Systematics of African rough-scaled lizards, with description of two new species from eastern Angola (Squamata: Lacertidae: Ichnotropis Peters, 1854)

627 Systematics of African rough-scaled lizards, with description of two new species from eastern Angola (Squamata: Lacertidae: Ichnotropis Peters, 1854) Werner Conradie1,2,3, Chad Keates3,4,5, Eli Greenbaum6, Javier Lobón-Rovira7,8, Krystal A. Tolley9, Max Benito10, Pedro Vaz Pinto7,8,11, Reuben V. van Breda12, Luke Verburgt3,13,14 1 Port Elizabeth Museum (Bayworld), Beach Road, Humewood, Gqeberha, 6013, South Africa 2 Department of Conservation Management, Natural Resource Science and Management Cluster, Faculty of Science, George Campus, Nelson Mandela University, George, 6529, South Africa 3 National Geographic Okavango Wilderness Project, Wild Bird Trust, Santon, 2196, South Africa 4 South African Institute for Aquatic Biodiversity, Makhanda, 6140, South Africa 5 Ft. Lauderdale Research and Education Center, University of Florida, Davie, FL 33314, USA 6 Department of Biological Sciences, University of Texas at El Paso, El Paso, TX 79968, USA 7 CIBIO, Centro de Investigacao em Biodiversidade e Recursos Geneticos, InBIO Laboratorio Associado, Campus de Vairao, Universidade do Porto, 4485-661 Vairao, Portugal 8 BIOPOLIS Program in Genomics, Biodiversity and Land Planning, CIBIO, Campus de Vairao, 4485-661 Vairao, Portugal 9 Department of Zoology, University of Johannesburg, Auckland Park, Johannesburg, 2006, South Africa 10 Universidad Internacional Menéndez Pelayo, Calle de Isaac Peral 23, 28040 Madrid, Spain 11 Fundação Kissama, Rua 60 Casa 560, Lar do Patriota, Luanda, Angola 12 Unit for Environmental Sciences and Management, North-West University, Potchefstroom, 2531, South Africa 13 Enviro-Insight CC, Unit 8 Oppidraai Office Park, Pretoria, 0050, South Africa 14 Department of Zoology and Entomology, University of Pretoria, Pretoria, 0001, South Africa https://zoobank.org/F811EE38-D26A-4C49-A863-D2800F54BA7B Corresponding author: Werner Conradie ([email protected]) Academic editor Uwe Fritz | Received 13 July 2025 | Accepted 3 October 2025 | Published 21 November 2025 Citation: Conradie W, Keates C, Greenbaum E, Lobón-Rovira J, Tolley KA, Benito M, Vaz Pinto P, van Breda RV, Verburgt L (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. https://doi.org/10.3897/vz.75.e167366 Abstract Ichnotropis is a genus of medium-sized lacertids endemic to sub-Saharan Africa, characterised by rough head shields. The genus currently comprises six nominal species distributed across much of southern, central, and eastern Africa. Some species are apparently active at only certain times of the year, resulting in limited specimen collections and severely hampering research. This scarcity of material has historically made comprehensive systematic reviews of the genus difficult and has led to the description of numerous regional morphological variants as distinct species or subspecies. Material collected in recent years has enabled us to provide a new phylogenetic hypothesis of Ichnotropis using two mitochondrial genes (16S and ND4) and two nuclear genes (c-mos and RAG-1). Our phylogenetic dataset includes 56 individuals representing five of the six currently recognised species (excluding I. chapini). Additionally, the broad geographical sampling of the widespread I. capensis group has allowed us to explore the taxonomic status of several species and subspecies within the group. As a result, we demonstrate the monophyly of Ichnotropis in relation to other African lacertids and present the most comprehensive phylogeny of the genus to date. We also provide the first phylogenetic placements for I. tanganicana and I. grandiceps, which allows us to validate their taxonomic statuses. Furthermore, we recovered a new cryptic species closely related to I. grandiceps, and identified several well-supported clades within the I. capensis group, all corroborated by multi-locus species delimitation analyses. One of these clades is described herein as a new species, while the remaining taxa of interVertebrate Zoology 75, 2025, 627–672 | DOI 10.3897/vz.75.e167366 Copyright Werner Conradie et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Conradie W et al.: Systematics of African rough-scaled lizards 628 est are discussed and highlighted for future investigation. Based on our findings, we recommend the following taxonomic revisions: Ichnotropis longipes and I. macrolepidota should remain synonyms of I. capensis; I. bivittata pallida and I. capensis nigrescens are treated as a junior synonyms of I. bivittata; and I. overlaeti is considered a junior synonym of I. tanganicana. Although we could not determine the phylogenetic placement of I. chapini due to the lack of genetic material, its head morphology and scalation support its reassignment to the I. bivittata group. Thus, it is retained as a valid species pending the availability of new material for further taxonomic actions. In conclusion, this study resolves several long-standing taxonomic issues within one of Africa’s most understudied lacertid genera and lays a solid foundation for future research on the genus Ichnotropis. Keywords Africa, cryptic species, reptiles, sub-Sahara, taxonomy Introduction The family Lacertidae comprises 388 recognised species and numerous subspecies distributed across Africa, Europe, and Asia (Uetz et al. 2025). Although the main species hotspot can be found in the Palearctic, particularly in arid zones such as northern Africa and the Arabian Peninsula, substantial diversity also occurs farther south. Central and southern Africa—spanning much of sub-Saharan Africa, from south of the Congo River Basin to the southernmost tip of the continent—harbours at least 65 recognised species across 11 genera (Uetz et al. 2025). This accounts for approximately 17% of global lacertid diversity, underscoring the region’s significance as a centre of endemism and evolutionary diversification within the family. Despite its richness, the lacertid fauna of central and southern Africa remains comparatively understudied (Tolley et al. 2016), highlighting a gap in our knowledge on the family’s biogeography and evolutionary history. In recent years, several phylogenetic studies have attempted to address this gap by focussing on central and southern African lacertid genera, including Adolfus (Greenbaum et al. 2011, 2018), Pedioplanis (Makokha et al. 2007; Conradie et al. 2012; Childers et al. 2021; Parrinha et al. 2021), Meroles (Edwards et al. 2012, 2013a), Nucras (Edwards et al. 2013b; Branch et al. 2019; Bauer et al. 2019, 2020, 2025; Baptista et al. 2020), and Heliobolus (Marques et al. 2022a). Some of these and other studies have also focused on alpha taxonomy (Greenbaum et al. 2011; Edwards et al. 2013a; Englender et al. 2013; Wagner et al. 2014). Collectively, these studies have improved our knowledge of lacertid systematics and diversity, resulting in the description of multiple new species. Despite this progress, detailed phylogenetic and phylogeographic studies are still lacking for certain genera, such as Holaspis, Ichnotropis, Latastia, and Tropidosaura. The African lacertid genus Ichnotropis Peters, 1854 comprises several small to medium-sized, rough-scaled, terrestrial species that inhabit mesic to xeric savannas in central and southern Africa (Branch 1998; Spawls et al. 2018; Pietersen et al. 2021; Benito et al. 2025). Interestingly, the genus includes several sympatric species that are presumed to have an annual reproductive strategy, in which they breed asynchronously, and this might be a strategy to reduce interspecific competition (Broadley 1967a, 1974, 1979; Jacobsen 1987). However, our knowledge of this ecological phenomenon is hampered by the lack of robust ecological studies on this genus. Knowledge of the genus is based largely on the work of Boulenger (1921) and subsequent species descriptions by de Witte and Laurent (1942), Laurent (1952), Marx (1956) and Broadley (1967b), as well as a recent comprehensive synthesis of the genus provided by van den Berg (2017). Six species are currently recognised as valid: Ichnotropis bivittata Bocage, 1866; Ichnotropis capensis (Smith, 1838); Ichnotropis chapini Schmidt, 1919; Ichnotropis grandiceps Broadley, 1967; Ichnotropis microlepidota Marx, 1956; and Ichnotropis tanganicana Boulenger, 1917. Furthermore, the subspecies, I. bivittata pallida Laurent, 1964 is considered to be valid, while additional species or subspecies (i.e., I. capensis nigrescens Laurent, 1952; I. macrolepidota Peters, 1864; I. longipes Boulenger, 1902; I. overlaeti de Witte & Laurent, 1942) have been described but are currently not considered valid, or are controversial (Uetz et al. 2025). Confusion therefore persists in the literature regarding the number of accepted species, their diagnostic characteristics, and synonyms (van den Berg 2017). Thus, in the absence of a strong phylogenetic framework for Ichnotropis that can be used to clarify species