombensis in tympanum size (when visible, between 0.5 and 0.6 times internarial distance, vs. conspicuous and around 0.7 times internarial distance), and ventral patterning (few speckles along midline of chest, and dark line in front of arm insertion vs. immaculate). It differs from P. fenoulheti in parotoid glands width (thinner or around same width of eye diameter vs. wider than eye diameter). It differs from P. grandisonae in tympanum size (when visible, between 0.5 and 0.6 times internarial distance, vs. same width or wider than internarial distance) and skin texture (rough vs. leathery). It differs from P. grindleyi in pale occipital and sacral patches (present vs. absent), dorsal spines (small vs. large), and ventral coloration (few dark speckles along midline of chest, and dark line in front of arm insertion vs. dark thick ventral marbling). It differs from P. hoeschi in ventral patterning (few speckles along midline of chest, and dark line in front of arm insertion vs. immaculate). It differs from P. jordani in shape of parotoid glands (kidney-shaped vs. a cluster of glands) and pale occipital patch (present vs. absent). It differs from P. lughensis in conspicuousness of parotoid glands (conspicuous vs. inconspicuous). It differs from P. kavangensis in foot webbing (non-serrated vs. serrated), dorsal patterning (absent vertebral line vs. present), and ventral patterning (few speckles along midline of chest, and dark line in front of arm insertion vs. immaculate). It differs from P. parkeri in development of parotoid glands (elevated vs. flattened). It differs from P. vertebralis in dorsal patterning (absent vertebral line vs. present), ventral patterning (few speckles along midline of chest, and dark line in front of arm insertion vs. distinct thick dark blotches), and conspicuousness of parotoid glands (conspicuous vs. inconspicuous).

External morphology. Small (SVL 31.9 mm), robust gravid female (Fig. 15, Fig. 16B, C, all measurements in Table 7). SVL approximately 2.9 times head width, 3.9 times head length, 2.8 times thigh length, 2.9 times tibiofibula length, and 4.8 times forearm length. Head rounded in dorsal view. Head length approximately 0.7 head width. Snout profile rounded, snout projecting slightly beyond upper jaw. Rostral tip rounded in dorsal, ventral and lateral views. Eyes projecting laterally just beyond eyelids and not beyond margins of head in dorsal view. Around a third of the eye projecting above dorsal margin of head in lateral view. Interorbital distance approximately 0.9 times eye diameter and 1.4 times internarial distance. Eye diameter approximately 0.9 times eye-nostril distance, and 2.7 times naris to rostral tip distance. Naris small, oval, directed dorsolaterally. Tympanum visible under dissecting microscope, approximately 0.3 times eye diameter. Canthus rostralis sharp. Loreal region concave. Limbs and digits robust and well-developed. Tarsal fold absent. Digits of manus and pes elongated. Finger III length approximately 0.5 times hand length. Relative length of fingers: III > I > IV = II. Finger tips rounded, not expanded to discs. Fingers with prominent subarticular tubercles that are single at the base of fingers II, III and IV, double at the base of finger I, and bilobate under fingers III and IV, on the right hand; and always double or bilobate, except at the base of fingers III and IV, which are single, on the left hand. Plants of hands beset with supernumerary tubercles. Digits with subdigital tubercles, often double. Inner and outer metacarpal tubercles very well developed (Fig. 15C), approximately oval in shape, the first being around twice as long as the latter. Webbing between manual digits absent. Toe IV length approximately 0.6 times foot length. Relative length of toes IV > III > V > II > I. Toe tips rounded, not expanded to discs. Prominent subarticular tubercles that are single at toe bases, the remaining being double. Double subdigital tubercles. Toes without a margin of web. Webbing between toes vestigial, only at the base of toes, webbing margin not serrated. Prominent oval inner and outer metatarsal tubercles, the first around twice longer than the latter. Inner metatarsal tubercle around half the length of toe I.

Figure 15. 

Holotype of Poyntonophrynus nambensis sp. nov. (ZMB 91787, female) in preservative. A Dorsal view. B Ventral view. C Palm of right hand. D Palm of right foot. Photos by F. Tillack. Short scale bars represent 1 mm (A, B), black scale bars represent 5 mm (C, D).

