taxonID	type	description	language	source
C8A5014707175FF7A796491D5BD7CAC4.taxon	description	Figures 4, 5, 6, 7; Table 3	en	Conradie, Werner, Keates, Chad, Greenbaum, Eli, Lobón-Rovira, Javier, Tolley, Krystal A., Benito, Max, Vaz Pinto, Pedro, van Breda, Reuben V., Verburgt, Luke (2025): Systematics of African rough-scaled lizards, with description of two new species from eastern Angola (Squamata: Lacertidae: Ichnotropis Peters, 1854). Vertebrate Zoology 75: 627-672, DOI: 10.3897/vz.75.e167366
C8A5014707175FF7A796491D5BD7CAC4.taxon	description	General description. A medium-sized lacertid with a rounded snout and strongly striated head scales. Nostril pierced between three nasals; the supranasals in broad contact behind the rostral; single frontonasal as broad as long; paired prefrontal scales in broad contact medially; prefrontal mostly in contact with the anterior supraocular (separate in BE _ RMCA _ Vert. R. 40 [I. overlaeti paratype], BE _ RMCA _ Vert. R. 14641 [I. capensis nigrescens holotype] and NMZB-UM 16358), separated from the first supraciliary by a smaller scale (rarely in contact); two large supraoculars, which are separated from the supraciliaries by one row (or rarely two rows anteriorly) of small scales (7 – 9) and preceded by a cluster of 2 – 5 smaller scales; 1 – 3 smaller post-supraocular scales; paired frontoparietal scales in broad contact; two parietals separated by an interparietal; occipital scale not reaching much past parietals; two loreal scales present, the anterior one smaller than the posterior; posterior loreal is separated from the anterior supraocular by two smaller scales; subocular in contact with lip; 3 – 6 (mostly 4) supralabials anterior to the subocular and two posteriorly; 6 – 9 (mostly 6) infralabials; 5 (rarely 6) chin shields, with the anterior three (rarely four) in broad contact; 3 – 4 (mostly 4) supraciliaries; 29 – 40 midbody scale rows; 8 – 10 longitudinal rows of enlarged ventral plates; 22 – 31 transverse ventral scale rows; 17 – 24 subdigital lamellae under the 4 th toe; 10 – 14 femoral pores per thigh. Size: Adult specimens varied from 42.2 – 75.0 mm (mean: 63.2 mm) SVL and 85.0 – 156 mm (mean: 109.7 mm) TAIL. Largest female: 71 mm SVL (FMNH 74288 – Serra do Moco, Angola); largest male: 75 mm SVL (NMZB-UM 16358 – Chitau, Angola). Colouration (Fig. 4): The dorsal side of the head, body and tail varies from brown to coppery red, sometimes with dark brown to black paired blotches. The flanks are dark brown to black, typically with two pale dorsolateral stripes. The upper stripe, usually two scales wide, originates behind the eye and extends onto the tail. The lower stripe begins anteriorly at the supralabials, tracing posteriorly through the ear, over the arm, and to the groin, though it may not be distinctly defined at midbody. Between these two stripes lies a broad band of coppery brown to black scales, interspersed with scattered black markings. Beneath the lower pale stripe there are scattered brown to black markings, sometimes accompanied by orange spots or blotches that extend onto the venter. During the breeding season, males exhibit more prominent orange flanks (extending onto the lower side of the tail), while the white stripes and lateral sides of the head become vividly yellow anteriorly. Diagnostic narrowly-spaced yellow or orange spots above the arm extend backwards for about a third of the body in both sexes. Dorsal tail with scattered white specks and black bars. The venter is typically plain white, although some individuals may have a grey venter or scattered fine grey to black specks.	en	Conradie, Werner, Keates, Chad, Greenbaum, Eli, Lobón-Rovira, Javier, Tolley, Krystal A., Benito, Max, Vaz Pinto, Pedro, van Breda, Reuben V., Verburgt, Luke (2025): Systematics of African rough-scaled lizards, with description of two new species from eastern Angola (Squamata: Lacertidae: Ichnotropis Peters, 1854). Vertebrate Zoology 75: 627-672, DOI: 10.3897/vz.75.e167366
C8A5014707175FF7A796491D5BD7CAC4.taxon	distribution	Distribution. Ichnotropis bivittata is known from Angola’s central plateau, with its range extending northward into western DRC, the Republic of the Congo, and southeastern Gabon (Fig. 3). Laurent (1964) reported both I. overlaeti de Witte & Laurent, 1942 and I. bivittata occurring sympatrically at Alto Cuilo, Lunda-Sul Province, Angola. During a recent field expedition to Alto Cuilo, the presence of I. bivittata was confirmed, and re-examination of historical DM (Dundo Museum) material attributed to I. overlaeti revealed it to be rather assignable to I. capensis sensu lato, based on a narrower and sharper head profile, the prefrontal in contact with the anterior supraocular and absence of any dorsolateral spots. In the present study, we also document the occurrence of I. aff. capensis — herein described as a new species — from Mona Quimbundo, approximately 62 km east of Alto Cuilo. These findings indicate that three distinct Ichnotropis species occur in the Miombo woodland of northeastern Angola. Historical records of I. bivittata from eastern Angola (Manaças 1963) require re-evaluation, as they may be referable to the I. capensis group or possibly to I. tanganicana.	en	Conradie, Werner, Keates, Chad, Greenbaum, Eli, Lobón-Rovira, Javier, Tolley, Krystal A., Benito, Max, Vaz Pinto, Pedro, van Breda, Reuben V., Verburgt, Luke (2025): Systematics of African rough-scaled lizards, with description of two new species from eastern Angola (Squamata: Lacertidae: Ichnotropis Peters, 1854). Vertebrate Zoology 75: 627-672, DOI: 10.3897/vz.75.e167366
