Bocage (1866) described Ichnotropis bivittatus (= bivittata) based on a series of specimens collected from Duque de Bragança [= Calandula], Malanje Province, Angola, despite Günther (based on material that Bocage had sent to him) considering them to be the same as I. capensis. Boulenger (1887) followed Günther and did not consider I. bivittata to be a valid species. This prompted Bocage (1895) to relegate I. bivittata into the synonymy of I. capensis. It was not until Boulenger (1921) reviewed the family Lacertidae that I. bivittata was reinstated as a valid species. This taxonomic revision was followed by most subsequent authors, except for a brief period when I. bivittata was regarded as a subspecies of I. capensis (Laurent 1952; Hellmich 1957; Manaças 1963; Robertson et al. 1963). The above confusion led to much of the historical Angolan material being incorrectly assigned to I. capensis (Marques et al. 2018).

When Laurent (1964) described the subspecies I. b. pallida he distinguished it from the nominotypical form based on its duller dorsal colouration and differences in head scalation—specifically, its less pronounced keeled head striations, distinct interparietal shape, and small frontoparietals that were separated by the interparietal (see Ceríaco et al. 2020a: fig. 30). However, colouration in Ichnotropis, as in many lacertids, is highly variable and influenced by factors such as substrate, age and sex, rendering it an unreliable taxonomic character—except when comparing adult breeding male material, where it can provide useful diagnostic insights. Furthermore, the head scalation observed in the holotype appears to be aberrant, as the scalation differs from a topotypic specimen from Humpata (PEM R17934; Fig. 7). Specifically, the configuration where the frontoparietals are separated by the interparietal, which in turn is in contact with the frontal, has not been observed in any other Ichnotropis specimens examined in this study, including the topotypic Humpata specimen. Although phylogenetic analyses reveal notable divergence between the Humpata specimen and other I. bivittata samples, further research is necessary before making definitive taxonomic decisions regarding the validity of I. b. pallida. Therefore, we currently treat I. b. pallida as a junior synonym of I. bivittata.

In the same paper, Laurent (1952) described Ichnotropis capensis nigrescens based on two specimens exhibiting darker ventral surfaces. Notably, the paratype (BE_RMCA_Vert.R.1869), originating from Luluabourg [= Kananga, Kasaï-Central Province, DRC], had previously been designated as a paratype in the description of Ichnotropis overlaeti by de Witte and Laurent (1942). The initial classification of nigrescens as a subspecies of I. capensis was guided by Boulenger’s (1921) key, which emphasised the separation of the prefrontal from the anterior supraocular. Subsequently, Loveridge (1933) synonymised this subspecies with I. capensis. Upon examining the type specimens, along with two additional specimens housed at RMCA (BE_RMCA_Vert.R.15925 and BE_RMCA_Vert.R.16240) from Ndwa Village near Bolobo—proximate to the holotype’s locality—it was observed that they possess a short and rounded snout, a character consistent with members of the I. bivittata group (see Fig. 2). Consequently, these specimens are transferred to the I. bivittata group instead of I. capensis. Specifically, the holotype (BE_RMCA_Vert.R.14671) and the two additional specimens are assigned to I. bivittata sensu lato based on the presence of closely spaced pale spots (possibly yellow in life) above the forelimb, whereas the Kananga paratype (BE_RMCA_Vert.R.1869) is reassigned to I. tanganicana, based on shared morphological (supraocular in contact with supraciliaries) and colouration characteristics detailed in the species account below (evenly spaced white dorsolateral spots; described as being blue by de Witte and Laurent 1942). Given the substantial sequence divergence observed in our limited I. bivittata material, the name nigrescens may be applied to northern populations, particularly those from the Republic of the Congo, DRC and Gabon, should future studies support the recognition of a distinct species in this region.

Ichnotropis capensis nigrescens Laurent, 1952: 201 (new synonymy); Ichnotropis bivittata pallida Laurent, 1964: 64 (new synonymy).

BMNH 1946.9.3.47–48 (1866.6.11.3–4), ZMB 5827 [additional syntypes in Lisbon Museum were probably destroyed by a fire in 1978], collected from Duque de Bragança [= Calandula], Malanje Province, Angola by F.A.P. Bayão.

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: 109.7mm) 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.

Figure 4. 

Photographs in life of Ichnotropis bivittata depicting the closely spaced yellow spots just posterior to the forelimbs. Specimens photographed at A Cuito town (PEM R23530), Angola; B Cambau (FKH-0833), Angola; C Cangandala National Park, Angola; D Luando Reserve (CHL0675), Angola; and E Humpata (PEM R17934 – I. b. pallida), Angola. Photographs: A – Werner Conradie; B, C, D – Pedro Vaz Pinto; E – William R. Branch.

