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Review Article
A treasure trove of endemics: two new species of snake-eyed skinks of the genus Panaspis Cope, 1868 (Squamata, Scincidae) from the Serra da Neve Inselberg, southwestern Angola
expand article infoMariana P. Marques‡§|, Diogo Parrinha‡|, Manuel Lopes-Lima‡|, Arthur Tiutenko#, Aaron M. Bauer¤, Luis M. P. Ceríaco‡|«
‡ Universidade do Porto, Vairão, Portugal
§ Carnegie Museum of Natural History, Section of Amphibians and Reptiles, Pittsburgh, United States of America
| Biodiversity and Land Planning, CIBIO, Vairão, Portugal
¶ Departamento de Biologia, Faculdade de Ciências da Universidade do Porto, Porto, Portugal
# Friedrich-Alexander-Universität Erlangen-Nürnberg, Erlangen, Germany
¤ Villanova University, Villanova, United States of America
« Universidade Federal do Rio de Janeiro, Rio de Janeiro, Brazil

Corresponding author: Mariana P. Marques ( mptlmarques@gmail.com )

Academic editor: Oliver Hawlitschek
© 2024 Mariana P. Marques, Diogo Parrinha, Manuel Lopes-Lima, Arthur Tiutenko, Aaron M. Bauer, Luis M. P. Ceríaco.
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.
Citation: P. Marques M, Parrinha D, Lopes-Lima M, Tiutenko A, Bauer AM, Ceríaco LMP (2024) A treasure trove of endemics: two new species of snake-eyed skinks of the genus Panaspis Cope, 1868 (Squamata, Scincidae) from the Serra da Neve Inselberg, southwestern Angola. Evolutionary Systematics 8(2): 167-182. https://doi.org/10.3897/evolsyst.8.121103
ZooBank: urn:lsid:zoobank.org:pub:9BEB46EA-7B8A-4880-B5D7-1163D37F4AE1
Open Access

Abstract

Four species of the genus Panaspis – P. cabindae, P. wahlbergii, P. maculicollis and P. mocamedensis – are currently known from Angola. The analysis of recently collected specimens from Serra da Neve Inselberg, an isolated mountain located in northern Namibe Province, revealed unexpected taxonomic diversity in the group. Using an integrative taxonomy approach based on morphological and DNA sequence data, with both mitochondrial (16S) and nuclear (RAG-1) genes, we were able to distinguish two distinct populations, described here as two new species, Panaspis ericae sp. nov. and P. mundavambo sp. nov. Both species are assumed to be endemic to the inselberg. This reinforces our notion of southwestern Angola as a hotspot of skink diversity, and highlights the urgent need for the conservation of Serra da Neve.

Key Words

Reptiles, integrative taxonomy, Africa, herpetofauna, cryptic species

Introduction

African snake-eyed skinks of the genus Panaspis Cope, 1868, are represented in Angola by four different species: Panaspis aff. breviceps (Peters, 1873), in the escarpment areas of Kwanza Sul and Kwanza Norte provinces; P. cabindae (Bocage, 1866), extending from central Africa to the central highlands of Angola; P. maculicollis Jacobsen & Broadley, 2000, in the south-eastern province of Cuando-Cubango; P. mocamedensis Ceríaco, Heinicke, Parker, Marques & Bauer, 2020, in the southwestern province of Namibe; and P. wahlbergii (Smith, 1849), on the central plateau of Angola (Ceríaco et al. 2020). The systematic placement of the genus has been a matter of debate for decades (Fuhn 1969, 1972; Perret 1973, 1975; Greer 1974; Schmitz et al. 2005; Jesus et al. 2007), until the recent molecular phylogeny by Medina et al. (2016) provided a much-needed clarification. In the last decade, several new species of Panaspis have been described from sub-Saharan Africa: P. thomensis Ceríaco, Soares, Marques, Bastos-Silveira, Scheinberg, Harris, Brehm & Jesus, 2018, from São Tomé Island, in the Gulf of Guinea (Soares et al. 2018); P. namibiana Ceríaco, Branch & Bauer, 2018, from Central Namibia (Ceríaco et al. 2018a); P. tsavoensis Kilunda, Conradie, Wasonga, Jin, Peng, Murphy, Malonza & Che, 2019, from southern Kenya (Kilunda et al. 2019); P. mocamedensis from southwestern Angola (Ceríaco et al. 2020) and P. annettesabinae Colston, Pyron & Bauer, 2020 from central Ethiopia (Colston et al. 2020).