B08CEE791EB95AD8B7EB0ABAE6E2C34D.taxon	description	Figures 20, 21, 22, 23; Table 6	en	Conradie, Werner, Keates, Chad, Greenbaum, Eli, Lobón-Rovira, Javier, Tolley, Krystal A., Benito, Max, Vaz Pinto, Pedro, van Breda, Reuben V., Verburgt, Luke (2025): Systematics of African rough-scaled lizards, with description of two new species from eastern Angola (Squamata: Lacertidae: Ichnotropis Peters, 1854). Vertebrate Zoology 75: 627-672, DOI: 10.3897/vz.75.e167366
B08CEE791EB95AD8B7EB0ABAE6E2C34D.taxon	description	General description. A medium-sized lacertid with a narrow and depressed snout. Head scalation strongly striated. Nostril pierced between three nasals; the supranasals are in broad contact behind the rostral; single frontonasal, as broad as long; paired prefrontal scales in broad contact medially; prefrontal separated from the anterior supraocular (only in contact in 15 out of 245 specimens examined) and separated from supraciliaries by a smaller scale; two large supraoculars, which are separated from the supraciliaries by one (very rarely two) row of small scales (4 – 9) and preceded by a cluster of 3 – 10 smaller scales; two loreal scales present, which are separated from the anterior supraocular by 2 – 3 scales; 1 – 2 post-supraoculars; subocular in contact with the lip; 3 – 6 (mostly four) supralabials in front of the subocular; 5 – 8 (mostly six) infralabials; five chin shields, with the anterior three in broad contact; 3 – 5 (mostly four) supraciliaries; 25 – 42 (average: 36.7) midbody scale rows; 8 – 10 (average: 8.8) longitudinal rows of enlarged ventral plates; 20 – 31 (average: 25.8) transverse ventral scale rows; 16 – 26 (average: 21.6) subdigital lamellae under the 4 th toe; 6 – 15 femoral pores per thigh. Size: Adult specimens varied from 40.0 – 67.8 mm (mean: 54.6 mm) SVL and 69.5 – 149.0 mm (mean: 110.4 mm) TAIL. Largest female: 65 mm SVL (NMZB-UM 9228 – Umtali, Zimbabwe); largest male: 67.8 mm SVL (BE _ RMCA _ Vert. R. 7785 – Dilolo, DRC). Colouration (Fig. 20): In males, the flanks feature a striking, broad black longitudinal band that originates at the tip of the snout, passes through the eye, and extends posteriorly well beyond the hind limbs, gradually fading towards the tip of the tail. This black band is bordered by two distinct white stripes: The upper stripe begins just behind the eye, while the lower stripe originates at the rostral plate, crosses the tympanum, and runs parallel to the black band along the length of the body. Below the lower white stripe, a vivid reddish-orange stripe is especially prominent on the anterior flanks. A secondary short black line also originates at the snout, runs across the supralabials along the side of the head, and terminates anterior to the insertion of the forelimbs. The main black band on the flank is often scattered with small white spots, particularly towards the posterior end of the body. In breeding males, the white stripes on the head and neck, as well as the gular region, become infused with a bright yellow hue. The dorsal surface is a rich reddish-brown, adorned with scattered dark brown speckling. Females exhibit a more subdued colouration, with an overall grey-brown tone that is lighter on the ventral side. A single, less pronounced dark black stripe originates at the snout, passes through the eye, and continues along the flanks, gradually fading towards the tail. Juveniles and subadults are often grey in colouration with a white dorsolateral stripe. The venter is mostly white, but some specimens exhibit grey colouration with scattered black specks.	en	Conradie, Werner, Keates, Chad, Greenbaum, Eli, Lobón-Rovira, Javier, Tolley, Krystal A., Benito, Max, Vaz Pinto, Pedro, van Breda, Reuben V., Verburgt, Luke (2025): Systematics of African rough-scaled lizards, with description of two new species from eastern Angola (Squamata: Lacertidae: Ichnotropis Peters, 1854). Vertebrate Zoology 75: 627-672, DOI: 10.3897/vz.75.e167366
B08CEE791EB95AD8B7EB0ABAE6E2C34D.taxon	distribution	Distribution. Widespread, occurring across several countries in southern Africa, including Angola, Namibia, Botswana, Zambia, Zimbabwe, Mozambique, and parts of South Africa and Malawi (Fig. 19). Historical records from north-eastern Angola and DRC assigned to I. capensis or I. overlaeti need to be re-evaluated in light of this study and might be assignable to either I. tanganicana, I. bivittata or I. longicorpa sp. nov. (see new species description below).	en	Conradie, Werner, Keates, Chad, Greenbaum, Eli, Lobón-Rovira, Javier, Tolley, Krystal A., Benito, Max, Vaz Pinto, Pedro, van Breda, Reuben V., Verburgt, Luke (2025): Systematics of African rough-scaled lizards, with description of two new species from eastern Angola (Squamata: Lacertidae: Ichnotropis Peters, 1854). Vertebrate Zoology 75: 627-672, DOI: 10.3897/vz.75.e167366
284FB215FB1A56028F821F613482C363.taxon	description	Figure 8; Table 3	en	Conradie, Werner, Keates, Chad, Greenbaum, Eli, Lobón-Rovira, Javier, Tolley, Krystal A., Benito, Max, Vaz Pinto, Pedro, van Breda, Reuben V., Verburgt, Luke (2025): Systematics of African rough-scaled lizards, with description of two new species from eastern Angola (Squamata: Lacertidae: Ichnotropis Peters, 1854). Vertebrate Zoology 75: 627-672, DOI: 10.3897/vz.75.e167366