Figure 5. 

Syntype (ZMB 5827) of Ichnotropis bivittata from Duque de Bragança [= Calandula], Malanje Province, Angola. Photographs of body in A dorsal and B ventral views, and head in C dorsal, D lateral and E ventral views. Scale bars represent 10 mm. Photographs: Frank Tillack.

Figure 6. 

Holotype (BE_RMCA_Vert.R.14641) of Ichnotropis capensis nigrescens from Bolobo, Democratic Republic of the Congo. Photographs of body in A dorsal and B ventral views, and head in C dorsal, D lateral and E ventral views. Scale bars represent 10 mm. Photographs: Max Benito.

Figure 7. 

Topotypic (PEM R17934) Ichnotropis bivittata pallida from Humpata, Huila Province, Angola. Photographs of body in A dorsal and B ventral views, and head in C dorsal, D lateral and E ventral views. Scale bars represent 10 mm. Photographs: Werner Conradie.

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.

Ichnotropis bivittata inhabits wet Miombo woodlands, preferring open, sandy areas suitable for thermoregulation and foraging. It is a diurnal, terrestrial species and an active forager, primarily preying on small arthropods such as ants, beetles, and termites (Pietersen et al. 2021). Activity peaks during warmer periods and declines in cooler or wetter conditions.

When I. chapini was described, it was differentiated from its congeners based on the presence of an anterior supraloreal, thus having two anterior loreal scales (Schmidt 1919). However, additional material collected from Adra in northeastern DRC does not possess any anterior supraloreal (de Witte 1933; Laurent 1952). Despite the limited material available, the scalation observed in the type specimen appears to be anomalous. This is further supported by the fact that we have only recorded this condition (an anterior supraloreal) once for all of the other Ichnotropis specimens examined (n = 432). As in the previous species, I. chapini was assigned to the I. capensis group based on the observation that the prefrontal is separated from the anterior supraocular (Boulenger 1921). Examination of high-resolution photographs of the holotype (Fig. 8) and physical examination of additional material from RMCA showed that this species belongs to the I. bivittata group, based on the more rounded head (Fig. 2). This species also seems geographically well isolated from other species in the genus, but this might just be an effect of under-sampling. The seasonality of Ichnotropis species makes them hard to observe outside or the breeding season. Consequently, coupled with the absence of any modern material and thus molecular data, we retain this species as valid until more data become available.

Figure 8. 

Holotype (AMNH 10674) of Ichnotropis chapini from Aba, Haut-Uele Province, Democratic Republic of the Congo. Photographs of body in A dorsal and B ventral views, and head in C dorsal, D lateral and E ventral views. Scale bars represent 10 mm. Photographs: Lauren Vonnahme.

AMNH 10674, adult female, collected from Aba, Haut-Uele Province, DRC in July 1911.

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.

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.

Very little is known about this species, but it is expected to have similar habitat requirements to other Ichnotropis species.

Described based on five specimens retrieved from the crop of a Dark Chanting Goshawk (Melierax metabates) at the base of Serra do Moco (the geographical feature of Serra do Moco which includes the highest peak in Angola at 2620 m a.s.l., is often colloquially referred to as Mount Moco) (Marx 1956). Parker (1936) was actually the first to document this species as I. bivittata from Serra do Moco and alluded to its smaller dorsal scales. Remarkably, these smaller dorsal scales were one of the main diagnostic features when I. microlepidota was described. However, its taxonomic status has been disputed in the past (Mayer 2013) because of its resemblance to I. bivittata and the lack of precise locality data, given that the type series was found in the crop of a dark chanting goshawk (Marx 1956). No additional specimens were collected until PVP collected a topotypic specimen in October 2020 at Serra do Moco (Benito et al. 2025). Thanks to this new material, Benito et al. (2025) provided the first phylogenetic placement of the species, validating its taxonomic status and demonstrating that this species belongs to the I. bivittata group.

Figure 9. 

Photographs in life of Ichnotropis microlepidota (MHNCUP-REP0983) from Serra do Moco, Huambo Province, Angola (adapted from Benito et al. 2025). Photographs in A dorsolateral view of the full body and B lateral view of the head. Photographs: Pedro Vaz Pinto.

Figure 10. 

Ichnotropis microlepidota (MHNCUP-REP0983) specimen from Serra do Moco, Huambo Province, Angola (adapted from Benito et al. 2025). Photographs of body in A dorsal and B ventral views, and head in C dorsal, D lateral and E ventral views. Scale bars represent 10 mm. Photographs: Max Benito.