Figure 16. 

Pictures of live Poyntonophrynus nambensis sp. nov. A Female paratype (FKH-0458) with greyish coloration. B, C Female holotype (ZMB 91787) with dorsal and ventral coloration, respectively. D Pair in axillary amplexus [female (CHL0326), and male (BMNH 2021.7540) paratypes]. E Female paratype (BMNH 2021.7539) with brighter shades of brown and orange. F Eggs (CHL0472). Photos by W.R. Branch (B–F).

Dorsal skin rough, with brown-tipped conical spines on dorsum, arms, legs, dorsal and lateral surface of snout, top of head, and outer ring of tympanum. Ventral, gular skin, and ventral surface of limbs granular with no spines. Dorsum with very prominent rounded glandular warts, located mostly around pale mid-dorsal and sacral blotches, and dorsolaterally towards the flanks (Fig. 16B). Parotoid glands elongated, elevated, with clearly discernible margins, with a curved outer margin, placed dorsolaterally and extending from behind the eye to slightly beyond the forearm insertion. Both kidney-shaped, but the right one consisting of a single mass, and the left broken in the middle, consisting of a junction of two masses.

Color. In life, dorsal coloration consists of a set of pale beige blotches distributed along the vertebral region, forming a symmetrical pattern (Fig. 16B). These include a thick conspicuous chevron-shaped occipital blotch extending between eyes and directed posteriorly, connected to a mid-dorsal roundish blotch (the connection between these two blotches resembling an hourglass shape), and a small rounded sacral blotch. Rest of the dorsum with dark-, coppery- and orangey-brown mottling. Dark brown glandular warts located around the mid-dorsal blotch and towards the flanks. A thin dark brown interorbital bar, forming the anterior border of the beige occipital patch. Anterior to it, one continuous interorbital brown thin chevron directed posteriorly. Brown dorsum of head and snout. A dark brown line extending from anterior corner of eye to tip of snout. A pale beige blotch under both eyes. Conspicuous beige blotch above arm insertion. Parotoid glands brown. Eyelid brick orange. Iris olden. Pupil black ellipsoid. Flanks with dark brown (corresponding to glandular warts) and coppery markings on a pale grey background, becoming faint towards the venter. Dorsal surface of forelimbs and fingers III and IV pale beige, with dark brown cross-bands thinner than the beige inter-spaces. Dorsal surface of fingers I and II pale grey. Dorsal surface of hindlimbs (thighs and crus) beige with two dark brown cross-bands thinner than the beige inter-spaces. Dark cross-bands on thigh, crus and feet touching when legs flexed. Ventral skin almost immaculate white, with dark spots placed in midline of pectoral region, and a dark line curving down in front of insertion of arm (Fig. 16C). Plantar surface of manus and pes grey with white tubercles. Preserved specimen, after six years in ethanol, pale beige and grey blotches turned pale grey, and dark blotches (shades of brown) turned dark grey (Fig. 15). A large grey blotch on mid chest corresponding to area where formalin was injected for preservation, unrelated to natural pigmentation (Fig. 15B).

Osteology (Fig. 17). Skull wider than long, without ornamentation on dermal roofing bones. Jaw joint anterior to otic region. Parotic plate incompletely ossified but synostosed to frontoparietal. Premaxillae lacking teeth, with robust pars dentalis and robust alary process that is taller than wide, and widely separated from nasals. Maxillae and mandibles without teeth. Quadratojugals thin and elongate with broad articulation with maxillae. Pterygoids slender, with long medially curved anterior ramus with broad articulation with adjacent maxilla, short posterior ramus approaching jaw joint, and short medial ramus approaching prootic. Vomers large and plate-like, without teeth, with short and pointy mediolateral processes, and reduced anterior processes. Neopalatines thin flat rods, approaching but not in contact with edges of sphenethmoid, maxillae, and anterior ramus of pterygoid distally. Septomaxillae present at anterior margin of nasal capsule. Prominent sphenethmoid co-ossified across midline and visible in dorsal view between nasals and frontoparietals. Parasphenoid narrowing anteriorly and exhibiting a small bifurcation at its anterior extent. Squamosals reduced, with dorsalmost otic region, reduced zygomatic ramus present, and vestigial ventral ramus separate from main bone. Prootic poorly ossified. Columella well ossified, with pars interna plectri developed as an expanded knob on medial end of pars media plectri, which is antero-posteriorly compressed. Posteromedial processes of hyoid ossified and slender.