284FB215FB1A56028F821F613482C363.taxon	description	General description. A medium-sized lacertid with a robust, rounded snout. Head scalation moderately striated. Nostril pierced between three nasals; the supranasals are in broad contact behind the rostral; single frontonasal, as broad as long; paired prefrontal scales in broad contact medially; prefrontal not in contact with anterior supraocular and separated from the supraciliaries by a smaller scale; two large supraoculars, which are separated from the supraciliaries by one row of small scales (6 – 8) and preceded by a cluster of 2 – 3 (3 median) smaller scales; one post-supraocular scale; two loreal scales present, which are separated from the anterior supraocular by two scales (except in the holotype, where the anterior loreal is divided to form a supraloreal on both sides and on the left side of BE _ RMCA _ Vert. R. 3657); subocular in contact with lip; 4 – 5 (mostly 4) supralabials in front of subocular; 6 – 7 (mostly 6) infralabials; five chin shields, with the anterior 2 – 3 in broad contact (in the holotype only the first two chin shields are in contact, while in BE _ RMCA _ Vert. R. 3657 the third chin shield is in narrow contact anteriorly); 3 – 5 (mostly 4) supraciliaries; 34 – 35 midbody scale rows; 8 – 10 longitudinal rows of enlarged ventral plates; 24 – 25 transverse ventral scale rows; 18 – 20 subdigital lamellae under 4 th toe; 8 – 9 femoral pores per thigh. Size: Adult specimens varied from 53.8 – 58.0 mm (median: 55.0 mm) SVL and 77 mm TAIL (all specimens’ tails missing or truncated; this measurement is based on Schmidt 1919). Largest female: 58 mm SVL (AMNH 10674 – holotype); largest male: 55.0 mm SVL (BE _ RMCA _ Vert. R. 3656 – Adra, DRC). Colouration (based on preserved specimens; Fig. 8): Dorsal surface uniformly greyish brown, with scattered darker brown to black scales. A distinct lateral white stripe originates at the subocular region, bordered both dorsally and ventrally by narrow black lines; this stripe extends over the forelimbs but does not reach the hind limbs in females (AMNH 10674 and BE _ RMCA _ Vert. R. 3657), but reaches the hind limbs in the male (BE _ RMCA _ Vert. R. 3656). A second faint dorsolateral line is present above the lower white stripe, and only extends to just posterior of the forearms in females, while in the male this stripe is more prominent and extends to just above the hind limbs. Between these lines are a series of transverse black spots in the females, each spanning 2 – 3 scales in width and approximately half a scale in length, located at the tips of the scales. In the male the space between the two white stripes forms a prominent black band with scattered black scales. This band extends onto the temporal and snout area. Dorsally, two similar series of transverse black markings flank the vertebral region, extending laterally to the dorsolateral stripe. Ventral scales and chin shields are white with subtle grey margins. The two outermost ventral rows are punctuated with small brown dots in females, but form a continuous narrow black band between the limbs in the male and extends onto the supralabials. Supralabials and infralabials are irregularly mottled with light and dark pigmentation. Limbs greyish brown dorsally, transitioning to a lighter tone on the ventral surfaces.	en	Conradie, Werner, Keates, Chad, Greenbaum, Eli, Lobón-Rovira, Javier, Tolley, Krystal A., Benito, Max, Vaz Pinto, Pedro, van Breda, Reuben V., Verburgt, Luke (2025): Systematics of African rough-scaled lizards, with description of two new species from eastern Angola (Squamata: Lacertidae: Ichnotropis Peters, 1854). Vertebrate Zoology 75: 627-672, DOI: 10.3897/vz.75.e167366
284FB215FB1A56028F821F613482C363.taxon	distribution	Distribution. Only known from northeastern DRC in the vicinity of Aba (Fig. 3). Given this locality’s proximity to the border with South Sudan (<10 km), it is likely to occur in the latter country.	en	Conradie, Werner, Keates, Chad, Greenbaum, Eli, Lobón-Rovira, Javier, Tolley, Krystal A., Benito, Max, Vaz Pinto, Pedro, van Breda, Reuben V., Verburgt, Luke (2025): Systematics of African rough-scaled lizards, with description of two new species from eastern Angola (Squamata: Lacertidae: Ichnotropis Peters, 1854). Vertebrate Zoology 75: 627-672, DOI: 10.3897/vz.75.e167366
F151507D3A5851ADB7FAAC88776348DD.taxon	description	Figures 15, 16; Table 4	en	Conradie, Werner, Keates, Chad, Greenbaum, Eli, Lobón-Rovira, Javier, Tolley, Krystal A., Benito, Max, Vaz Pinto, Pedro, van Breda, Reuben V., Verburgt, Luke (2025): Systematics of African rough-scaled lizards, with description of two new species from eastern Angola (Squamata: Lacertidae: Ichnotropis Peters, 1854). Vertebrate Zoology 75: 627-672, DOI: 10.3897/vz.75.e167366