FMNH 74285, adult male, collected from the ‘foot of Mount Moco’ [= Serra do Moco], Huambo Province, Angola, by Gerd Heinrich on 19 September 1954.

FMNH 74283–84 (females), FMNH 74286–87 (males); same collection details as holotype.

MHNCUP-REP0983, adult male, collected at Serra do Moco, Huambo Province, Angola (–12.4554°, 15.1632°, 2300 m a.s.l.), on 18 October 2020 by Pedro Vaz Pinto (Benito et al. 2025); juvenile specimen collected at Serra do Moco, Huambo Province, Angola, 1500–1900 m a.s.l., in March 1934 by Karl Jordan (Parker 1936).

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 (FMNH74285, 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

Currently only known from the slopes of Serra do Moco, in the central Angolan highlands (Fig. 3).

The specimens from the type series were preyed upon by a dark chanting goshawk (Melierax metabates) (Marx 1956). The specimen collected by PVP (MHNCUP-REP0983) was found during the day on top of an exposed small rock in open montane grassland, with thick vegetation cover at 2300 m a.s.l. (Benito et al. 2025). The montane habitat in Serra do Moco is mainly formed by a thick layer of grass and small bushes as well as many rocks underneath. This type of habitat is likely to hinder the species detectability.

This species was described from the ‘East Coast [of] Lake Tanganyika’ in modern-day Tanzania based on a single subadult specimen that was collected in 1896. When Boulenger (1917) described I. tanganicana, he ascribed the holotype to a subadult male. However, after our examination of high-resolution photographs of the type specimen it was not possible to sex it, so we regarded it as an unsexed subadult specimen. Since its description, no additional material has been documented. However, due to the vague description provided by Boulenger (1917), the taxonomic status of this species has been questioned by some authors (Mayer 2013). On the other hand, based on some diagnosable head scalation features (i.e., supraoculars in contact with supraciliaries), this species was preliminary retained as valid in subsequent years (Spawls et al. 2002, 2018; van den Berg 2017; Uetz et al. 2025).

In this study, an adult female specimen collected from the mid-elevation Miombo woodlands west of the Kabobo Plateau, DRC (MTSN 9947; Fig. 11E) agreed with the description of I. tanganicana based on the supraocular arrangement, (i.e., anterior supraocular in direct contact with the supraciliaries), and the colouration (bronzy olive dorsum with three fine black stripes on nape). However, Boulenger (1917), in his description of the type specimen after 20 years of preservation, did not document the unique, evenly-spaced blue dorsolateral spots observed in the new DRC specimen (Fig. 11E). Based on this new information about the dorsal colouration, we revisited the literature, examined known museum specimens (previously ascribed to I. bivittata in eastern DRC and adjacent Zambia and Tanzania) and consulted online citizen science platforms.

Figure 11. 

Photographs in life of Ichnotropis tanganicana from across its range, depicting the evenly spaced dorsolateral blue spots. Photographs from A Rukwa, Tanzania; B Cambua, Democratic Republic of the Congo; C Rumphi, Malawi (https://www.inaturalist.org/observations/146895735); D Nyika National Park, Malawi (https://www.inaturalist.org/observations/146684850); E Kindingi, Lake Tanganyika (MTSN 9947), Democratic Republic of the Congo; and F Upemba National Park, Democratic Republic of the Congo (https://www.inaturalist.org/observations/249778421). Photographs: David Lloyd-Jones; B – Colin Tilbury, C – Marc Henrion, D – Tim Brammer, E – Wandege Muninga, D – Naftali Honig.

Figure 12. 

Holotype (BMNH 1946.9.3.49) of Ichnotropis tanganicana from ‘East coast of Lake Tanganyika’, Tanzania. Photographs of body in A ventral and B dorsal views, and head in C dorsal, D lateral and E ventral views. Photographs: Patrick Campbell.

Of special interest is the case of the first specimens of I. bivittata from Ipemi, Udzungwa Mountains, Tanzania, documented by Loveridge (1933). He states that, in comparison to the type, he regards his specimens as conspecific with I. bivittata and distinct from I. tanganicana, of which he also examined the type. However, he provides no further details. In his description of the specimens’ colour, he offered a detailed account of the colouration as follows: ‘…series of blotches which is rather more black than chestnut-brown having the appearance of ocelli by reason of a blueish-white central spot in each …’. Examination of high resolution images of the two Ipemi specimens in the Museum of Comparative Zoology (MCZ R30836–7) confirmed the presence of the unique blue lateral spots (although faded to white in preservative) and the dorsal colouration, but the supraoculars were not in contact with the supraciliaries, as reported in the type specimen (BMNH 1946.9.3.49) of I. tanganicana. This difference might have been the reason why Loveridge (1933) considered his material to be conspecific with I. bivittata rather than I. tanganicana.