boundaries, and a detailed morphological revision, this confusion is likely to continue. In recent years, collections of Ichnotropis across its range have improved, particularly due to a concerted effort to carry out biodiversity surveys in under-sampled regions such as Angola (Conradie et al. 2016, 2022a; Benito et al. 2025), the Democratic Republic of the Congo (DRC) (Keates 2024) and Zambia (Pietersen et al. 2017). This has allowed for validation of the taxonomic status of some described species within a phylogenetic framework in the current study, as well as providing a more informed knowledge baseline regarding the diversity and evolution of this group in central and southern Africa. Through this process, we aim to stabilise the taxonomy of the group and lay a foundation for future work. Vertebrate Zoology 75, 2025, 627–672 629 Material and Methods Sampling Over the past decade, multiple new Ichnotropis specimens were collected across central and southern Africa, especially from Angola, Democratic Republic of the Congo (DRC), Mozambique, South Africa and Zambia ( Table 1). Initial species identifications were based on key diagnostic features (e.g., dorsolateral colouration, head scalation, supraocular–supraciliar-prefrontal contacts) reported in the literature (Boulenger 1921; Marx 1956; Broadley 1967b; van Berg 2017), supplemented by examination of type and topotypic or near-topotypic material (see Morphology below), and by considering geographic proximity to the respective type localities. DNA samples were collected from either liver, muscle or tail tips and preserved in 99% ethanol after which voucher specimens were fixed in 10% formalin and transferred to 70% ethanol for long-term storage at the Port Elizabeth Museum (PEM, South Africa), the Colecção Herpetológica do Lubango (CHL, Angola), the National Museum Namibia (NMNW, Namibia), the Museu de História Natural e da Ciência–Universidade do Porto (MHNC-UP, Portugal), the Museo delle Scienze di Trento (MUSE, Italy), and the Fundação Kissama Collection (FKH, Angola). Representative material was also deposited with the Ministry of Environment, Luanda, Angola (MINAMB), and the Museu de História Natural de Maputo, Mozambique (MHNM). For all newly collected specimens, geographic coordinates were recorded in decimal degrees (WGS84 datum, four decimal places) using a handheld GPS, and elevation in meters above sea level (a.s.l.). DNA extraction, amplification and sequencing DNA was isolated from tissue samples using a standard salt extraction method (Aljanabi and Martinez 1997). Standard Polymerase Chain Reaction (PCR) procedures were utilised to amplify one partial mitochondrial ribosomal gene (16S rRNA [16S]), one partial mitochondrial gene (NADH-dehydrogenase subunit 4 [ND4]), and two partial nuclear genes (oocyte maturation factor [c-mos], recombination activating gene 1 [RAG-1]). Each amplification was conducted with a PCR mixture of 25 µl total volume, containing 12.5 µL Taq DNA Polymerase 2x Master Mix (Ampliqon; 3 mM MgCl2, 0.4 mM dNTPs and Ampliqon Taq DNA polymerase), 2 µl forward primer (10 µM), 2 µl reverse primer (10 µM), and 8.5 µl of extracted genomic DNA (20–50 ng/µl) and water combined. The cycling profile for all the genes was as follows: Initial denaturation step at 94°C for 5 min, followed by 30–37 cycles of 94°C for 30 s, 42–58°C for 45 s, and 72°C for 45 s, with a final extension at 72°C for 8 min. The cycling profile for the genes differed only in the annealing temperature and the number of cycles (Table S1). The prepared PCR products were sent to Macrogen Corporation in Amsterdam, The Netherlands, for purification and sequencing with forward primers. Phylogenetic analyses For the phylogenetic analyses, 46 Ichnotropis individuals were sequenced, supplemented with sequences from 10 individuals available from GenBank. Six Meroles squamulosus individuals were used as outgroup taxa ( Table 1). For each gene, new sequences were checked and edited using BioEdit Sequence Alignment Editor v.7.2.5 (Hall 1999), and then aligned with the GenBank sequences in MEGA v.7.0.27 (Tamura et al. 2013), using the ClustalW v.1.6 alignment algorithm with default settings for alignment parameters (Thompson et al. 1994). Single gene maximum likelihood (ML) trees were created in IQ-TREE v.2.1.3 (Nguyen et al. 2015) to visually check the placement of sequences and compare topologies. DAMBE v.7.3.1 (Xia 2018) was used to test for saturation using the individual as well as combined first and second codon positions of each protein-coding gene. As none of the genes were found to be saturated, the genes were not partitioned by codons. In addition, congruence between individual gene datasets, as well as between mitochondrial and nuclear datasets, were tested using 100 replicates of the partition-homogeneity test (PHT) (Farris et al. 1994, 1995) in PAUP* v.4.0a169 (Swofford 2003). All gene-tree combinations were congruent, allowing for the creation of a concatenated dataset of 2128 base pairs for further phylogenetic analyses, with the individual gene alignments joined using SequenceMatrix v.1.8.2 (Vaidya et al. 2011). The optimal partition scheme and best-fitting models of molecular evolution were selected using ModelFinder implemented in IQ-TREE (Chernomor et al. 2016; Minh et al. 2021). The following settings were used: -p partition file (each partition has its own evolution rate), a greedy strategy and the FreeRate heterogeneity model excluded (only invariable sites and Gamma rate heterogeneity considered) (Chernomor et al. 2016; Kalyaanamoorthy et al. 2017). As MrBayes is not able to implement all the IQ-TREE models, the -mset mrbayes command was used to ensure that only models that were compatible with MrBayes were selected. The best-fitting model schemes selected for each dataset were as follows: 16S: GTR+G, ND4: GTR+I+G, c-mos+RAG-1: HKY+G. Maximum likelihood phylogenies were generated in IQ-TREE, using a random starting tree and the best-fitting model schemes selected for each dataset (as selected above). The ultrafast bootstrap approximation (UFBoot) method (Hoang et al. 2018) was implemented using 5000 replicates and a minimum correlation coefficient of 0.99. To ensure accuracy, the analysis was run twice to confirm that independent ML searches recovered the same topologies. Bayesian inference (BI) was run using MrBayes v.3.2.7a (Ronquist et al. 2012) on the CIPRES Science Gateway XSEDE (http://www.phylo.org; Miller et al. 2010) using the gene-partitioned scheme and model selection identified by ModelFinder implemented in IQ- Conradie W et al.: Systematics of African rough-scaled lizards 630 Table 1. Material used for the phylogenetic analyses, including sample number, museum catalogue number, country, locality, geographic coordinates, and GenBank/ENA accession numbers. Abbreviations: Aaron M. Bauer field numbers (AMB), British Museum of Natural History (BMNH), California Academy of Sciences (CAS), Colecção Herpetológica do Lubango (CHL), Chad Keates field series (CKD), Enviro-Insight (EI), Fundação Kissama Collection (FKH). Krystal Tolley field numbers (KTH, RSP, WP), Marius Burger field numbers (MBUR), Museu de História Natural e da Ciência - Universidade do Porto (MHNCUP), Ninda Baptista field numbers (NB), Museu de História Natural de Maputo (MHNM), National Museum Namibia (NMNW), Museo delle Scienze di Trento (MUSE), Pedro Vaz Pinto field numbers (P, L series), Port Elizabeth Museum (PEM), Reuben V. van Breda field numbers (RE), Sebastian Kirchof field numbers (SK), Stuart V. Nielsen field numbers (SVN), Thomas Branch field numbers (TB), Werner Conradie field numbers (WC, ANG), William R. Branch field numbers (WRB). Missing data or unavailable information is indicated as NA. Sample No. Museum catalogue No. Species Country Locality Latitude Longitude 16S ND4 RAG-1 c-mos WC-4515 PEM R23530 Ichnotropis b. bivittata Angola West of Cuito town on Aludungo road −12.3278 16.9067 PV357721 OZ347967 PV412867 NB0675 CHL0675 Ichnotropis b. bivittata Angola Luando Integral Nature Reserve −10.2772 16.9533 PV357719 PV412839 PV412865 PV390641 P1-318 FKH-0833 Ichnotropis b. bivittata Angola Cambau −10.1048 15.2182 PV357720 PV412840 PV412866 PV390642 KTH09-075 PEM R17934 Ichnotropis b. pallida Angola 7 km East of Humpata −14.9820 13.4352 HF547775 HF547731 HF547694 ABC2 NA Ichnotropis capensis Namibia Katima Mulilo −17.5066 24.2688 JX962898 JX963023 JX962916 AMB-6001 NMNW Ichnotropis capensis Namibia Road to Tsumkwe −19.4600 19.7200 DQ871148 HF547732 DQ871206 AMB-6067 CAS 209602 Ichnotropis capensis South Africa Kosi Bay, KwaZulu-Natal −26.9400 32.8200 DQ871149 HF547733 DQ871207 ANG-311 PEM R20495 Ichnotropis capensis Angola 8.5 km North of Rito −16.6232 19.0535 PV357725 PV412844 OZ347936 BMNH 2019. 2745 BMNH 2019. 2745 Ichnotropis capensis Zambia Sioma Ngwezi National Park −16.8987 23.5985 MK464418 BMNH 2019. 2746 BMNH 2019. 2746 Ichnotropis capensis Zambia Chavuma Farm −13.0701 22.9288 MK464417 BMNH 2019. 2747 BMNH 2019. 