F151507D3A5851ADB7FAAC88776348DD.taxon	description	General description. A large, robust lacertid with a pointed snout. Head scalation weakly striated. Nostril pierced between three nasals; the supranasals are in broad contact behind the rostral; single frontonasal, as broad as long; paired prefrontal scales in broad contact medially; prefrontal separated from the anterior supraocular by a smaller scale (except on the right-side of TM 86237) and separated from supraciliaries by a smaller scale (except on the right-side of TM 38309); two large supraoculars, which are separated from the supraciliaries by one row of small scales (5 – 9) and preceded by a cluster of smaller scales (3 – 7); 2 – 3 post-supraocular scales; two loreal scales present, which are separated from the anterior supraocular by two scales; subocular in contact with lip; 4 – 5 (mostly five) supralabials in front of subocular; 5 – 7 (mostly six) infralabials; five chin shields, with the anterior three in broad contact; 4 – 5 (mostly five) supraciliaries; 44 – 47 (average: 45.6) midbody scale rows; 10 longitudinal rows of enlarged ventral plates; 27 – 31 (average: 28.3) transverse ventral scale rows; 20 – 26 subdigital lamellae under the 4 th toe; 8 – 14 femoral pores per thigh. Size: Adult specimens varied from 57.2 – 77.9 mm (mean: 65.6 mm) SVL and 103.4 – 148.0 mm (mean: 124.9 mm) TAIL. Largest female: 77.9 mm SVL (RE 211206 D 1 / NMNW R 12212 – Khaudum, Namibia); largest male: 70 mm SVL (USNM 163989 – 40 km W of Mohembo, Botswana). Colouration (in preservative; Fig. 16): Above pale grey-brown, with darker stippling and a few scattered dark black spots on the body and tail. A poorly defined dark brown dorsolateral band extends from the neck to the groin, where it breaks up into a line of lateral spots on the tail. Sides of the head and lower flanks white. Venter white. In juveniles or subadults (Fig. 15), the dark brown lateral band is replaced by a mustard-coloured band (Pietersen et al. 2017).	en	Conradie, Werner, Keates, Chad, Greenbaum, Eli, Lobón-Rovira, Javier, Tolley, Krystal A., Benito, Max, Vaz Pinto, Pedro, van Breda, Reuben V., Verburgt, Luke (2025): Systematics of African rough-scaled lizards, with description of two new species from eastern Angola (Squamata: Lacertidae: Ichnotropis Peters, 1854). Vertebrate Zoology 75: 627-672, DOI: 10.3897/vz.75.e167366
F151507D3A5851ADB7FAAC88776348DD.taxon	distribution	Distribution. Known from northeastern Namibia and adjacent Botswana, and from western Zambia (Fig. 13). The apparent gap in distribution between northeastern Namibia / Botswana and western Zambia likely reflects a lack of sampling, and the species’ range is believed to be more continuous.	en	Conradie, Werner, Keates, Chad, Greenbaum, Eli, Lobón-Rovira, Javier, Tolley, Krystal A., Benito, Max, Vaz Pinto, Pedro, van Breda, Reuben V., Verburgt, Luke (2025): Systematics of African rough-scaled lizards, with description of two new species from eastern Angola (Squamata: Lacertidae: Ichnotropis Peters, 1854). Vertebrate Zoology 75: 627-672, DOI: 10.3897/vz.75.e167366
08667256A08C5902B4CF3B9BB2A2F97A.taxon	description	Figures 24, 25; Tables 6, 7	en	Conradie, Werner, Keates, Chad, Greenbaum, Eli, Lobón-Rovira, Javier, Tolley, Krystal A., Benito, Max, Vaz Pinto, Pedro, van Breda, Reuben V., Verburgt, Luke (2025): Systematics of African rough-scaled lizards, with description of two new species from eastern Angola (Squamata: Lacertidae: Ichnotropis Peters, 1854). Vertebrate Zoology 75: 627-672, DOI: 10.3897/vz.75.e167366
08667256A08C5902B4CF3B9BB2A2F97A.taxon	etymology	Etymology. The species name longicorpa is the feminine form of the Latin adjective longicorpus, derived from longus (long) and corpus (body), referring to this species’ elongate body.	en	Conradie, Werner, Keates, Chad, Greenbaum, Eli, Lobón-Rovira, Javier, Tolley, Krystal A., Benito, Max, Vaz Pinto, Pedro, van Breda, Reuben V., Verburgt, Luke (2025): Systematics of African rough-scaled lizards, with description of two new species from eastern Angola (Squamata: Lacertidae: Ichnotropis Peters, 1854). Vertebrate Zoology 75: 627-672, DOI: 10.3897/vz.75.e167366
08667256A08C5902B4CF3B9BB2A2F97A.taxon	diagnosis	Diagnosis. Assigned to Ichnotropis due to the absence of a well-defined collar, digits not serrated or fringed, subdigital lamellae keeled, and subocular bordering the lip. A slender Ichnotropis with a single frontonasal; subocular bordering the lip; a single anterior loreal; feebly developed head shield striations, prefrontals well separated from the anterior supraocular; and supraciliaries separated from the supraoculars by a series of smaller scales. The new species can be distinguished from other Ichnotropis species based on a combination of the following characteristics: Prefrontals well separated from the anterior supraocular (versus mostly in contact in I. bivittata, I. microlepidota and I. tanganicana); lower number (34 – 41) of midbody scales rows (44 – 47 in I. grandiceps and 43 – 48 in I. robusta sp. nov.); small, depressed head and pointed snout (versus large robust head and rounded snout in I. grandiceps and I. robusta sp. nov.); four supralabials anterior to the subocular (versus mostly five in I. grandiceps and I. robusta sp. nov.); distinctive occipital scale usually extending posteriorly well beyond the level of the parietals (versus large trapeziform occipital wedged between the parietals, not protruding past parietals in I. grandiceps and I. robusta sp. nov.). The new species resembles I. capensis sensu lato in its narrow, pointed snout, with the prefrontals well separated from the anterior subocular. It differs in that the new species exhibits black spots on the chin shields and gular scales (versus immaculate in I. capensis sensu lato) and the absence of a clear upper white dorsolateral stripe that separates the dark black lateral band from the dorsal brown vertebral band (versus present in most I. capensis sensu lato). In the phylogenetic analysis, the uncorrected p distances show that the new species differs by> 5.9 % for 16 S and> 12.3 % for ND 4 sequence divergence from other Ichnotropis species (Table 2).	en	Conradie, Werner, Keates, Chad, Greenbaum, Eli, Lobón-Rovira, Javier, Tolley, Krystal A., Benito, Max, Vaz Pinto, Pedro, van Breda, Reuben V., Verburgt, Luke (2025): Systematics of African rough-scaled lizards, with description of two new species from eastern Angola (Squamata: Lacertidae: Ichnotropis Peters, 1854). Vertebrate Zoology 75: 627-672, DOI: 10.3897/vz.75.e167366