Additionally, de Witte and Laurent (1952) again mentioned these unique dorsolateral blue spots in the colour description of I. overlaeti: “… from this place it is sometimes replaced by a series of small blue spots more or less bordered with black, extending to the base of the hind limbs; blue spots are also present on the upper band, between the front and hind limbs.” (translation from French to English). When we examined the type specimens of I. overlaeti at the RMCA (Fig. 13), we not only confirmed the remnants of blue lateral spots and the nape colouration, but we also confirmed the presence of contact between the supraoculars and the supraciliary scales, in agreement with I. tanganicana. Nevertheless, this feature was only present in the holotype (BE_RMCA_Vert.R.9691) and one of the original paratypes (BE_RMCA_Vert.R.1869, later used as a paratype for I. nigrescens). However, the other paratype material conformed morphologically to either I. bivittata (see above) or I. capensis sensu lato (see below).

Figure 13. 

Holotype (BE_RMCA_Vert.R.9691) of Ichnotropis overlaeti from Kapanga, Haut-Katanga Province, Democratic Republic of the Congo. Photographs of body in A ventral and B dorsal (note the evenly spaced white dorsolateral spots indicated by the arrows) views, and head in C dorsal, D lateral and E ventral views. Scale bars represent 10 mm. Photographs: Max Benito.

Other published sources showing photographs of I. bivittata (sic) with blue spots include de Witte (1933: plate 2, fig. 1) from southeastern DRC, Spawls et al. (2018: 202, bottom right) from southwestern Tanzania, and Phadima et al. (2024: 22; also on iNaturalist 146895735) from northwestern Malawi. Additional records were also found on iNaturalist (191773297, 146684850, 147210660, 87417155) and ReptileMap (169500) from DRC, Malawi and Zambia

Based on the combined evidence, all the above material can thus be confidently assigned to I. tanganicana. We therefore take this opportunity to expand on the original description of I. tanganicana and synonymise I. overlaeti with I. tanganicana.

Ichnotropis overlaeti de Witte & Laurent, 1942: 173 (new synonymy).

BMNH 1946.9.3.49 (96.5.14.14), collected from ‘East Coast [of] Lake Tanganyika’, Tanzania, presented to the museum by Mr. WH. Nutt in 1896.

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.

Known from western Tanzania, south to northern Malawi, and eastward to northern Zambia and southern DRC (Fig. 3).

The Lukwati specimen was discovered in grassland adjacent to Brachystegia woodland. This specimen exhibited peculiar leg-tucking behaviour, wherein it raised its body and folded its legs to the sides (Spawls et al. 2018; Lloyd-Jones pers. comm.). A gravid female was observed laying eggs in January (iNaturalist 146684850). Shelled eggs in the oviducts of one specimen measured 13.5 mm × 6.5 mm (Robertson et al. 1963). Stomach contents were documented to contain Acrididae, Mantidae, Isoptera, and Araneae (Robertson et al. 1963).

This is the most recently described species of Ichnotropis. It was described from the western Zambezi Region in north-eastern Namibia, based on only three specimens, and was distinguished from sympatric I. capensis based on its larger size, rounded head and dorsal colouration (Broadley 1967b). This is a rarely documented species and it is only known from the type series, four additional specimens collected from north-eastern Namibia (Haacke 1970), one specimen from Khaudum, Namibia (van Breda 2023), and one specimen from western Zambia (Pietersen et al. 2017). Conradie et al. (2022a) tentatively assigned material from eastern Angola to this species based on shared morphology, but phylogenetic analyses (see Results) recover it as a separate sister lineage, which represents a candidate new species described below.

USNM 163989, an adult male, collected ‘25 miles west of Mohembo, Botswana, on the border of the Caprivi Strip (South West Africa)’, Namibia by T.N. Liversedge and S.W. Goussard on 20 May 1967.

NMZB-UM 16278 (male) and USNM 163990 (juvenile); same collection details as holotype.

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 (RE211206D1/NMNW R12212 – 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).

Figure 15. 

Photographs in life of juvenile Ichnotropis grandiceps. Specimens photographed from A Ngonye Falls (TM 86237), Zambia and B Chitokoloki, Zambia. Photographs: A – Darren Pietersen; B – Frank Willems.

Figure 16. 

Ichnotropis grandiceps (RE211206D1/NMNW R12212) specimen from Khaudum, Namibia. Photographs of body in A ventral and B dorsal views, and head in C dorsal, D lateral and E ventral views. Photographs: Werner Conradie.