2747 Ichnotropis capensis Zambia Lukwakwa −12.6608 24.4370 MK464416 BMNH 2019. 2750 BMNH 2019. 2750 Ichnotropis capensis Zambia Nanzila Plains, Kafue National Park −16.2814 25.9168 MK464415 BOX23-C02 PEM R24748 Ichnotropis capensis South Africa Tembe Elephant Park, KwaZulu-Natal −27.0217 32.4583 OZ347416 OZ347968 EI-0437 NA Ichnotropis capensis South Africa Lepalale, Limpopo −23.6391 27.5981 OZ347969 OZ347937 OZ347957 EI-0443 PEM R25370 Ichnotropis capensis South Africa Lepalale, Limpopo −23.6391 27.5981 OZ347970 OZ347958 EI-0444 PEM R25371 Ichnotropis capensis South Africa Lepalale, Limpopo −23.6391 27.5981 OZ347417 OZ347971 OZ347938 OZ347959 L-18 PEM R22069 Ichnotropis capensis Angola Gambos, Foster‘s farm −15.8500 14.6833 PV357726 PV412845 OZ347939 MOZ14-356 NHNM Ichnotropis capensis Mozambique Chizavane, Zona Braza Lodge −25.0137 34.0376 OZ347418 OZ347972 OZ347940 MOZ14-357 PEM R21112 Ichnotropis capensis Mozambique Chizavane, Zona Braza Lodge −25.0137 34.0376 OZ347419 OZ347973 OZ347941 MOZ14-358 PEM R21113 Ichnotropis capensis Mozambique Chizavane, Zona Braza Lodge −25.0137 34.0376 OZ347420 OZ347974 OZ347942 OZ347960 NB0771 CHL0771 Ichnotropis capensis Angola Bicuar National Park −15.2435 14.8915 PV357727 PV412846 PV412870 PV390644 NB0772 CHL0772 Ichnotropis capensis Angola Bicuar National Park −15.2435 14.8915 PV357728 PV412847 PV412871 PV390644 NB0779 CHL0779 Ichnotropis capensis Angola Bicuar National Park −15.1049 14.8403 PV357729 PV412848 PV412872 PV390644 NB1116 CHL1116 Ichnotropis capensis Angola Cusseque −13.6851 17.0795 PV357730 PV412849 OZ347943 PV390647 NB1123 CHL1123 Ichnotropis capensis Angola Cusseque −13.6782 17.0832 PV357732 PV412851 PV390649 NB1124 CHL1124 Ichnotropis capensis Angola Cusseque −13.6782 17.0832 PV357733 PV412852 NB1138 CHL1138 Ichnotropis capensis Angola Cusseque -13.6782 17.0832 PV357734 PEM R15556 PEM R15556 Ichnotropis capensis Mozambique 7 km North of Chibuto −24.6231 33.5661 OZ347975 RE211206B1 NMNW R11561 Ichnotropis capensis Namibia Khaudum −18.2876 20.9897 OZ347421 OZ347976 OZ347944 RE211206B3 NMNW R11562 Ichnotropis capensis Namibia Khaudum −18.2876 20.9897 OZ347977 OZ347945 SK13128 NA Ichnotropis capensis Namibia Naye-Naye −19.9235 20.6976 MN015330 MN030223 Vertebrate Zoology 75, 2025, 627–672 631 Sample No. Museum catalogue No. Species Country Locality Latitude Longitude 16S ND4 RAG-1 c-mos WC-3660 PEM R22021 Ichnotropis capensis Zambia Ngonye Falls −16.6736 23.5969 OZ347422 OZ347978 OZ347946 OZ347961 WC-6797 PEM R27394 Ichnotropis capensis Angola Quembo River bridge camp −13.5275 19.2806 PV357731 PV412850 PV412873 PV390648 WC12-A191 PEM R20009 Ichnotropis capensis Angola HALO Cuito Cuanavale office −15.1392 19.1436 PV357724 PV412843 OZ347947 WP031 NA Ichnotropis capensis Namibia 50 km North of Gobabis −22.0000 19.1400 HF547734 HF547695 WRB INH2 NA Ichnotropis capensis Mozambique Inhassoro −21.7143 35.2103 OZ347423 OZ347979 OZ347948 WC-4585 PEM R23525 Ichnotropis capensis Angola Quembo River source −13.1095 19.0061 PV357722 PV412841 PV412868 PV390643 WC-4618 PEM R23500 Ichnotropis capensis Angola Quembo River source −13.1360 19.0453 PV357723 PV412842 PV412869 RE211206D1 NMNW R12212 Ichnotropis grandiceps Namibia Khaudum −18.2876 20.9897 OZ347424 OZ347980 P3-059 MHNCUP-REP 0983 Ichnotropis longicorpa sp. nov. Angola Cuemba −12.1707 18.2257 PV357739 PV412857 OZ347962 P9-035 MHNCUP-REP 0984 Ichnotropis longicorpa sp. nov. Angola Mona Quimbundo −10.0583 19.8056 PV357737 PV412855 OZ347949 OZ347963 TB-44 PEM R19903 Ichnotropis longicorpa sp. nov. Angola Camp Chiri, Miombo forest/camp −9.3969 20.4319 PV357735 PV412853 PV412874 TB-46 PEM R19905 Ichnotropis longicorpa sp. nov. Angola Camp Chiri, Miombo forest/camp −9.3969 20.4319 PV357738 PV412856 PV412876 WC-4557 PEM R23409 Ichnotropis longicorpa sp. nov. Angola Lungwebungu River camp bridge crossing −12.5835 18.6660 PV357741 PV412859 PV412877 WC-4560 PEM R23531 Ichnotropis longicorpa sp. nov. Angola Sombanana village river −12.3071 18.6235 PV357736 PV412854 PV412875 OZ347964 WC-6291 PEM R23996 Ichnotropis longicorpa sp. nov. Angola Lake Tchanssengwe −12.4140 18.6442 PV357740 PV412858 OZ347950 P0-44 MHNCUP-REP 0983 Ichnotropis microlepidota Angola Serra do Moco – Canjonde −12.4261 15.1478 PV357742 PV412860 PV412878 PV390651 WC-3969 PEM R23306 Ichnotropis robusta sp. nov. Angola 4 km upstream from Cuanavale River source −13.0508 18.8973 PV357715 PV412835 PV412862 PV390640 WC-3994 PEM R23279 Ichnotropis robusta sp. nov. Angola Cuanavale River source −13.0903 18.8940 PV357718 PV412838 PV412864 WC-4056 PEM R23362 Ichnotropis robusta sp. nov. Angola drive to Cuanavale River Camp from Samanunga village −13.0380 18.8298 OZ347425 OZ347981 OZ347951 WC-4816 PEM R23420 Ichnotropis robusta sp. nov. Angola Cuando River source −13.0035 19.1275 PV357716 PV41283 CKD-457 PEM R28456 Ichnotropis tanganicana DRC Upemba National Park −9.0442 26.9966 OZ347426 OZ347982 OZ347952 CKD-432 PEM R28448 Ichnotropis tanganicana DRC Upemba National Park −9.0442 26.9966 OZ347427 OZ347983 OZ347953 OZ347965 CKD-433 PEM R28449 Ichnotropis tanganicana DRC Upemba National Park −9.0442 26.9966 OZ347428 OZ347984 OZ347954 OZ347966 CKD-442 PEM R28452 Ichnotropis tanganicana DRC Upemba National Park −9.0442 26.9966 OZ347429 OZ347985 OZ347955 MTSN 9947 MUSE-VER 09947 Ichnotropis tanganicana DRC Kindingi, West of Kabobo Plateau −5.2626 28.9076 OZ347430 OZ347956 ABH3 Meroles squamulosus Mozambique unknown JX962896 EF632221 EF632266 ABH9 Meroles squamulosus Tanzania Laela −8.7500 32.1833 JX962897 JX963022 JX962915 MBUR00872 Meroles squamulosus South Africa Cleveland, Limpopo −24.0219 31.1991 LT745784 LT745812 LT745838 RSP373 Meroles squamulosus South Africa Venetia Limpopo Reserve, Limpopo −22.2661 29.3329 HF547777 HF547737 HF547699 SVN362 PEM R19626 Meroles squamulosus South Africa Lapalala Game Reserve, Limpopo Landmanslust, Limpopo −23.8759 28.3061 HF547776 HF547736 HF547697 WP125 Meroles squamulosus South Africa Rooipoort Nature Reserve Northern Cape −28.5937 24.2100 HF547778 HF547738 HF547701 Conradie W et al.: Systematics of African rough-scaled lizards 632 TREE. Two parallel runs of the MCMC were run for 20 million generations, each with four independent chains, and trees were sampled every 1000 generations using BEAGLE (Ayres et al. 2019). A burn-in of 20% was used to generate the consensus tree. Tracer v.1.7.2 (Rambaut et al. 2018) was used to assess the effective sample size (ESS) for the run parameters. The ESS values were found to be above 200, indicating that the burn-in was adequate. Both the ML and BI trees were generated using FigTree v.1.4.4 (Rambaut 2018). Nodes with bootstrap support (BS) ≥ 95% for ML analyses as well as posterior probabilities (PP) ≥ 0.95 for the BI analyses (Huelsenbeck and Rannala 2004) were regarded as well supported. Species delimitation analyses were performed to explore species boundaries and elucidate whether there was potential cryptic diversification within Ichnotropis. Mitochondrial genes (16S, ND4) were combined for species delimitation analyses, excluding the outgroup taxa, and the sequences were trimmed to 1134 bp to minimise missing data in the datasets. Several different delimitation analyses were run: Automatic Barcode Gap Discovery (ABGD), Assemble Species by Automatic Partitioning (ASAP), Poisson Tree Processes (PTP), Multi-rate Poisson Tree Process (mPTP), and Bayesian Poisson Tree Processes (bPTP). Alignments were prepared and uploaded onto the ABGD Web Interface (https://bioinfo. mnhn.fr/abi/public/abgd/abgdweb.html, web version 22 May 2023) and the ASAP Web Interface (https://bioinfo.mnhn.fr/abi/public/asap/asapweb.html, web version 22 May 2023) as FASTA files. For ABGD, the following settings were used: Standard pairwise distance (p distance) metrics, minimum barcode gap width (1), intraspecific divergence minimum (0.001) and maximum (0.1) (Puillandre et al. 2012). For ASAP, the Simple Distance (p distance) substitution model was used (Puillandre et al. 2021). Multi-locus ML phylogenies were created for the two concatenated mitochondrial genes in IQ-TREE as outlined above. The phylogenies were rendered as unrooted nexus files and uploaded onto the bPTP web server (http://species.h-its.org/ptp; Zhang et al. 2013) for the PTP and bPTP analyses, and the mPTP analysis was conducted using the web server at http://mptp.h-its.org/#/ tree (Kapli et al. 2016). Uncorrected pairwise distances (p distances) were estimated in MEGA X (Kumar et al. 2018) for mitochondrial genes (16S, ND4). Sequences were trimmed to minimise missing data in the datasets and p distances were estimated using uniform rates, pairwise deletion of remaining data, and 500 bootstrap replicates. Morphology We examined all newly collected material in the collections of the National Museum of Namibia (NMNW), Windhoek, Namibia, and Port Elizabeth Museum (PEM). Additional morphological data were gathered from the following museum collections: PEM, Ditsong National Museum of Natural History, Pretoria (TM); Musée Royal de