08667256A08C5902B4CF3B9BB2A2F97A.taxon	description	Holotype description (Fig. 25). Adult male measuring 67.7 mm SVL and 160 mm TAIL (2.4 × SVL). Body moderately depressed; head distinctly depressed, almost twice as long as broad (HL 15.8 / HW 8.5 mm), its length equivalent to 23.3 % of SVL, expanded in the temporal region and very distinct from the neck. Adpressed hind limb just reaching the anterior edge of ear opening. The foot length is longer than the head length (FL 19.2 / HL 15.8 mm). Dorsal head shields very feebly striated and keeled; nostril pierced between three nasals, the supranasals in broad contact behind the rostral; frontonasal as long as broad (2.2 × 2.2 mm); prefrontals much longer than broad (2.7 × 1.5 mm), in broad contact medially, not reaching the anterior supraoculars (separated by a small keeled scale), in contact with the anterior and posterior loreal; frontal more than twice as long as its maximum width between the posterior tips of the prefrontals (4.6 × 2.1 mm), rounded anteriorly and strongly narrowed posteriorly; paired frontoparietals longer than broad (3.0 × 2.0 mm); parietals longer than broad (3.8 × 2.6 mm), extending posteriorly, widely separated by a large interparietal and occipital, the posterior margin extending past the posterior borders of the parietals; three keeled temporal scales bordering the parietal, the first one longest, followed by the second and third (smallest); two enlarged supraoculars, the anterior supraocular slightly longer than the posterior one and longer than its distance from the posterior loreal (2.4 mm vs. 1.5 mm), in contact with the posterior half of the frontal, separated from the posterior loreal by two smaller keeled scales; the anterior supraoculars are preceded by a cluster of five smaller keeled scales, the posterior supraocular is followed by three smaller keeled post-supraocular scales, the two supraoculars are separated from the supraciliaries by a single row of nine small keeled scales. Five supraciliaries, the first two much longer than the others and forming a long oblique suture. Lower nasal in contact with the rostral, first supralabial, and anterior loreal (narrow contact on left side); postnasal small, in contact with the other two nasals, anterior loreal, and frontonasal. Two loreals, the posterior one much larger and divided below; four supralabials anterior to the subocular, whose lower border on the lip is much shorter (3 ×) than the upper border; three supralabials posterior to subocular; temporal scales strongly keeled; a narrow tympanic shield on the upper anterior corner of the vertically elongate ear opening. Lower eyelid scaly with a median series (4 – 5) of vertically elongate scales. Six infralabials; five pairs of large chin shields, the first three pairs in median contact; gular scales imbricate; no collar. Dorsal scales rhombic, strongly keeled and imbricate; laterals smaller and feebly keeled, passing gradually into the smooth, rounded ventral plates, which are broader than long; 39 scales around the middle of the body; ventral plates in nine longitudinal and 30 transverse rows between the fore- and hind limbs; preanal scales irregular; scales on upper surfaces of limbs rhombic, strongly keeled, and imbricate; 12 / 10 femoral pores on each side; subdigital lamellae pluricarinate and spinulose, 22 under the 4 th toe; caudal scales strongly keeled above and below, except those just posterior to the vent, which are smooth.	en	Conradie, Werner, Keates, Chad, Greenbaum, Eli, Lobón-Rovira, Javier, Tolley, Krystal A., Benito, Max, Vaz Pinto, Pedro, van Breda, Reuben V., Verburgt, Luke (2025): Systematics of African rough-scaled lizards, with description of two new species from eastern Angola (Squamata: Lacertidae: Ichnotropis Peters, 1854). Vertebrate Zoology 75: 627-672, DOI: 10.3897/vz.75.e167366
08667256A08C5902B4CF3B9BB2A2F97A.taxon	distribution	Distribution. Only recorded from the headwaters of the Lungwebungu and Cuando Rivers in central Angola, northwards to the DRC border (Fig. 19). Some specimens from Mabwe River, Upemba National Park, DRC (IRSNB 7895, 7897, 7907 – 9, 78728), exhibit the same distinct dark brown to black gular markings and might be assigned to this species. If confirmed, this new species could be more widely distributed than currently thought.	en	Conradie, Werner, Keates, Chad, Greenbaum, Eli, Lobón-Rovira, Javier, Tolley, Krystal A., Benito, Max, Vaz Pinto, Pedro, van Breda, Reuben V., Verburgt, Luke (2025): Systematics of African rough-scaled lizards, with description of two new species from eastern Angola (Squamata: Lacertidae: Ichnotropis Peters, 1854). Vertebrate Zoology 75: 627-672, DOI: 10.3897/vz.75.e167366