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.

Ontogenetic colour differences have been observed between juveniles and adults (this study). Found in sympatry with I. capensis sensu lato. Associated with Baikiaea woodland on deep Kalahari alluvial sands and hard lime-rich soils in open woodland (Haacke 1970; Pietersen et al. 2021).

Ichnotropis cf. grandiceps – Conradie et al. (2022a: 198); Ichnotropis aff. grandiceps – Benito et al. (2025: 893).

PEM R23420 (field number WC-4816), adult male, collected from Cuando River source (–13.0035°, 19.1275°, 1343 m a.s.l.), Moxico Province, Angola by Werner Conradie and James Harvey on 21 November 2016.

4 specimens: a) PEM R23361 (field number WC-4063) and PEM R23362 (field number WC-4056), adult females, collected on the road between Cuanavale River source camp and Samanunga village (–13.0380°, 18.8298°, 1605 m a.s.l.), Moxico Province, Angola by Werner Conradie and Luke Verburgt on 13 March 2016; b) PEM R23421, adult male, same collection details as holotype; c) PEM R23482 (field number WC-4804), adult male, collected from Cuando River source, trap 4 (–13.0016°, 19.1296°, 1372 m a.s.l.), Moxico Province, Angola by Werner Conradie and James Harvey on 15 November 2016.

12 specimens: a) PEM R23279–80; INBAC (no number), collected from Cuanavale River source lake (–13.0933°, 18.8940°, 1367 m a.s.l.), Moxico Province, Angola by Werner Conradie on 1 March 2016; b) PEM R23299–300, grassland west of Cuanavale River source en route to Samanunga village (–13.0751°, 18.8848°, 1366 m a.s.l.), Moxico Province, Angola by Werner Conradie and Luke Verburgt on 16 March 2016; c) PEM R23303–9, trap 4 km upstream from Cuanavale River source lake (–13.0508°, 18.8973°, 1380 m a.s.l.), Moxico Province, Angola by Werner Conradie from 28 February to 15 March 2016.

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.

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 1967b; 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 16S and >16.3% for ND4 sequence divergence from other Ichnotropis species (Table 2).

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.

(In life, breeding colouration; similar to Fig. 17A): Above uniform reddish-brown, with small black spots on the lower body and anterior third of tail; dorsolateral bands as in preserved colouration, but more vivid; gular, chin shields and posterior section of white dorsolateral band bright yellow to anterior of the forelimb insertion, fading to just behind the arm; front limbs brick red dorsally, hind limbs dark grey anteriorly and brick red posteriorly; venter (except gular) white. Colouration (in preservative; Fig. 18): Above pale grey-brown, with a few scattered dark brown to black spots (not covering more than one scale) on lower body and upper tail; a well-defined broad (covering 3–4 scales at midbody) black dorsolateral band extends from the tip of the snout to the groin; below this black dorsolateral band is a second narrow white band (covering 1–2 scales at midbody) which also extends from the tip of the snout to the groin; below this white band is another narrow black band extending from the mental, along the edge of the jaw (edge of supra- and infralabials), to just behind the forelimb insertion, where it fades to fine specks towards the groin. Limbs dorsally brown and ventrally white; venter white.

Figure 17. 

Photographs in life of Ichnotropis robusta sp. nov. Photographs of A adult male (PEM R23482) from Cuando River source, Angola; B adult female (PEM R23362) from the road between Cuanavale River source camp and Samanunga village, Angola; C juvenile (PEM R23304) from 4 km upstream of Cuanavale River source lake, Angola; and D juvenile (PEM R23300) from the road between Cuanavale River source camp and Samanunga village, Angola. Photographs: A, B, C – Werner Conradie; D – Luke Verburgt.

Figure 18. 

Holotype (PEM R23420) of Ichnotropis robusta sp. nov. from Cuando River source, Moxico Province, Angola. Photographs of body in A ventral and B dorsal views, and head in C dorsal, D lateral and E ventral views. Scale bars represent 10 mm. Photographs: Werner Conradie.