l’Afrique Centrale, Tervuren, Belgium (RMCA); and Institut Royal des Sciences Naturelles de Belgique, Bruxelles, Belgium (IRSNB). The morphological dataset was further supplemented with data from primary literature (Boulenger 1921; Loveridge 1933; de Witte and Laurent 1942; Laurent 1952, 1964; Broadley 1967b; Haacke 1970) and unpublished data of D.G. Broadley and W.R. Branch. The final dataset incorporates morphological data from the type specimens of: Ichnotropis bivittata, I. capensis nigrescens, I. chapini, I. macrolepidota, I. microlepidota, and I. overlaeti. Additionally, high-resolution images of the type specimens of I. longipes and I. tanganicana were consulted, as well as key specimens in the Dundo Museum, Angola (DM) and Museum of Comparative Zoology, USA (MCZ). List of material examined can be found in the Appendix. This represents all the relevant type material, except for the type of Ichnotropis capensis, which remains unaccounted for in the Natural History Museum, London (BMNH). The available material enables us to confidently assign our specimens to known species and to make informed taxonomic decisions. Scale nomenclature, scale counts, and measurements used in the descriptions follow previous studies on African Lacertidae (Conradie et al. 2012; Branch et al. 2019; Parrinha et al. 2021; Benito et al. 2025), and were adjusted as needed to address the morphology of Ichnotropis. The following measurements were taken in millimetres (mm) using a digital calliper (accuracy of 0.01 mm) with the aid of a Nikon SMZ1270 microscope: Snout–vent length (SVL, tip of the snout to the posterior edge of the cloaca); tail length (TAIL, tip of tail to posterior edge of the cloaca, measured only for specimens with complete original tails); total length (TL, combined SVL and tail length); head length (HL, from the anterior edge of the occipital scale to the tip of the snout); head width (HW, widest part of head –usually measured just behind the eye); head height (HH, measured just behind the eyes); snout to front of arm (S-FL, from tip of snout to anterior insertion of forelimb); eye diameter (ED, from top anterior to the posterior edge of eye); snout to eye distance (SE, from tip of snout to anterior edge of eye); eye-to-eye distance (EE, from anterior edge of one eye to anterior edge of the other eye); tympanum length (Tymp-L, at its widest part vertically); lower jaw length (LJL, anterior edge of the jaw bone to tip of lower jaw); inter-limb length (ILL, distance between axillary and inguinal regions); forelimb length (FLL, from elbow to wrist); hind limb length (HLL, from knee to heel); hind foot length (HFL, from ankle to tip of fourth toe, excluding claw); fourth finger length (FFL, excluding claw), fourth toe length (FTL, excluding claw), length of anterior supraocular scale (SO), distance between anterior supraocular to second loreal (SO-L, measurement between the closest point of the anterior supraocular to the posterior edge of the second loreal), frontal scale width (FNW, at its widest point), and frontal scale length (FNL). All measurements, except for EE, FNW, FNL, were taken on the right side of the body. The following scalation details were recorded with the aid of a dissecting microscope: Number of supralabials (SL, anterior to the subocular); number of infralabials (IL); the number and condition of the nasal scales; the Vertebrate Zoology 75, 2025, 627–672 633 number and condition of the loreals; scalation condition of the lower eyelids; number of supraciliaries (SC); the condition of temporal scales and the elongate temporal plate (scales between parietals and temporal scales); the degree of head striations (ridges) present on the dorsal head; the condition of the tympanum opening and the tympanic shield; the condition of the frontonasals (FN), prefrontals (PF, whether the PF is in contact with anterior SO and if the PF is in contact with 1st SC), frontal (F), interparietal (IP), parietals (P) and occipital scales (O); number of supraoculars (SO) [note: for this study we restrict the supraoculars to the two larger scales and refer to the cluster of 1–4 smaller scales posterior to the 2nd SO as the post-supraoculars]; number of smaller scales in front of the anterior SO, touching frontal, prefrontal and 1st SC; number of granules in contact with the two large SO and SC; number of paired chin shields (CS, and the number in contact); midbody scale rows (MSR); longitudinal ventral scale rows (LVSR, counted midway between foreand hind limbs); transverse rows of ventrals (TVSR, counted from the axilla to the groin); femoral pores (right/left); and subdigital lamellae under the 4th toe (LUFT). To investigate the morphological variation between Ichnotropis species and to compare them with previously published material (Benito et al. 2025), two separate principal component analyses (PCA) were run on adult specimens. We considered specimens to be adults if the SVL was larger than 40 mm, as that was the smallest size at which we could clearly observe the hemipenal bulge in males. First, a PCA was performed on the full dataset, which included all measurement variables (Dataset 1). The initial analysis revealed that head-related measurements (HH, HL, HW) accounted for most of the variation. To determine whether other variables contributed notable variation, a second PCA was run on a reduced dataset that excluded these head measurements (Dataset 2). All the variables were first size-corrected using a linear regression with body size (SVL) as the covariate, and the residuals were used as input variables for the PCA. Variables with communalities > 0.5 were retained in the analysis, a varimax rotation was applied, and vectors with eigenvalues > 1.0 were extracted (Tabachnick and Fidel 2019). The resulting principal component (PC) scores were saved and subsequently used as input for a multivariate analysis of variance (MANOVA), with species as the fixed factor. Differences between species were evaluated post hoc using Tukey’s HSD test. All analyses were conducted in RStudio v.2023.09.1+494 (RStudio Team 2022). In order to explore other potential diagnostic characters between species, we tested the morphological variation in Dataset 1 and 2 across different taxa using permutational ANOVAs (PERMANOVAs) with the package RRPP (Collyer and Adams 2018) implemented in RStudio v.2023.09.1+494 (RStudio Team 2022). Variables were size-corrected (SVL) and log-transformed prior to the analyses to mitigate the effects of size and multicollinearity. Finally, standard boxplots were used to visually represent the variables that were significantly different between species. Mapping To enable production of contemporary geographic distribution maps for all Ichnotropis species, observation locations were sourced from published datasets (e.g., van den Berg 2017; Marques et al. 2018), museum databases (PEM, TM, RMCA, IRSNB), and other citizen science repositories (http://www.inaturalist.org; http://vmus.adu. org.za [records were download before the website shut down]). Each record obtained from online sources was checked for diagnostic features mentioned in this study to confirm species identifications. Those which could not be confidently identified were excluded from the mapping exercise. However, historical specimens that were not examined, or for which identification could not be verified, were tentatively mapped based on their initial identification in the original publication or museum catalogue. The online GeoNames gazetteer (http://www.geonames. org) or the GEOLocate Web Application (https://www. geo-locate.org/web/WebGeoref.aspx) was used to georeference all historical data lacking precise location information. Finally, all valid observation records were mapped using QGIS v.3.2 (http://qgis.org). Data used for mapping can be found at: https://doi.org/10.6084/m9. figshare.30285421. Results Phylogenetic analyses Both maximum likelihood and Bayesian inference analyses recovered identical topologies for the concatenated dataset (Figs 1, S1, S2), with strong support at most major nodes. Although the mitochondrial and nuclear gene trees produce different topology in the placement of I. microlepidota, these differences were not considered to represent strong phylogenetic conflict (see Figs S3, S4). Our phylogenetic results recovered four distinct clades within Ichnotropis. Ichnotropis tanganicana was consistently supported as sister to all other congeners in all analyses, from which it is highly divergent (Table 2). All species delimitation methods supported its distinct species status and further subdivided it into two lineages (Fig. 1). Ichnotropis microlepidota was recovered as sister to the I. bivittata clade, though this relationship was not strongly supported in the BI analysis. Nevertheless, all species delimitation analyses favoured its specific status, with notably high pairwise p distances for 16S and ND4 genes (Table 2). The I. bivittata clade contained only four samples and amongst