22C20657A1BE521C8EACCEAEF5DF86DB.taxon	description	Figures 9, 10; Table 3	en	Conradie, Werner, Keates, Chad, Greenbaum, Eli, Lobón-Rovira, Javier, Tolley, Krystal A., Benito, Max, Vaz Pinto, Pedro, van Breda, Reuben V., Verburgt, Luke (2025): Systematics of African rough-scaled lizards, with description of two new species from eastern Angola (Squamata: Lacertidae: Ichnotropis Peters, 1854). Vertebrate Zoology 75: 627-672, DOI: 10.3897/vz.75.e167366
22C20657A1BE521C8EACCEAEF5DF86DB.taxon	description	General description. A medium-sized, robust lacertid with a rounded snout and strongly striated and keeled head scales. Nostril pierced between three nasals; the supranasals are in broad contact behind the rostral; single frontonasal, as broad as long; paired prefrontal scales in broad contact medially; prefrontal in contact with the anterior supraocular and either in contact or narrowly separated from supraciliaries by a smaller scale; two large supraoculars, preceded by a single scale (documented by Marx 1956 as a small supraocular); the anterior supraocular is in broad or narrow contact with the 1 st supraciliary anteriorly; the posterior part of the anterior supraocular and the posterior supraocular are separated from the supraciliaries by one row of small scales (6); one post-supraocular scale; two loreal scales present, which are separated from the anterior supraocular by one scale; subocular in contact with lip; four supralabials in front of subocular; 6 – 8 infralabials (mostly seven); five chin shields, with the anterior three pairs in broad contact; four supraciliaries (Marx 1956 recorded five, but he included the posterior loreal); 43 – 50 midbody scale rows; 8 – 10 longitudinal rows of enlarged ventral plates; 26 – 30 transverse ventral scale rows; 16 – 19 subdigital lamellae under the 4 th toe; 10 – 13 femoral pores per thigh. Size: Adult specimens varied from 48.7 – 52.0 mm (mean: 50.4 mm) SVL and 69.8 mm TAIL (only one specimen with intact tail). Largest female: 51 mm SVL (FMNH 74283); largest male: 52 mm SVL (FMNH 74285, 74286). Colouration (Fig. 9): The dorsal pattern features a light brown central band extending from just behind the head to the hind limbs. This band is bordered on each side by two broken rows of black blotches. Along the lateral sides of the body, two cream to yellow longitudinal stripes run from the level of the ear openings posteriorly to the hind limbs. Between these stripes lie a series of paired white ocelli, each bordered externally by black rings. Below the lower lateral stripe there is a continuous row of single white ocelli. The dorsal surface of the head is brown, mottled with black speckling across most scales. The mouth is bordered in black, which fades to white along the upper portion of the supralabials and the lower portion of the infralabials. The first row of chin shields is entirely black, while rows two through five are bicoloured — black medially and white laterally. The throat (gular region) is pale red-orange, interspersed with black scales and marked by two distinct bright yellow-orange spots located beneath the posterior ends of the lower jaws. The ventral surface is uniformly white	en	Conradie, Werner, Keates, Chad, Greenbaum, Eli, Lobón-Rovira, Javier, Tolley, Krystal A., Benito, Max, Vaz Pinto, Pedro, van Breda, Reuben V., Verburgt, Luke (2025): Systematics of African rough-scaled lizards, with description of two new species from eastern Angola (Squamata: Lacertidae: Ichnotropis Peters, 1854). Vertebrate Zoology 75: 627-672, DOI: 10.3897/vz.75.e167366
22C20657A1BE521C8EACCEAEF5DF86DB.taxon	distribution	Distribution. Currently only known from the slopes of Serra do Moco, in the central Angolan highlands (Fig. 3).	en	Conradie, Werner, Keates, Chad, Greenbaum, Eli, Lobón-Rovira, Javier, Tolley, Krystal A., Benito, Max, Vaz Pinto, Pedro, van Breda, Reuben V., Verburgt, Luke (2025): Systematics of African rough-scaled lizards, with description of two new species from eastern Angola (Squamata: Lacertidae: Ichnotropis Peters, 1854). Vertebrate Zoology 75: 627-672, DOI: 10.3897/vz.75.e167366
59681C2BC5B251A2A570E125D6B18DE3.taxon	description	Figures 17, 18; Tables 4, 5	en	Conradie, Werner, Keates, Chad, Greenbaum, Eli, Lobón-Rovira, Javier, Tolley, Krystal A., Benito, Max, Vaz Pinto, Pedro, van Breda, Reuben V., Verburgt, Luke (2025): Systematics of African rough-scaled lizards, with description of two new species from eastern Angola (Squamata: Lacertidae: Ichnotropis Peters, 1854). Vertebrate Zoology 75: 627-672, DOI: 10.3897/vz.75.e167366
59681C2BC5B251A2A570E125D6B18DE3.taxon	etymology	Etymology. The species name robusta is the feminine form of the Latin adjective robustus, meaning ‘ robust’ or ‘ sturdy’, in reference to the large, heavy-built adults of this species.	en	Conradie, Werner, Keates, Chad, Greenbaum, Eli, Lobón-Rovira, Javier, Tolley, Krystal A., Benito, Max, Vaz Pinto, Pedro, van Breda, Reuben V., Verburgt, Luke (2025): Systematics of African rough-scaled lizards, with description of two new species from eastern Angola (Squamata: Lacertidae: Ichnotropis Peters, 1854). Vertebrate Zoology 75: 627-672, DOI: 10.3897/vz.75.e167366