Figure 17. 

CT-scan of Poyntonophrynus nambensis sp. nov. holotype (ZMB 91787, female). A Skeleton in dorsal view. B Lateral, dorsal and ventral views of skull (left to right). C Pectoral girdle in ventral view. D Ventral views of right hand and right foot (left to right). Blue indicates the parotic plate, orange indicates the columella. Scale bars represent 5 mm.

Eight distinct, procoelous, non-imbricating, not synostosed presacral vertebrae. Atlas without transverse processes, with widely separated cotyles. Sacrum procoelous with laterally expanded transverse processes, bearing expanded diapophyses. Urostyle long and thin with weakly developed dorsal ridge on proximal half, a small pointy posteriorly directed protrusion on the right side of its right cotyle, and bicondylar articulation with sacrum. Pectoral girdle firmisternal, with widely spaced and slender coracoids. Clavicles slender, nearly reaching one another. Scapulae are stout, directed laterally but strongly curving dorsally at lateral extent. No visible ossified sternum or omosternum. Pelvic girdle comprising ilium, pubis, and ischium. Shaft of ilium long and slender, lacking a dorsal crest. Radioulna shorter than humerus. Humerus bearing a ventral crest on proximal half, and without medial and lateral crest. Phalangeal formula for manus 2–2–3–3. Ossified prepollex formed by two elements, a rounded proximal element, and a thinner, elongated and pointy distal element. Tips of terminal manual phalanges weakly expanded into small knobs. Tibiofibula around same length as femur. Phalangeal formula for pes 2–2–3–4–3. Tips of terminal pedal phalanges weakly expanded as in fingers. A single ossified prehallux.

Males (SVL 24.4–26.4 mm, n = 6) smaller than females (30.7–34.9 mm, n = 8). Males with a dense covering of minute dark asperities on upper and inner surfaces of finger I and to a lesser extent on the upper medial surface of finger II and inner metacarpal tubercle. Males with smoother skin than females, with few spines on snout, different from females, which have a rough dorsal skin. Parotoid glands on males less conspicuous and more flattened than on females.

Measurements of the type series are presented in Table 7, and variation in selected body ratios is summarized in Figure 4. Relative length of fingers sometimes III > IV > II = I. Relative length of toes sometimes IV > III > V > II > I, rarely IV > V > III > II > I. Variation on subarticular tubercles based on five paratypes (ZMB 91788–9 and BMNH 2021.7538–40). Subarticular tubercles at the base of fingers: always double or bilobate on fingers I and II; always single on finger III except for ZMB 91788 on right hand, which is bilobate, and for BMNH 2021.7540 and ZMB 91788 on both hands, which are double; always single on finger IV except for BMNH 2021.7540 on right hand, and for BMNH 2021.7539 on left hand, which are double. Subarticular tubercles at the joint between finger phalanges: always double on finger III; always single on finger IV, except for BMNH 2021.7538 on right hand, which is double, and for ZMB 91788 on right hand, which is absent. Subarticular tubercles at the base of toes: always double on toe I, except for BMNH 2021.7538 on both feet and for BMNH 2021.7540 on left foot, which are single, and for ZMB 91788 and BMNH 2021.7539 on left feet, which are not visible; always single on toe II, except for BMNH 2021.7539 which are double on both feet; always single on toes III–V. Subarticular tubercles at the joint between toe phalanges: always double on toe III; always double on toe IV except for ZMB 91790, distal one on right foot, which is single; always single on finger V, except for BMNH 2021.7540 on both feet and for ZMB 91788 on right foot which are absent, and ZMB 91789, which is double on both feet. Tympanum’s conspicuousness variable, being conspicuous (CHL 0326, FKH-0462), visible (BMNH 2021.7540, FKH-0377, FKH-0380, FKH-0457, ZMB 91788), and not visible in almost half of the collected specimens (BMNH 2021.7538–9, FKH-0378–9, FKH-0381, FKH-0458, ZMB 91789). Metatarsal tubercles oval, outer between half and two thirds the length of inner. Male paratypes usually with no spines, except on snout. Parotoid glands kidney-shaped, but very often in paratypes they look like the holotype’s left one, broken in the middle, consisting of two masses. Plants of hands and feet beset with supernumerary tubercles, less densely on feet than on hands. Spine tips vary in coloration between white, beige and brown.