The paratypes are in general agreement with the holotype in most regards, differing only in: Two large supraoculars, which are separated from the supraciliaries by a single row of smaller scales (7–10) and preceded by a cluster of smaller scales (3–5) (except in PEM R23482, the 1^st supraciliary is in narrow contact with the anterior supraocular on the right side); 1–3 post-supraocular scales; large occipital scale that separates the two interparietals and extends well past their posterior edge; two loreal scales present, which are separated from the anterior supraocular by 1–2 scales; 4–5 (mostly 5) supralabials in front of the subocular; 6–7 (mostly 6) infralabials; five chin shields, with the anterior three in broad contact; five supraciliaries; 43–48 (average: 45.0) midbody scale rows; 26–30 (average: 27.8) transverse ventral scale rows; 20–23 subdigital lamellae under the 4^th toe; 11–14 femoral pores per thigh. Size: Adult specimens varied from 71.9–78.8 mm (mean: 75.3 mm) SVL and 121–140.0 mm (mean: 133.0 mm) TAIL. Largest female: 78.8 mm SVL + 136 mm TAIL (PEM R23361); largest male: 73.5 mm SVL + 96.0 mm truncated tail (Holotype). Colouration (Fig. 17B): Females with numerous scattered black scales on back and tail, scattered white scales along the upper edge of the dorsolateral dark brown band, continuing onto the tail. Lower dorsolateral black band broken from behind the head to the tail.

Prefrontal separated from the anterior supraocular by 1–2 smaller scales (in contact in PEM R23300 and R23299 – both sides; PEM R23309 – left side only) and separated from supraciliaries by a smaller scale (except PEM R23305 and PEM R23299); two large supraoculars, which are separated from the supraciliaries by one row (two in PEM R23280 and R23303) of small scales (7–10) and preceded by a cluster of smaller scales (2–7) (except in PEM R23300, where the 1^st supraciliary is in narrow contact with the anterior supraocular on the right side; in narrow contact on the left side in PEM R23306; in PEM R23307, the 2^nd and 3^rd supraciliaries are in contact with the posterior supraocular on the right and anterior supraocular on the left); 1–2 post-supraocular scales; two loreal scales present, which are separated from the anterior supraocular by 1–2 scales; subocular in contact with the lip; 4–6 (mostly five) supralabials in front of the subocular; 6–7 (mostly six) infralabials; five chin shields, with the anterior three in broad contact; 5–6 (mostly five) supraciliaries; 43–47 (average: 45.0) midbody scale rows; 9–10 longitudinal rows of enlarged ventral plates; 30–33 (average: 32.4) transverse ventral scale rows; 20–25 subdigital lamellae under the 4^th toe; 11–13 femoral pores per thigh. Size: Juvenile specimens varied from 35.6–51.1 mm (mean: 44.6 mm) SVL and 68.0–100.9 mm (mean: 88.2 mm) TAIL. Colouration (Fig. 17C, D): In juveniles, the dorsum is grey anteriorly and orange posteriorly, with scattered black and white specks, and a narrow mustard coloured dorsolateral band. The dorsum of the tail is orange with scattered black and white scales. The venter is white.

Only recorded from the headwaters of the Okavango (Cuito and Cuanavale Rivers) and Cuando Rivers in central Angola (Fig. 14).

Ontogenetic colour differences have been observed between juveniles and adults. Breeding colouration males and gravid females were collected in November 2016, while non-breeding females and juveniles were found in February–March 2016. Juveniles were only observed on sandier areas around the source of the Cuanavale River, while two adult females were found on the elevated grassland ridges surrounding the river. Found in sympatry with I. capensis sensu lato.

Ichnotropis capensis was originally described from the “sandy deserts around Latakoo”, which corresponds to the present-day Kuruman area in the Northern Cape province, South Africa. Since its original description, no additional specimens have been collected from the type locality or proximate areas. The nearest known record today is from Giya Camp in southern Botswana, approximately 340 km north of the type locality. This gap in distribution may be due to limited sampling effort in the region (Tolley et al. 2023), or alternatively, it may reflect uncertainty or inaccuracy in the locality information provided by Smith (1838), who was traveling extensively across the northern provinces of South Africa at the time. Due to the uncertainty surrounding the type locality and the fact that the type specimen remains unaccounted for in the BMNH (P. Campbell, pers. comm. 23 January 2024), the designation of a neotype is recommended to further stabilise the taxonomic status of Ichnotropis capensis. However, we refrain from taking this action at present, because we lack comparative material from the southern and eastern parts of the species’ distribution, including areas near the type locality of Ichnotropis longipes (Mazoë and the region between Umtali and Marandellas). Although we have examined photographs of the I. longipes type specimens (all subadults), we choose not to assign this name to any of the remaining I. capensis clades until additional material becomes available and a neotype for I. capensis can be designated.