Despite this progress, further cryptic diversity awaiting formal description is expected in the genus. Such cryptic diversity has already been documented by Medina et al. (2016), and further field workers in biodiverse but poorly explored areas, and more are likely to be discovered. One such case deals with two putative new species we recently collected on the Serra da Neve Inselberg in northern Namibe Province, southwestern Angola. Serra da Neve is an isolated mountain of subvolcanic origin with a basal area of approximately 630 km2 and is the second highest peak of Angola at 2489 m (Pereira 1977). Serra da Neve lies in what Mendelsohn and Huntley (2023) define as “the southern escarpment landscape”, an area ranging from the Coporolo River in Benguela Province, Angola, to the Huab River in Kunene Region, Namibia. The inselberg is covered by a Miombo forest habitat, contrasting with the surrounding lowland habitats, which are mainly dominated by Namibian woodland savanna and arid areas of Namib Desert (Grandvaux-Barbosa 1970). Recent fieldwork at Serra da Neve revealed an impressive number of strictly endemic species, such as a bufonid frog of the genus Poyntonophrynus (Ceríaco et al. 2018), three geckos (genera Lygodactylus, Afroedura, and Rhoptropus; Branch et al. 2020; Marques et al. 2020; Parrinha et al. 2024), one cordylid lizard of the genus Cordylus (Marques et al. 2019), and a legless skink of the genus Acontias (Marques et al. 2023a). Moreover, near or regional endemics also occur in the inselberg and its surroundings (Marques et al. 2024). Ceríaco et al. (2020) reported the presence of P. cabindae on the southern slopes of the inselberg, at the locality of Mamué (-13.8015, 13.1206). During a recent survey of the inselberg, several Panaspis specimens were collected both at the top of the mountain and in the surrounding lowlands. Some of these are representatives of two putative new species. Building on the data provided by Medina et al. (2016) and Ceríaco et al. (2020), we here provide an integrative approach to the revision of Serra da Neve’s Panaspis and describe two new taxa. Biogeographic and conservation considerations are also provided.

Materials and methods

Newly collected specimens were euthanized with MS-222 following standard practices (Simmons 2015), fixed in 10% buffered formalin in the field and subsequently transferred to 70% ethanol for long-term storage. Liver tissue was removed before formalin fixation and preserved in 95% ethanol. The specimens were deposited in the collections of the Museu Nacional de História Natural e da Ciência da Universidade de Lisboa, Lisboa, Portugal (MUNHAC). For mensural and meristic comparisons, we used the datasets presented by Ceríaco et al. (2018, 2020) which included Angolan and Namibian Panaspis specimens deposited in the collections at American Museum of Natural History (AMNH), New York, New York, USA; California Academy of Sciences (CAS), San Francisco, California, USA; Carnegie Museum of Natural History (CM), Pittsburgh, Pennsylvania, USA; Field Museum of Natural History (FMNH), Chicago, Illinois, USA; Museum for Comparative Zoology (MCZ), Harvard University, Cambridge, Massachusetts, USA; Florida Museum of Natural History (FLMNH), Gainsville, USA; Natural History Museum of Los Angeles County (LACM), Los Angeles, California, USA; University of Texas El-Paso Herpetological collection (UTEP), El-Paso, Texas, USA; Port Elizabeth Museum/Bayworld (PEM), Port Elizabeth, South Africa; Iziko South African Museum, Cape Town, South Africa (SAM); Ditsong National Museum of Natural History (TM), Pretoria, South Africa; National Museum of Namibia (formerly Staatsmusem Windhoek), Windhoek, Namibia (SMW); Instituto de Investigação Científica Tropical (IICT), Lisbon, Portugal; Museu de História Natural e da Ciência da Universidade do Porto (MHNC-UP), Porto, Portugal; Muséum national d’Histoire naturelle (MNHN), Paris, France; Naturhistorisches Museum (NHMW), Wien, Austria; Senckenberg Forschungsnstitut und Naturmuseum, Frankfurt am Main, Germany (SMF); and the Natural History Museum (BMNH), London, United Kingdom.