Coloration varying in shades of grey, beige and brown with some traces of brick orange and dark brown (Fig. 16). Color intensity of pale dorsal blotches varied from pale grey (BMNH 2021.7538, BMNH 2021.7540, CHL 0326, FKH-0380, ZMB 91788) to very dark grey (FKH-0377–81, FKH-0457–8, FKH-0462, ZMB 91789). Ventral markings, anterior surface of forearm, posterior surface of tarsus, and palm of manus and pes with similar variation of color intensity. Except for BMNH 2021.7539, which has an immaculate venter, all individuals have dark spots placed along midline of pectoral region, and a dark line curving down in front of insertion of arm, that vary in size and form (usually more elongated antero-dorsally on males, and less elongate on females) (Fig. 16C). Dorsal hourglass-shaped blotch only absent in three out of 14 paratypes (CHL 0326, FKH-0378–9) (Fig. 16A). Soles of hands and feet from pale to very dark grey.

Humeri of male ZMB 91788 with a well-developed medial distal crest and a more discrete lateral distal crest (Fig. 7). Regular urostyle, with no pointy protrusion. Parasphenoid trifurcated anteriorly. Squamosals with reduced ventral rami. Distal element of prepollex very elongated, more than on the female holotype.

The type series was collected in the rainy season, during the night, while breeding in small rocky pools on granite boulders, where two pairs were found in axillary amplexus (Fig. 16D). No advertisement call was heard. All specimens were caught either on moist bedrock or steep boulder faces, near forest streams or natural seepages, while the surrounding habitat included also extensive montane grasslands (Fig. 8). Poyntonophrynus nambensis sp. nov. seem to be opportunistic feeders, inferring from the stomach content of one gravid female (ZMB 91789), which had ten swarming termites, one beetle (around 1.5 mm), ten ants (nine of which very recently ingested) near 3 mm in length and all morphologically identical, the remnants of the head of a jumping spider, and other non-identified arthropod body parts. One of the females from the type series deposited 545 eggs after being captured. Eggs were arranged in a single string, had black and white poles (Fig. 16F), and a diameter of around 1.5 mm. Amphibian species found in sympatry were Hyperolius parallelus (Günther, 1858), Hyperolius cinereus Monard, 1937, Phrynobatrachus cf. mababiensis FitzSimons, 1932, Ptychadena oxyrhynchus (Smith, 1849), Tomopterna tuberculosa (Boulenger, 1882), and P. cf. fernandae (lineage A).

This species is only known from the region of Serra da Namba, 1730–1840 m a.s.l., in the Angolan highlands. It has been recorded in various sites in relatively close proximity along the south and western slopes of the main mountain of Namba. So far, it has not been found in the escarpment zone or in other surveyed mountains in the highlands, suggesting that it may be endemic to Namba. The species appears to be locally common and its rupicolous habitat is probably not threatened, but until more research is conducted, we suggest it to be listed as Data Deficient (DD) as per IUCN Red List categories (IUCN 2022).

The specific epithet nambensis (Pt.) is a reference to the Namba mountains. This is the largest and more preserved relic of Afromontane forest and montane grasslands in Angola. However, it lacks official protection. We suggest “Namba pygmy toad” and “sapo pigmeu da Namba” as English and Portuguese common names respectively.