them, topotypic material of I. b. pallida (KTH09-075), which exhibited high intraspecific variation. This sample differed markedly from other I. b. bivittata samples, with divergence values comparable to those in the I. capensis group, but lower than between other species (Table 2). Additionally, all species delimitation analyses—except mPTP— Conradie W et al.: Systematics of African rough-scaled lizards 634 identified each I. bivittata lineage as a candidate species (Fig. 1). Ichnotropis grandiceps was recovered as a sister taxon to the I. capensis group. Furthermore, newly collected material from Angola was recovered as a distinct lineage, sister to a sample from Namibia (RE211206D1) collected from near the type locality of I. grandiceps, and with high divergence in both mitochondrial markers (Table 2). All delimitation analyses supported the distinctiveness of the Angolan lineage as a candidate new species (Fig. 1). Within the I. capensis group, high levels of intraspecific variation were detected, with three major clades (Clades 1–3) returned in the phylogenetic analysis. Species delimitation analyses recovered between three and seven candidate species among these clades, which do not necessarily agree with the three major clades. Sequence divergence among Clade 1 with Clades 2 and 3 (collectively referred to as I. capensis sensu lato) was >5% for 16S and ~12% for ND4 (Table 2), comparable to species-level thresholds amongst other African Lacertidae (~2–12% 16S and 7–23% ND4; Conradie et al. 2012; Edwards et al. 2013a; Branch et al. 2019; Parrinha et al. 2021). Furthermore, I. capensis sensu lato was consistently supported as distinct across all species delimitation methods. It should be noted that the intra-specific variation within I. capensis sensu lato, while surprisingly high, was not comparable with species level divergence (Fig. 1; Table 2). Overall, we propose that the genetic evidence, taken with other evidence (see below) suggests that each of these groups can be considered separately evolving metapopulations under the general lineage concept of species (de Queiroz 1998; see below). Morphology The two PCAs produced similar results despite analysing different subsets of original variables. In PCA1 (Dataset 1), PC1 (37.3%) and PC2 (13.11%) together explained 50.41% of the total variation, with PC1 primarily correlated to head-related variables (HL and SE) and PC2 correlated to limb measurements (FTL and HFL; Fig. 2; Table S2). Similarly, in PCA2 (Dataset 2), PC1 (36.24%) and PC2 (11.41%) accounted for 47.65% of the total variation, in which PC1 is also correlated with head morphology (SE and LJL), while PC2 correlated to the same variable as in previous analysis, suggesting consistent underlying morphological patterns regardless of variable inclusion. The MANOVA showed significant differences between the species for only PC2 (P = 0.000) in both PCAs, and for PC1 of PCA2 (P = 0.007; Table S2). Post hoc pairwise comparisons (Tukey’s HSD) revealed no signifTable 2. Mean sequence divergences (uncorrected p distances) between Ichnotropis species for 16S and ND4 genes, given as percentages. The numbers in the diagonal grey boxes represent the mean intraspecific sequence divergences and standard errors, numbers below the diagonal grey boxes represent the mean interspecific sequence divergences, while numbers above the diagonal grey boxes represent standard errors of the interspecific sequence divergences. n/c – was not possible to estimate sequence divergences. 16S 12345678 1I. capensis sensu lato 3.0 ± 0.5 0.9 1.5 1.4 1.4 1.4 1.7 1.6 2I. longicorpa sp. nov. 5.9 2.6 ± 0.8 1.5 1.4 1.3 1.2 1.6 1.6 3I. robusta sp. nov. 11.2 12.0 0.2 ± 0.2 1.1 1.4 1.3 1.7 1.5 4I. grandiceps 11.5 11.8 6.3 n/c 1.4 1.3 1.7 1.6 5I. b. pallida 11.6 10.5 9.9 10.3 n/c 0.9 1.7 1.5 6I. b. bivittata 11.5 9.8 8.8 9.3 5.4 3.8 ± 0.8 1.6 1.4 7I. microlepidota 16.4 16.1 16.4 16.1 14.4 13.9 n/c 1.8 8I. tanganicana 15.8 14.5 12.9 13.8 13.1 12.2 18.4 1.8 ± 0.4 ND4 1I. capensis sensu lato 7.7 ± 0.6 0.8 1.3 1.3 1.4 1.1 1.3 1.2 2I. longicorpa sp. nov. 12.3 6.6 ± 0.6 1.3 1.3 1.4 1.1 1.3 1.2 3I. robusta sp. nov. 22.1 19.6 1.4 ± 0.3 1.3 1.6 1.2 1.5 1.4 4I. grandiceps 22.3 20.4 15.3 n/c 1.6 1.2 1.5 1.4 5I. b. pallida 21.5 20.6 22.3 20.7 n/c 1.1 1.6 1.7 6I. b. bivittata 19.2 17.6 20.0 19.6 12.8 9.4 ± 0.9 1.3 1.3 7I. microlepidota 24.0 21.6 24.4 24.4 22.6 20.6 n/c 1.4 8I. tanganicana 19.1 18.0 21.7 21.8 22.7 19.4 21.2 0.1 ± 0.1 Vertebrate Zoology 75, 2025, 627–672 635 icant differences between species for PC1 (P > 0.05), but significant differences were detected for PC2 in both analyses (Table S3). The main differences detected were between species of different groups (e.g., I. bivittata versus I. capensis; I. capensis versus I. grandiceps) but not within the different groups (Table S3). Figure 1. IQ-TREE maximum likelihood consensus phylogeny for Ichnotropis with likelihood bootstrap support values (above) and Bayesian posterior probabilities (below) indicated at each node (see key in top left). The coloured bars to the right of the phylogeny summarise the results for each of the species delimitation analyses. Inset image: I. capensis sensu stricto. The scale bar represents substitutions/site. Conradie W et al.: Systematics of African rough-scaled lizards 642 Ichnotropis chapini Schmidt, 1919 Chapin’s rough-scaled lizard Figure 8; Table 3 Taxonomic note. 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. Holotype. AMNH 10674, adult female, collected from Aba, Haut-Uele Province, DRC in July 1911. 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 Table 3. Summary of morphological data for the Ichnotropis bivittata group. Measurements are all shown in millimetres (mm). Values are given as a range, with mean ± standard deviation in parenthesis. Juveniles were excluded from the measurements, but were included in the scalation data. For abbreviations see Materials and Methods section. n = sample size. Characters I. bivittata I. chapini I. microlepidota I. tanganicana n = 39 n = 3 n = 6 n = 32 SVL 42.2–75.0 (63.2 ± 8.33) 53.8–58.0 (55.6 ± 2.15) 48.7–52.0 (50.4 ± 1.45) 41.0–60.0 (53.9 ± 4.21) TAIL 85–156 (109.7 ± 15.47) 77 69.8 55.6–107.9 (81.7 ± 12.15) HL 12.1–15.3 (13.6 ± 1.22) 11.7–12.9 (12.2 ± 0.64) 12.6 11.2–14.2 (12.6 ± 0.79) HW 6.0–9.9 (8.3 ± 1.08) 7.0 –8.7 (7.7 ± 0.89) 6.7–7.3 (7.0 ± 0.20) 6.1–8.7 (7.6 ± 0.51) HH 5.4–8.7 (7.2 ± 0.90) 6.5 5.3–6.7 (5.9 ± 0.62) 5.2–8.4 (6.8 ± 0.71) ED 4.1–4.7 (4.4 ± 0.29) 4.2–5.2 (4.7 ± 0.70) 3.6 3.9–4.5 (4.2 ± 0.18) SE 3.9–6.8 (5.6 ± 0.90) 4.9–5.9 (5.4 ± 0.53) 4.3–5.2 (4.7 ± 0.37) 5.1–6.6 (5.8 ± 0.36) LL 14.2–17.2 (15.6 ± 1.6) 11.5 –13.8 (12.7 ± 1.63) 13.3 12.3–16.0 (13.8 ± 1.02) IL 23.6–33.9 (26.7 ± 3.31) 24.3–33.0 (28.8 ± 4.33) 22.5 19.4–29.4 (24.8 ± 3.0) FLL 5.8–9.4 (7.5 ± 1.13) 6.1–6.9 (6.5 ± 0.52) 5.4 4.7–8.7 (6.5 ± 0.80) HLL 9.0–12.4 (10.4 ± 1.28) 9.2–9.5 (9.3 ± 0.23) 7.4 7.9–10.7 (9.2 ± 0.79) TAIL/SVL 1.3–2.4 (1.8 ± 0.28) 1.3 1.4 1.2–2.0 (1.6 ± 0.23) HL/SVL 0.2 (0.2 ± 0.01) 0.2 (0.2 ± 0.02) 0.3 0.2–0.3 (0.2 ± 0.02) ES/HL 0.4–0.5 (0.4 ± 0.02) 0.4–0.5 (0.4 ± 0.02) 0.4 0.4–0.5 (0.5 ± 0.01) HW/HL 0.5–0.7 (0.6 ± 0.05) 0.3–0.7 (0.6 ± 0.10) 0.6 0.6 (0.6 ± 0.03) MSR 29–40 34–35 43–50 28–42 LVSR 8–10 8–10 8 –10 8–10 TVSR 22–31 24–25 26–30 20–27 SL 3–6 (mostly 4) 4–5 (mostly 4) 4 3–5 (mostly 4) IL 6–9 (mostly 6) 6–7 (mostly 6) 6–8 (mostly 7) 5–7 (mostly 6) SC 3–4 (mostly 4) 3–5 (mostly 4) 4 4–5 (mostly 4) LUFT 17–24 18–20 16–19 17–22 Femoral pores 10–14 8–9 10–13 10–15 Vertebrate Zoology 75, 2025, 627–672 643 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 4th 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. 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. Habitat and Natural History. Very little is known about this species, but it is expected to have similar habitat requirements to other Ichnotropis species. 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. Conradie W et al.: Systematics of African rough-scaled lizards 644 Ichnotropis microlepidota Marx, 1956 Mount Moco rough-scaled lizard Figures 9, 10; Table 3 Taxonomic note. 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. Holotype. FMNH 74285, adult male, collected from the ‘foot of Mount Moco’ [= Serra do Moco], Huambo Province, Angola, by Gerd Heinrich on 19 September 1954. Paratypes. FMNH 74283–84 (females), FMNH 74286– 87 (males); same collection details as holotype. 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. Vertebrate Zoology 75, 2025, 627–672 645 Additional material. 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). 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 1st 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 4th 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 Distribution. Currently only known from the slopes of Serra do Moco, in the central Angolan highlands (Fig. 3). Habitat and Natural History. The specimens from the type series were preyed upon by a dark chanting goshawk (Melierax melabates) (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. Ichnotropis tanganicana Boulenger, 1917 Tanzanian rough-scaled lizard Figures 11–13; Table 3 Taxonomic note. 