59681C2BC5B251A2A570E125D6B18DE3.taxon	diagnosis	Diagnosis. Assigned to Ichnotropis due to the absence of a well-defined collar, digits not serrated or fringed, subdigital lamellae keeled, and subocular bordering the lip. A large Ichnotropis with a single frontonasal; subocular bordering the lip; a single anterior loreal; feebly developed head shield striations; prefrontals well separated from the anterior supraocular; and supraciliaries separated from the supraoculars by a series of smaller scales. The new species can be distinguished from other Ichnotropis species based on a combination of the following characters: Prefrontals well separated from the anterior supraocular (versus mostly in contact in I. bivittata, I. microlepidota and I. tanganicana); high number of midbody scale rows (43 – 48 versus 25 – 42 in I. capensis sensu lato); large, robust head and rounded snout (versus small depressed head and pointed snout in I. capensis sensu lato); four (46 %) to five (50 %) supralabials anterior to the subocular (versus mostly four in I. capensis sensu lato); distinctive large trapeziform occipital wedged between the parietals, not protruding past parietals (versus occipital usually extending posteriorly, well beyond the level of the parietals in I. capensis sensu lato). The new species resembles I. grandiceps in its large size, robust, rounded head; prefrontals well separated from anterior subocular; high midbody scale rows (43 – 48 versus 44 – 47) and genetic similarity. Due to the lack of comparative adult material of I. grandiceps, no clear morphological and colouration differences could be observed between the two species. However, the two species exhibit clear differences in habitat preferences. All I. grandiceps material have either been found in drier Zambezian Baikiaea woodlands or Combretum-Vachellia bushveld (Broadley 1967 b; Haacke 1970; Pietersen et al. 2017) at lower elevations (less than 1000 m a. s. l.), while the new species is associated with the higher elevations (above 1300 m a. s. l.) of the Angolan Plateau, which consists of moister Angolan Miombo woodland. In the phylogenetic analysis, the uncorrected p distances show that the new species differs by> 6.7 % for 16 S and> 16.3 % for ND 4 sequence divergence from other Ichnotropis species (Table 2).	en	Conradie, Werner, Keates, Chad, Greenbaum, Eli, Lobón-Rovira, Javier, Tolley, Krystal A., Benito, Max, Vaz Pinto, Pedro, van Breda, Reuben V., Verburgt, Luke (2025): Systematics of African rough-scaled lizards, with description of two new species from eastern Angola (Squamata: Lacertidae: Ichnotropis Peters, 1854). Vertebrate Zoology 75: 627-672, DOI: 10.3897/vz.75.e167366
59681C2BC5B251A2A570E125D6B18DE3.taxon	description	Holotype description (Fig. 18). Adult male measuring 73.5 mm SVL and 96 mm TAIL (regenerated). Body moderately depressed; head not depressed, 1.7 times as long as broad (HL 18.9 / HW 11.1 mm), its length equivalent to 25.7 % of snout – vent length, expanded in the temporal region and very distinct from the neck. Adpressed hind limb reaches the anterior ear opening. The foot length is almost equal to the head length (FL 18.3 / HL 18.9 mm). Upper head shields very feebly striated and keeled; nostril pierced between three nasals, the supranasals in broad contact behind the rostral; frontonasal slightly broader than long (2.6 × 2.4 mm); prefrontals much longer than broad (4.3 × 1.7 mm), in broad contact medially, not reaching the anterior supraoculars (separated by a small keeled scale), and separated from the anterior loreal by a small keeled scale; frontal twice as long as its maximum width between the posterior tips of the prefrontals (6.2 × 3.0 mm), rounded anteriorly and strongly narrowed posteriorly; frontoparietals longer than broad; parietals longer than broad (5.4 × 2.8 mm), extending posteriorly, widely separated by a large interparietal and occipital, the latter small and its posterior margin level with the posterior borders of the parietals; an elongate keeled upper temporal shield borders the parietal; two supraoculars, the anterior supraocular longer than its distance from posterior loreal (2.3 mm vs. 1.8 mm), and in contact with posterior half of frontal; the second is smaller, separated from the supraciliaries by nine (right) / eight (left) small keeled scales (except the 4 th supraciliary on the left side, which is in narrow contact with the second supraocular); two post-supraocular scales; five supraciliaries, the first two much longer than the others and forming a long oblique suture. Lower nasal in contact with the rostral, first supralabial and anterior loreal; postnasal small, in contact with the other two nasals, frontonasal, and anterior loreal; two loreals, the posterior one much larger; four supralabials anterior to the subocular, whose lower border on the lip is much shorter (2.5 ×) than the upper; three supralabials posterior to the subocular; temporal scales strongly keeled; a narrow tympanic shield on the upper anterior edge of the vertically elongate ear opening; lower eyelid scaly with a median series of vertically elongate scales. Six infralabials; four (right) and five (left) large chin shields, the first two (right) and three (left) in median contact; gular scales imbricate; no collar. Dorsal scales rhomboid, strongly keeled and imbricate, lateral scales smaller and feebly keeled, passing gradually into the smooth, rounded ventral plates, which are broader than long; 44 scales around the middle of the body; ventral plates in 10 longitudinal and 29 transverse rows between fore- and hind limbs; preanal scales irregular; scales on upper surfaces of limbs rhomboid, strongly keeled, and imbricate; 12 femoral pores on each side; subdigital lamellae pluricarinate and spinulose, 21 under the 4 th toe; caudal scales strongly keeled above and below, except those just posterior to the vent, which are smooth.	en	Conradie, Werner, Keates, Chad, Greenbaum, Eli, Lobón-Rovira, Javier, Tolley, Krystal A., Benito, Max, Vaz Pinto, Pedro, van Breda, Reuben V., Verburgt, Luke (2025): Systematics of African rough-scaled lizards, with description of two new species from eastern Angola (Squamata: Lacertidae: Ichnotropis Peters, 1854). Vertebrate Zoology 75: 627-672, DOI: 10.3897/vz.75.e167366