Jacobsen et al. (2010) noted the presence of disjunct populations of Ichnotropis capensis between the east coast of southern Africa and the interior. Populations from the eastern coastal regions of South Africa and adjacent Mozambique form a distinct monophyletic clade that differs genetically by 4.1–6.4% 16S uncorrected p distance from other I. capensis populations. Although this coastal material could be referred to Ichnotropis macrolepidota Peters, 1854—originally described from Lourenço Marques (now Maputo), Mozambique—our analysis indicates that the observed genetic differences fall within the expected range of intraspecific variation and likely reflect geographic separation between populations. This was further supported by the lack of clear morphological or geographical separation. As such, we consider Ichnotropis macrolepidota a junior synonym of I. capensis. Nevertheless, if future studies demonstrate that the eastern coastal populations represent a distinct operational taxonomic unit (OTU), the name I. macrolepidota should be resurrected for it.

Algyra capensis Smith, 1838: 94; Tropidosaura Dumerelii Smith, 1849: appendix 7; Ichnotropis macrolepidota Peters, 1854: 617; Ichnotropis longipes Boulenger, 1902: 17.

BMNH 1865.5.4.56, collected from the ‘Sandy deserts around Latakoo’ [= Kuruman], Northern Cape, South Africa by Andrew Smith.

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.

Figure 20. 

Photographs in life of Ichnotropis capensis sensu lato from A Cuando River source, Angola (adult male), B Cuanavale River source, Angola (adult female), C Xai-Xai, Mozambique (adult male), D Ngonye Falls, Zambia (adult female), E Zambezi Region, Namibia (adult male), F Kosi Bay, South Africa (adult male). Photographs: A–D – Werner Conradie; E, F – William R. Branch.

Figure 21. 

Ichnotropis capensis sensu lato (PEM R27396) from middle Quembo River bridge camp, Moxico Province, Angola. Photographs of body in A ventral and B dorsal views, and head in C dorsal, D lateral and E ventral views. Scale bars represent 10 mm. Photographs: Werner Conradie.

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).

Figure 22. 

Syntype (ZMB 6123) of Ichnotropis macrolepidota from Lourenço Marques [= Maputo], Mozambique. Photographs of body in A ventral and B dorsal views, and head in C dorsal, D lateral and E ventral views. Scale bars represent 10 mm. Photographs: Frank Tillack.

Figure 23. 

Syntype (BMNH 1946.8.4.23) of Ichnotropis longipes from Mazoë, Zimbabwe. Photographs of body in A ventral and B dorsal views, and head in C dorsal, D lateral and E ventral views. Scale bars represent 10 mm. Photographs: Patrick Campbell.

This species prefers arid to mesic savanna habitats. It is a diurnal lizard, actively foraging for small invertebrates such as termites, spiders, beetles, and grasshoppers. Females lay up to nine eggs per clutch, typically during the summer months from October to November. The eggs measure approximately 5.5–7.0 mm by 8.5–9.5 mm. The incubation period ranges from 56 to 77 days, with hatchlings emerging between January and March. Females may produce up to two clutches within a single breeding season.

Ichnotropis capensis overlaeti – Laurent (1950: 12, in part); Ichnotropis capensis – Conradie et al. (2022a: 198, in part); Ichnotropis aff. capensis – Benito et al. (2025: 893).

PEM R23410 (field number WC-4558), adult male, collected from Lungwebungu River camp bridge crossing, (–12.5835°, 18.6660°, 1304 m a.s.l.), Moxico Province, Angola by Werner Conradie and Luke Verburgt on 22 October 2016.

6 specimens: a) PEM R23409 (field number WC-4557), adult male, same collection details as holotype; b) PEM R23502 (field number WC-4522), and PEM R23531 (field number WC-4560), adult male and female respectively, collected from Sombanana Village (–12.3108°, 18.6239°, 1403 m a.s.l.), Moxico Province, Angola by Werner Conradie and Luke Verburgt on 9 October 2016; c) PEM R23505–7 (field number WC-4543, WC-4562 and WC-4563, respectively), adult males, collected from Lake Tchanssengwe (–12.4102°, 18.6348°, 1414 m a.s.l.), Moxico Province, Angola by Werner Conradie and Luke Verburgt on 21 October 2016.

9 specimens: a) PEM R19903 (field number TB 44) and PEM R19905 (field number TB 46), adult female and male respectively, collected from Camp Chiri, Miombo forest/camp (–9.3969°, 20.4319°, 1004 m a.s.l.), Lunda-Sul Province, Angola by Tom Branch on 24 October 2008; b) PEM R23977 (field number WC-6267), juvenile, collected from near Lungwebungu Trap 2 (–12.5820°, 18.6656°, 1208 m a.s.l.), Moxico Province, Angola by Werner Conradie and Alex Rebelo on 22 April 2018; c) PEM R23986 (field number WC-6266), juvenile, collected from Lungwebungu Trap 1 (–12.5801°, 18.6674°, 1298 m a.s.l.), Moxico Province, Angola by Werner Conradie and Alex Rebelo on 22 April 2018; d) PEM R23996–7 (field numbers WC-6291 and WC-6292, respectively), juveniles, collected from Lake Tchanssengwe (–12.4140°, 18.6442°, 1393 m a.s.l.), Moxico Province, Angola by Werner Conradie and Alex Rebelo on 23 April 2018; e) BE_RMCA_Vert.R.17490, 17492 (adult males) and BE_RMCA_Vert.R.17491 (adult female), collected from the Dundo region, Lunda-Norte Province, Angola by Barros Machado on 14 December 1947.