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. 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 Conradie W et al.: Systematics of African rough-scaled lizards 646 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 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. Vertebrate Zoology 75, 2025, 627–672 647 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. 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. Conradie W et al.: Systematics of African rough-scaled lizards 648 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). 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. Synonymy. Ichnotropis overlaeti de Witte & Laurent, 1942: 173 (new synonymy). Holotype. 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. 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 4th 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. Distribution. Known from western Tanzania, south to northern Malawi, and eastward to northern Zambia and southern DRC (Fig. 3). Habitat and Natural History. 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). The Ichnotropis grandiceps group Phylogenetically and morphologically, the I. grandiceps group includes I. grandiceps and a candidate new species from Angola described here, which share the following morphological features: Robust, broad and depressed head; the prefrontal always separated from the anterior supraocular; weak head striations; and uniform brown to red dorsum. This group is restricted to the Kalahari Basin, from central Angola to northern Namibia (Fig. 14). Vertebrate Zoology 75, 2025, 627–672 649 Ichnotropis grandiceps Broadley, 1967 Zambezi rough-scaled lizard Figures 15, 16; Table 4 Taxonomic note. 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. Holotype. 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. Paratypes. NMZB-UM 16278 (male) and USNM 163990 (juvenile); same collection details as holotype. 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 4th 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 14. Records of the Ichnotropis grandiceps group, based on all literature records (open circles), examined material (closed circles) and genetically analysed material (white centres). Respective type localities are indicated by arrows: Ig – I. grandiceps, and Ir – I. robusta sp. nov. Conradie W et al.: Systematics of African rough-scaled lizards 650 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. Table 4. Summary of morphological data for the Ichnotropis grandiceps group. Measurements are presented in millimetres (mm). Values are given as a range with mean ± standard deviation in parenthesis. Data for adults and subadults are presented separately. For abbreviations, see the Materials and Methods section. n = sample size. Characters I. grandiceps I. grandiceps I. robusta sp. nov. I. robusta sp. nov. n = 6 (adults) n = 4 (subadults) n = 5 (adults) n = 11 (subadults) SVL 57.2–77.9 (65.6 ± 7.52) 57.2–77.9 (65.6 ± 7.52) 71.9–78.8 (74.9 ± 3.12) 35.6–51.1 (44.6 ± 5.33) TAIL 103.4–148.0 (124.9 ± 16.21) 38.5–50.2 (44.9 ± 5.97) 121.0–140.0 (133.0 ± 8.29) 68.0–100.9 (88.2 ± 11.76) HL 12.7–18.2 (16.0 ± 2.11 9.9–13.5 (12.0 ± 4.558) 17.8–18.7 (18.3 ± 0.39) 9.5–04.7 (11.9 ± 1.60) HW 7.7–12.0 (9.3 ± 1.97) 5.8–7.9 (6.9 ± 1.08) 11.1–11.8 (11.5 ± 0.31) 5.6–8.5 (7.2 ± 1.07) HH 6.4–9.7 (7.8 ± 1.39) 5.4–6.8 (6.2 ± 0.72) 9.1–9.4 (9.3 ± 0.13) 5.8–6.8 (6.4 ± 0.41) ED 4.9–6.0 (5.2 ± 0.50) 3.0–4.6 (4.0 ± 0.90) 2.9–3.8 (3.2 ± 0.42) 2.3–2.7 (2.6 ± 0.16) SE 6.3–8.2 (7.0 ± 0.85) 4.6–6.0 (5.5 ± 0.73) 6.1–8.4 (8.3 ± 0.12) 5.4–6.0 (5.6 ± 0.25) Vertebrate Zoology 75, 2025, 627–672 651 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. Habitat and Natural History. 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 robusta sp. nov. Robust rough-scaled lizard https://zoobank.org/1E069D01-1F86-488F-9025-DEBDB47BEA00 Figures 17, 18; Tables 4, 5 Chresonymy. Ichnotropis cf. grandiceps – Conradie et al. (2022a: 198); Ichnotropis aff. grandiceps – Benito et al. (2025: 893). Holotype. 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. Paratypes. 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. Additional juvenile material. 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. 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. 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 subCharacters I. grandiceps I. grandiceps I. robusta sp. nov. I. robusta sp. nov. LL 14.3–18.3 (16.1 ± 1.99) 10.5–15.0 (13.3 ± 1.96) 20.5–24.2 (21.6 ± 1.53) 1.1–17.0 (13.8 ± 1.90) IL 24.8–38.9 (30.4 ± 5.99) 19.4–38.9 (26.1 ± 6.9) 33.4–40.1 (36.3 ± 2.38) 18.2–26.0 (22.0 ± 2.77) FLL 6.9–9.4 (7.8 ± 1.11) 4.2–6.5 (5.3 ± 1.15) 7.2–8.6 (8.3 ± 0.62) 3.8–6.4 (5.1 ± 0.89) HLL 10.6–13.7 (11.8 ± 1.35) 7.0–9.0 (8.3 ± 1.13) 12.8– 13.9 (13.4 ± 0.44) 6.5– 10.5 (8.7 ± 1.35) TAIL/SVL 1.3–2.2 (1.9 ± 0.34) 2.0 (2.0 ± 0.02) 1.6–1.9 (1.8 ± 0.17) 1.8–2.1 (2.0 ± 0.09) HL/SVL 0.2–0.3 (0.2 ± 0.01) 0.3 (0.3 ± 0.01) 0.2–0.3 (0.2 ± 0.01) 0.3 (0.3 ± 0.01) ES/HL 0.4–0.5 (0.5 ± 0.03) 0.4–0.5 (0.5 ± 0.02) 0.4–0.5 (0.5 ± 0.01) 0.4–0.5 (0.4 ± 0.00) HW/HL 0.5–0.7 (0.6 ± 0.05) 0.5–0.6 (0.6 ± 0.04) 0.6–0.7 (0.6 ± 0.02) 0.6–0.7 (0.6 ± 0.03) MSR 44–47 43–48 LVSR 10 9–10 TVSR 27–31 26–33 SL 4–5 (mostly 5) 4–6 (mostly 5) IL 5–7 (mostly 6) 5–7 (mostly 6) SC 4–5 (mostly 5) 4–5 (mostly 5) LUFT 20–26 20–26 Femoral pores 8–14 8–14 Conradie W et al.: Systematics of African rough-scaled lizards 658 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 4th 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. 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). Habitat and Natural History. This species prefers arid to mesic savanna habitats. It is a diurnal lizard, actively 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. Vertebrate Zoology 75, 2025, 627–672 659 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 longicorpa sp. nov. Long-bodied rough-scaled lizard https://zoobank.org/2D3A7A69-E837-4542-89C3-D4CC18A2A160 Figures 24, 25; Tables 6, 7 Chresonymy. 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). Holotype. 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. Paratypes. 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. Additional material. 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 WC6267), 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 ProvTable 6. Summary of morphological data for the Ichnotropis capensis group. Measurements are in millimetres (mm). Values are given as a range with mean ± standard deviation in parenthesis. Juveniles were excluded from the measurements, but were included in the scalation data. For abbreviations, see the Materials and Methods section. n = sample size. Characters I. capensis sensu lato I. longicorpa sp. nov. n = 256 n = 12 SVL 40.0–67.8 (54.8 ± 5.11) 62.7–71.2 (66.4 ± 2.37) TAIL 69.5–149.0 (110.4 ± 17.89) 117–160 (137.5 ± 15.94) HL 10.1–14.7 (12.7 ± 0.97) 13.1–17.2 (15.1 ± 1.16) HW 5.6–8.6 (7.2 ± 0.67) 7.8–9.4 (8.7 ± 0.46) HH 4.5–7.5 (5.9 ± 0.62) 6.1–8.5 (7.4 ± 0.61) ED 3.9–6.3 (4.5 ± 0.52) 2.4–5.9 (4.4 ± 0.98) SE 4.0–7.0 (5.2 ± 0.77) 6.0–7.3 (6.9 ± 0.40) LL 12.5–16.4 (14.2 ± 1.18) 14.4–18.7 (17.8 ± 1.27) IL 18.5–31.6 (25.5 ± 3.29) 27.4–31.6 (28.7 ± 1.42) FLL 5.0–8.2 (6.2 ± 0.70) 6.8–8.4 (7.5 ± 0.46) HLL 7.4–13.2 (10.5 ± 1.25) 10.6–13.5 (12.3 ± 0.96) TAIL/SVL 1.2–2.7 (2.0 ± 0.28) 1.9–2.4 (2.2 ± 0.22) HL/SVL 0.2–0.3 (0.2 ± 0.01) 0.2–0.3 (0.2 ± 0.01) ES/HL 0.4–0.5 (0.5 ± 0.02) 0.4–0.5 (0.4 ± 0.02) HW/HL 0.2–0.7 (0.6 ± 0.06) 0.5–0.6 (0.6 ± 0.03) MSR 25–42 34–41 LVSR 8–10 9–10 TVSR 20–31 25–31 SL 3–6 (mostly 4) 4–5 (mostly 4) IL 5–8 (mostly 6) 6–7 (mostly 6) SC 3–5 4 LUFT 16–26 19–24 Femoral pores 6–15 10–13 Conradie W et al.: Systematics of African rough-scaled lizards 660 ince, 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. 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. 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 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. Vertebrate Zoology 75, 2025, 627–672 661 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). 