59681C2BC5B251A2A570E125D6B18DE3.taxon	distribution	Distribution. Only recorded from the headwaters of the Okavango (Cuito and Cuanavale Rivers) and Cuando Rivers in central Angola (Fig. 14).	en	Conradie, Werner, Keates, Chad, Greenbaum, Eli, Lobón-Rovira, Javier, Tolley, Krystal A., Benito, Max, Vaz Pinto, Pedro, van Breda, Reuben V., Verburgt, Luke (2025): Systematics of African rough-scaled lizards, with description of two new species from eastern Angola (Squamata: Lacertidae: Ichnotropis Peters, 1854). Vertebrate Zoology 75: 627-672, DOI: 10.3897/vz.75.e167366
CBA5F90B078E5964B4D1B7B663DFDD50.taxon	description	Figures 11, 12, 13; Table 3	en	Conradie, Werner, Keates, Chad, Greenbaum, Eli, Lobón-Rovira, Javier, Tolley, Krystal A., Benito, Max, Vaz Pinto, Pedro, van Breda, Reuben V., Verburgt, Luke (2025): Systematics of African rough-scaled lizards, with description of two new species from eastern Angola (Squamata: Lacertidae: Ichnotropis Peters, 1854). Vertebrate Zoology 75: 627-672, DOI: 10.3897/vz.75.e167366
CBA5F90B078E5964B4D1B7B663DFDD50.taxon	description	General description. A medium-sized lacertid with a robust, rounded snout. Head scalation weakly to moderately striated. Nostril pierced between three nasals; the supranasals are in broad contact behind the rostral; single frontonasal, as broad as long; paired prefrontal scales in broad contact medially; prefrontal mostly in contact with the anterior supraocular (n = 29 in contact, seven not in contact; three in contact on one side only) and separated from supraciliaries by a smaller scale; two large supraoculars, which are either in direct contact (n = 15) or separated (n = 18) from the supraciliaries by a series of small scales; those that are not in contact are separated by one row of small scales (3 – 9) and preceded by a cluster of 1 – 6 (1.7 average) smaller scales; one post-supraocular scale; two loreal scales present, which are separated from the anterior supraocular by two scales; subocular in contact with lip; 3 – 5 (mostly 4) supralabials in front of subocular; 5 – 7 (mostly six) infralabials; five chin shields, with the anterior three in broad contact; 4 – 5 (mostly four) supraciliaries; 28 – 42 (average: 36.0) midbody scale rows; 8 – 10 (average: 8.4) longitudinal rows of enlarged ventral plates; 20 – 27 (average: 22.8) transverse ventral scale rows; 17 – 22 subdigital lamellae under the 4 th toe; 10 – 15 femoral pores per thigh. Size: Adult specimens varied from 41.0 – 60.0 mm (mean: 53.9 mm) SVL and 55.6 – 107.9 mm (mean: 81.7 mm) TAIL. Largest female: 60 mm SVL (NMZB-UM 24433 – Misuku Hills, Malawi); largest male: 56 mm SVL (NMZB-UM 24432 – Misuku Hills, Malawi). Colouration (Fig. 11): The top of the head and the anterior part of the body are coppery red, sometimes with three clearly defined black stripes on the nape. The anterior part of the dorsum is grey with scattered brown paired blotches with black edging, extending onto the tail. The flanks are dark brown to black, typically with interrupted white dorsolateral stripes. The upper stripe originates behind the eye and extends onto the neck and then breaks up into smaller white blotches. The lower stripe begins anteriorly at the supralabials, tracing posteriorly through the ear and over the arm, breaking into smaller white blotches on the anterior third of the body. Between these two stripes lies a broad dark brown to black band. Diagnostic, evenly-spaced green to blue spots start above the arm and extend posteriorly to the groin in both sexes. Beneath the lower interrupted white stripe / blotches lies another narrow band of brown to black scales, sometimes accompanied by orange spots or blotches extending onto the venter. During the breeding season, males exhibit more prominent orange flanks, while the lower white stripe and lateral head become vivid yellow anteriorly. The specimen from the DRC (MTSN 9947) exhibits a bright orange lower jaw. The venter is typically plain white but can have light grey colouration.	en	Conradie, Werner, Keates, Chad, Greenbaum, Eli, Lobón-Rovira, Javier, Tolley, Krystal A., Benito, Max, Vaz Pinto, Pedro, van Breda, Reuben V., Verburgt, Luke (2025): Systematics of African rough-scaled lizards, with description of two new species from eastern Angola (Squamata: Lacertidae: Ichnotropis Peters, 1854). Vertebrate Zoology 75: 627-672, DOI: 10.3897/vz.75.e167366
CBA5F90B078E5964B4D1B7B663DFDD50.taxon	distribution	Distribution. Known from western Tanzania, south to northern Malawi, and eastward to northern Zambia and southern DRC (Fig. 3).	en	Conradie, Werner, Keates, Chad, Greenbaum, Eli, Lobón-Rovira, Javier, Tolley, Krystal A., Benito, Max, Vaz Pinto, Pedro, van Breda, Reuben V., Verburgt, Luke (2025): Systematics of African rough-scaled lizards, with description of two new species from eastern Angola (Squamata: Lacertidae: Ichnotropis Peters, 1854). Vertebrate Zoology 75: 627-672, DOI: 10.3897/vz.75.e167366