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.

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 16S and >12.3% for ND4 sequence divergence from other Ichnotropis species (Table 2).

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.

(In life, breeding colouration; Fig. 24A): The dorsum varies from grey on the head and nape to reddish-brown on the dorsum and grey on the tail. The side of the body has a dark black band that originates on the snout and run posteriorly to the tail, where it disappears at the tip. Below this black band is a white stripe that originates on the snout, runs below the eye to the front limb, is less distinct between the fore- and hind limb, and then fades onto the tail. Below this white line is another black stripe that originates on the snout, running along the edges of the supra- and infralabials to the front limbs. The white stripes on the sides of the head and the gular region are pale yellow. Below the black band and white flank stripe (that appears as scattered white and grey spots in places) is an orange band. Limbs are brick red and grey. Chin shields and gular scales have scattered black blotches of varying sizes. The venter is white with scattered black specks. Colouration (in preservative; Fig. 25): Above pale grey-brown; a well-defined broad (covering 3–4 scales at midbody) black dorsolateral band extends from the tip of the snout to the groin; below this black dorsolateral band is a narrow white band (covering one scale at midbody) which extends from the tip of the snout to just posterior to the front limb, fading towards the groin and tail base. Below this white band is another narrow black band extending from the mental, along the edge of the jaw (edge of supra- and infralabials) to just posterior to the forelimb insertion. Flanks bear a light brown band (two scales wide); gular and chin shields with scattered black spots or blotches; limbs dorsally brown and ventrally white; 2–3 white spots on the anterior surfaces of the legs; venter white with scattered black specks.

Figure 24. 

Photographs in life of Ichnotropis longicorpa sp. nov. Photographs of A adult male (PEM R23410) from Lungwebungu River camp bridge crossing, Angola, and B adult female (PEM R23531) from Sombanana village, Angola. Photographs: Werner Conradie.

Figure 25. 

Holotype (PEM R23410) of Ichnotropis longicorpa sp. nov. from Lungwebungu River camp bridge crossing, Moxico Province, Angola. Photographs of body in A ventral and B dorsal views, and head in C dorsal, D lateral and E ventral views. Scale bars represent 10 mm. Photographs: Werner Conradie.

The paratypes are in agreement with the holotype in scalation, with only minor variation: Prefrontal always separate from the anterior supraocular by one scale (except PEM R19905 on right side); frontonasal always separate from the 1^st supraciliaries (except in PEM R23409 and PEM R23506); two (rarely three) scales separating anterior supraocular from the posterior loreal; cluster of 3–9 scales in front of the anterior supraocular; single row of 6–9 scales separating the supraoculars from the supraciliaries; 1–4 post-supraoculars; four supraciliaries; 4–5 supralabials; 6–7 infralabials; five chin shields, with first three in contact (PEM R23505 has six chin shields, with the first four in contact on the left side); 9–10 transverse ventral plates; 25–31 longitudinal ventral plates; 34–41 midbody scale rows; 19–24 subdigital lamellae under the 4^th toe; 10–13 femoral pores on each thigh. PEM R23409 exhibits some aberrant head scalation in that the anterior loreal seems to be divided, forming a supraloreal that separates the anterior loreal from the frontonasal and the parietal, and the posterior loreal is divided into two scales. Size: Adult specimens varied from 62.7–71.2 mm (mean: 65.3 mm) SVL and 117.0–160.0 mm (mean: 141.2 mm) TAIL. Largest female: 65.1 mm SVL (PEM R19905 – Camp Chiri, Angola); largest male: 71.2 mm SVL (BE_RMCA_Vert.R.17492 – Dundo, Angola). Colouration of all males are in agreement with the holotype. The paratype female (PEM R23531; Fig. 24B) is duller in colouration, almost uniformly reddish-brown dorsally and grey laterally, with no white stripes or black bands.

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.

This species was not found to be sympatric with any other Ichnotropis species, but it occurs in close geographical proximity to I. capensis sensu lato and I. robusta sp. nov. This species is associated with wet Miombo woodland.