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 Table 7. Measurements (in mm) and scale counts for the type series of Ichnotropis longicorpa sp. nov. Scale counts are given as Right/Left. For abbreviations, see the Materials and Methods section. t = truncated. Catalogue No. PEM R23410 PEM R23409 PEM R23502 PEM R23505 PEM R23506 PEM R23507 PEM R23531 Type Status Holotype Paratype Paratype Paratype Paratype Paratype Paratype Sex Male Male Male Male Male Male Female SVL 67.7 63.0 64.3 63.9 65.4 66.2 63.1 TAIL 160.0 146.0 149.0 92t 127.0 148.0 117.0 HL 15.6 15.8 15.6 14.7 15.3 15.8 13.1 HW 8.5 8.6 9.4 8.9 8.9 9.2 7.8 HH 7.1 7.3 7.5 7.3 7.1 7.5 6.1 ED 3.1 3.6 3.6 2.4 2.8 3.8 3.0 SE 6.6 6.6 6.7 6.7 6.6 7.2 6.1 LL 17.6 18.0 18.1 16.8 17.3 18.7 15.2 IL 30.8 31.4 29.0 28.9 31.2 29.5 31.6 FL L 7.4 7.8 8.2 7.0 7.5 7.3 6.8 HLL 13.5 13.1 12.6 13.3 13.5 13.2 11.5 TAIL/SVL 2.4 2.3 2.3 1.9 2.2 1.9 HL/SVL 0.2 0.3 0.2 0.2 0.2 0.2 0.2 ES/HL 0.4 0.4 0.4 0.5 0.4 0.5 0.5 HW/HL 0.5 0.5 0.6 0.6 0.6 0.6 0.6 MSR 39 37 38 40 37 40 37 LVSR 9 9 9 9 9 10 9 TVSR 30 28 29 31 28 30 28 SL 4/4 4/4 4/4 4/5 4/4 4/4 4/5 IL 6/6 7/6 6/6 7/7 6/6 6/6 6/6 SC 4/4 4/4 4/4 4/4 4/4 4/4 4/4 LUFT 22 20 21 22 24 22 22 Femoral pores 12/10 10/10 12/12 12/11 13/12 11/10 10/10 Conradie W et al.: Systematics of African rough-scaled lizards 662 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 foreand 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 4th toe; caudal scales strongly keeled above and below, except those just posterior to the vent, which are smooth. Colouration. (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 foreand 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 supraand 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 supraand 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. Paratype and additional material variation. 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 1st 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 4th 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. 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. Habitat and natural history. 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. Vertebrate Zoology 75, 2025, 627–672 663 Key to the genus Ichnotropis Peters, 1854 1a Snout depressed and pointed, prefrontal separated from anterior supraocular by one or two smaller keeled scales ...2 1b Snout robust and rounded, prefrontal mostly in contact with anterior supraocular ...................5 (I. bivittata group) 2a 43–47 midbody scale rows, body robust, head broad, adult SVL > 70 mm ......................... 3 (I. grandiceps group) 2b 25–42 midbody scale rows, body slender, head narrow, adult SVL < 65 mm ............................. (I. capensis group) 3a Confined to the drier Zambezian Baikiaea woodlands or Combretum-Vachellia bushveld in western Zambia and the Zambezi Region of Namibia and adjacent Botswana; occurs below 1000 m a.s.l ................................I. grandiceps 3b Confined to the Angolan Plateau, which consists of moister Angolan Miombo woodland; occurs above 1300 m a.s.l. ................................................................................................................................................I. robusta sp. nov. 4a Long, slender body and head; dark black spots/blotches on chin shields and gular scales, no clear upper white dorsolateral line separating dorsolateral black band from dorsal brown vertebral band ............... I. longicorpa sp. nov. 4b Short body and head; no dark black spots/blotches on chin shields or gular scales, clear white upper dorsolateral line separating black dorsolateral band from dorsal brown vertebral band ............................. I. capensis sensu lato 5a ≥ 42 (42–50) midbody scale rows, known only from Mt Moco, Angola .......................................... I. microlepidota 5b ≤ 42 (28–42) midbody scale rows ..............................................................................................................................6 6a Presence of dorsolateral blue or yellow spots in life ..................................................................................................7 6b No dorsolateral blue or yellow spots, only known from north-eastern DRC ...............................................I. chapini 7a Prefrontals mostly separated from the anterior supraocular, which is often in contact with the 1st supraciliary; evenly spaced blue dorsolateral spots; occurs in southern DRC, Zambia, Malawi and Tanzania .................. I. tanganicana 7b Prefrontal mostly in contact with anterior supraocular, which is always in contact with the 1st supraciliary; closely spaced yellow spots above arm; occurs in Angola, eastern DRC, Republic of the Congo and Gabon ....... I. bivittata Discussion Although Ichnotropis species are locally abundant and widely distributed across sub-Saharan Africa, excluding West Africa, the genus remains among the most taxonomically neglected of African lacertids. This historical oversight is likely driven by a combination of strong seasonal activity patterns, which limit detectability outside of peak reproductive periods (Broadley 1967a, 1974, 1979; Jacobsen 1987; this study), and a legacy of taxonomic confusion dating back over a century (Boulenger 1921; Laurent 1952, 1964; van den Berg 2017; Benito et al. 2025). As a result, specimen collections have remained sparse in many regions, and the group’s true species richness has long been underestimated. Through expanded geographic sampling, covering much of the known ranges of I. bivittata and I. capensis, and the generation of the first genetic data for I. tanganicana and I. grandiceps, we provided the most comprehensive and geographically inclusive phylogenetic framework for Ichnotropis species to date. Species delimitation analyses, combined with a substantially enhanced morphological dataset, support the validity of most currently recognised species (with the exception that we could not validate the phylogenetic status of I. chapini) and justify the formal description of two new Angolan endemics: Ichnotropis robusta sp. nov. and I. longicorpa sp. nov. These findings underscore the continued importance of integrating molecular and morphological approaches in taxonomic revisions of morphologically conservative lineages. Conversely, our phylogenetic framework also facilitated the reassessment of several taxonomically ambiguous species and subspecies, many of which had not previously been evaluated using phylogenetic analyses. For instance, although I. b. pallida exhibits notable genetic divergence from other I. bivittata populations, we conservatively synonymise the subspecies with the nominal form until further data become available. Furthermore, based on our findings, we recommend retaining I. longipes and I. macrolepidota as synonyms of I. capensis sensu lato, due to the absence of consistent diagnostic morphological differences and inclusion of topotypic material in our phylogenetic analyses. Additionally, we propose synonymising I. c. nigrescens with I. bivittata, and treating I. overlaeti as a junior synonym of I. tanganicana, based on morphological congruence. Newly collected Ichnotropis material from west of the Kabobo Plateau and Upemba National Park in DRC allowed us to reassess the poorly known I. tanganicana, previously known only from its type specimen and a vague type locality (Boulenger 1917; Meiri et al. 2018). Our data confirm the species’ taxonomic validity, help revise the species description and suggest a broader geographic distribution than previously recognised. As a result, most historical records of I. bivittata from East Africa are here reassigned to I. tanganicana. While there was a more comprehensive sampling for I. capensis relative to I. bivittata, the limited material available for I. bivittata (n = 4, including topotypic I. b. pallida) exhibited moderate genetic divergence, and species delimitation analyses consistently identified these lineages as distinct. Given the broad unsampled distribution of I. bivittata, coupled with the high genetic diversity observed within the available dataset, increased geographic sampling, especially within topographically Conradie W et al.: Systematics of African rough-scaled lizards 664 complex Angola, DRC, Republic of the Congo and Gabon, may reveal additional cryptic diversity. Moreover, our species delimitation analyses recovered multiple putative candidate species within the I. capensis sensu lato complex. It must be noted that some of these lineages may correspond to previously described species, such as I. longipes, but resolving this will require targeted sampling from type localities or museomics (i.e., historical DNA [hDNA]) to assess potential synonymy or revalidation (McGuire et al. 2018; Raxworthy and Smith 2021; Zacho et al. 2021; Lalueza-Fox 2022; Letsch et al. 2025). Although we were unable to assess the phylogenetic placement of I. chapini due to the absence of genetic data, we recommend its provisional assignment to the I. bivittata group based on similarities in head morphology and scalation. Until further material becomes available for molecular analysis, we propose retaining I. chapini as a valid species. The species delimitation analyses confirmed the species status of both newly described and previously recognised taxa, less conservative methods also revealed potential cryptic diversity within I. tanganicana, I. bivittata, I. longicorpa sp. nov., and I. capensis sensu lato. We caution that single-locus approaches can overestimate species boundaries by conflating intraspecific variation with interspecific divergence, especially under conditions of incomplete lineage sorting or limited geographic sampling (e.g., Carstens et al. 2013; Sukumaran and Knowles 2017). To mitigate this risk, we employed five independent species delimitation methods, enabling cross-validation and allowing us to conservatively interpret only those lineages supported by multiple lines of evidence (morphology, colouration, ecology) as candidate species (e.g., Fujisawa and Barraclough 2013; Zhang et al. 2013; Kapli et al. 2016). Morphological analyses broadly support the three major clades recovered by the phylogenetic analyses, with diagnostic differences in head shape, scalation, and breeding colouration. These traits are largely consistent across multiple populations, making them valuable for species identification in the field. The I. capensis, I. grandiceps and I. bivittata groups exhibit distinctive morphological characteristics that aid in distinguishing them from each another, even in the absence of genetic data. However, I. tanganicana is an exception: It forms a deeply divergent mitochondrial lineage that is sister to all other Ichnotropis species in the phylogeny; nevertheless, it remains morphologically similar to all members of the I. bivittata group, being thus included as part of this group. The recognition of I. tanganicana as a distinct species is supported primarily by molecular divergence, unique geographic distribution, and its distinctive colouration – most notably the presence of evenly spaced blue lateral spots – underscoring the importance of integrating genetic and phenotypic data in resolving cryptic diversity within morphologically conservative lineages. Biogeographically, Ichnotropis shows its highest diversity across the northern and western parts of central and southern Africa, with the Kalahari Basin emerging as a hotspot for species richness for this group. In contrast, eastern Africa is represented by only a single species (i.e., I. tanganicana), while South Africa harbours just one taxon from the I. capensis complex. These patterns are shaped by regional habitat heterogeneity, historical barriers to gene flow, and likely also by limited historical sampling in large swathes of suitable habitat (Greenbaum et al. 2018). The description of two new Ichnotropis species from Angola contributes to the wave of reptile species discoveries in the region over the past two decades (Conradie et al. 2012, 2020a, 2020b, 2020c; Stanley et al. 2016; Branch et al. 2019, 2021; Marques et al. 2019a, 2019b, 2020, 2022a, 2022b, 2023a, 2023b, 2024; Ceríaco et al. 2020b, 2020c, 2020d, 2024; Hallermann et al. 2020; Lobón-Rovira et al. 2021, 2022, 2025a, 2025b; Parrinha et al. 2021, 2025a, 2025b, 2025c; Wagner et al. 2021; Bates et al. 2023; Röll et al. 2024). Since 2012, Angola’s known lacertid diversity has nearly doubled, from 10 to 18 species, mirroring growth in other reptile groups such as skinks and geckos (Conradie 2024). Given that large portions of Angola remain poorly surveyed, further species discoveries are likely to occur within the territory. The two new Ichnotropis species described here were discovered through intensive fieldwork in central and southeastern Angola, regions long underexplored due to decades of civil conflict and the difficult access to these areas (Conradie et al. 2021). Recent herpetological surveys in these areas have yielded numerous new species, expanded faunal records, and have provided comprehensive species inventories (e.g., Conradie et al. 2016, 2020a, 2020b, 2021, 2022a, 2022b, 2023; Nielsen et al. 2020), underscoring the importance of targeted fieldwork and systematic biodiversity assessments in historically inaccessible areas (Tolley et al. 2016). These two newly described Ichnotropis species (I. robusta sp. nov. and I. longicorpa sp. nov.) are currently considered Angolan endemics, though they may also occur in adjacent regions of DRC and Zambia. Finally, the Angolan highlands and adjacent Miombo woodlands appear particularly significant, harbouring several endemic or near-endemic taxa (Bauer et al. 2023; Becker et al. 2023). These findings align with previous research identifying this region as a hotspot of reptile endemism and support calls for enhanced conservation attention. Conclusion This study presents the most comprehensive phylogenetic and taxonomic revision of the genus Ichnotropis to date. By integrating mitochondrial and nuclear molecular data, detailed morphological assessments, and broad geographic sampling, we reveal that Ichnotropis lizards harbour more diversity than previously recognised. Our findings support the description of two new taxa, indicate multiple potential cryptic species and clarifies the taxonomic status of several historically ambiguous names. The recovered phylogenetic structure and strong geographic Vertebrate Zoology 75, 2025, 627–672 665 partitioning underscore the evolutionary significance of central and southern Africa, particularly the Angolan highlands, as a centre of diversification and endemism for reptiles. This work not only stabilizes the taxonomy of Ichnotropis, but also lays a robust foundation for future evolutionary, ecological and conservation studies for this group across its range. Acknowledgements This work was made possible through the collaboration and support of numerous individuals and institutions. We thank the Ministry of Environment of the Republic of Angola (MINAMB), particularly Dr. Miguel Xavier, Director of the Instituto Nacional de Biodiversidade e Conservação (INBC), for their institutional collaboration. Material was collected and exported under the following permits issued by MINAMB: 31/ GGPCC/2016 and 151/INBAC/MINAMB/2019. We also acknowledge the exceptional logistical assistance provided by Fundação Kissama, especially Vlady Russo. WC thanks the Wild Bird Trust, which administers the National Geographic Okavango Wilderness Project (2015–2019 National Geographic Society grant), and Chris Brooks, who organized the SAREP Aquatic Biodiversity Surveys of the upper Angola catchments of the Cubango-Okavango River Basin (May 2012) and the lower Cuito and Cuando River Basins (April 2013). We are also grateful to the Natural History Museum of Maputo, which endorsed and provided permits (315/MHN/E.27/2014) to carry out part of this work in Mozambique. This work received financial and logistical support from several institutions: The National Geographic Society Committee for Research and Exploration (CRE 9281-13); the South African National Biodiversity Institute; the National Research Foundation of South Africa (Grant #92776) for the 2014 Mozambique survey; and Khangela Safaris for camp logistics in 2014. CK thanks and acknowledges Upemba National Park, Forgotten Parks Foundation (DPF), Institut Congolais pour la Conservation de la Nature (ICCN), University of Lubumbashi, and Hankuzi Explorations for their assistance with facilitating the collection of valuable Congolese samples for this study. JLR is currently supported by Associação BIOPOLIS CIBIO Base FUI 2020–2023 (UIDB1 50027 i2020). This work was also partially supported by the Synthesis+ BE-TAF Project 2022 Grant obtained by JLR. We thank the CTM staff at CIBIO – especially Susana Lopes, Sofia Mourão, and Patrícia Ribeiro – for their dedicated laboratory support. EG acknowledges Ana Betancourt of the Border Biomedical Research Center (BBRC) Genomics Analysis Core Facility for technical services and facilities. This work was supported by Grant 5U54MD007592 from the National Institute on Minority Health and Health Disparities (NIMHD), part of the US National Institutes of Health (NIH). We are grateful to Garin Cael (RMCA) and Olivier Pauwels (RBINS) for providing access to relevant material from their herpetological collections. Photographs of key museum specimens were generously provided by: Eugen Behrens (Museo delle Scienze, MUSE), Lauren Vonnahme (American Museum of Natural History, AMNH), Patrick Campbell (Natural History Museum London, NHML), Joshua Mata (The Field Museum of Natural History, FMNH), Matthew Gage (Museum of Comparative Zoology, Harvard University, MCZ), and Frank Tillack (Museum für Naturkunde Berlin, ZMB). Sebastian Kirchhof (Natural History Museum Abu Dhabi, NHMAD) kindly provided additional sequences. We also thank the following individuals for their assistance in the field and sample collection: Ninda Baptista, Gabriella Bittencourt-Silva, Thomas Branch, William R. Branch, Hanlie Engelbrecht, James Harvey, Timóteo Júlio, Michele Menegon, Götz Neef, and Alex Rebelo. Reuben van Breda extends special thanks to Ed Netherlands, Ash Bullard, Haley Dutton, Bernie Jordaan, Bertha Buiswalelo, Francois Becker, and Louis du Preez for their help in the field, as well as Piet Beytell and Francois Jacobs and their team for facilitating fieldwork in Namibia. Specimens in Namibia were collected under the National Commission of Research, Science and Technology permit AN20191118. We are grateful to the late Don Broadley and Bill Branch for allowing us to incorporate their unpublished data on the genus. References Aljanabi SM, Martinez I (1997) Universal and rapid salt-extraction of high quality genomic DNA for PCR-based techniques. 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