Molecular methods

DNA was extracted from tissue samples of newly collected specimens using the Spin Column Animal Genomic DNA Miniprep Kit from Biobasic (Markham, Ontario, Canada), following the manufacturer’s instructions. The mitochondrial 16S rRNA and the nuclear Recombination Activating Protein 1 (RAG1) genes were then amplified following Ceríaco et al. (2020) with the following primers 16S (16SA 5’-CGCCTGTTTATCAAAAACAT-3’; 16SB 5’-CCGGTCTGAACTCAGATCACGT-3’ Palumbi et al. 1996) and RAG1 (RAG1f700 5’-GGAGACATGGACACAATCCATCCTAC-3’; RAG1r700 5’-TTTGTACTGAGATGGATCTTTTTGCA-3’; Bauer et al. 2007). Briefly, genes were amplified in 25 μL PCRs, with an initial denaturing temperature of 95 °C for 2 min, followed by denaturation at 95 °C for 35 seconds (s), annealing at 50 °C for 35 s, and extension at 72 °C for 95 s with 4 s added to the extension per cycle for 32 or 34 cycles (for the mitochondrial or nuclear gene, respectively).The mitochondrial 16S rRNA and the nuclear Recombination Activating Protein 1 (RAG1) genes were then amplified and sequenced for subsequent phylogenetic analyses. The successful amplifications underwent enzymatic purification and were then sequenced using the BigDye Terminator v3.1 Cycle sequencing protocol (Applied Biosystems, Waltham, MA, USA) with an automated Sequencer ABI3500xl Genetic Analyzer at the Centre for Molecular Analysis (CTM) in the Research Centre in Biodiversity and Genetic Resources (CIBIO), Vairão, Portugal. Two datasets, 16S and RAG1, were created using MAFFT software (version 7.304, Katoh and Standley 2013), which included newly sequenced individuals (refer to Table 1 for GenBank Accession numbers) and all previously available sequences of the same genes from Medina et al. (2016) and Ceriaco et al. (2020). The 16S and RAG1 datasets were then concatenated into a single dataset with 1,319 nucleotides (nt) (16S = 671 nt; RAG1 = 648 nt) for phylogenetic analyses. The outgroup used was a combination of those employed by Medina et al. (2016) and Ceriaco et al. (2020) (Table 1). Each partition was tested for substitution saturation using DAMBE 7 (Xia 2018) following the method of Xia et al. (2003). Little saturation was found, even under the assumption of an asymmetric tree (p < 0.001). Incongruence Length Difference (ILD) tests were conducted to investigate incongruence between the two concatenated genes (Farris et al. 1994). The tests showed no significant phylogenetic conflict between the 16S and RAG1 (p = 0.95). Pairwise sequence divergences (uncorrected p-distance) of 16S sequences were calculated using MEGA X. Phylogenies were estimated using Maximum Likelihood (ML) and Bayesian Inference (BI) methods in IQ-TREE v. 2.2.0 (Minh et al. 2020) and Mr Bayes v. 3.2.7a (Ronquist et al. 2012), respectively. The best-fitting nucleotide substitution models for each partition of the subsequent analyses were selected using JModelTest 2.1.10 (Darriba et al. 2012) under the corrected Akaike Information Criterion. The dataset was partitioned by gene and run using the GTR + I + G model for the 16S and GTR + G for the RAG1 partitions. BI analyses were performed using program-generated trees and four Markov chains with default incremental heating. Two independent runs of 15 × 106 generations were sampled at 1,000-generation intervals, resulting in a total of 15,000 trees. A burn-in of 10% was determined by assessing the convergence of log-likelihood and parameter values using Tracer v. 1.7.1 (Rambaut et al. 2018). Phylogenetic analyses were conducted for ML. An initial tree search was followed by 10 independent runs and 10,000 ultrafast bootstrap replicates. Maximum likelihood trees were constructed also for the RAG1 and 16S datasets alone, using IQ-TREE the same parameters as above. The haplotype network reconstruction of the RAG1 dataset was conducted in TCS 1.21 (Clement et al. 2000) with a statistical parsimony limit of 95%.

Table 1.
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Specimens used for genetic analysis and corresponding GenBank accession numbers for genes used in the study. See Materials and Methods section for collection abbreviations. Institution and field number acronyms not cited in the Material and Methods were retrieved from Medina et al. (2016) suppl. material.

Species Collection number Field number Locality Genbank
16S RAG1
Panaspis annettesabinae ZMNH H2019,2176 TJC264 ETHIOPIA: Oromia Region, 8 km SW of Bedele on the Metu-Bedele rd, Buno Bedele zone KU23675 –
Panaspis africanus MUNHAC/MB03-001133 UMRTGGPR-698 / PR357 SÃO TOMÉ AND PRÍNCIPE: Príncipe Island, Montalegre EU164475 DQ675337
Panaspis breviceps ZFMK 87662 MM 105 CAMEROON: Mawne KU236786 KU298714
Panaspis breviceps ZFMK 87663 MM 106 CAMEROON: Mawne KU236787 KU298715
Panaspis cabindae PEM R19467 ANG 21 ANGOLA: Lunda Norte Prov., Lagoa Carumbo KU236741 KU298690
Panaspis cabindae Uncatalogued PM 049 DRC: Luango-Nzambi, Bas-Congo KU236750 KU298697
Panaspis cabindae Uncatalogued PM 050 DRC: Luango-Nzambi, Bas-Congo KU236751 KU298698
Panaspis cabindae PEM R21594 WRB 810 ANGOLA: Bengo Prov., Riverine Forest, Bengo KU236765 KU298705
Panaspis cabindae PEM R20256 WRB 804 ANGOLA: Zaire Prov., Soyo KU236768 KU298708
Panaspis cabindae CAS 263553 AMB 10315 ANGOLA: Namibe Prov., Mamué MN846689 MN850709
Panaspis cabindae FLMNH 187242 – ANGOLA: Namibe Prov., Mamué MN846690 MN850710
Panaspis cabindae MUNHAC/MB03-001088 – ANGOLA: Malanje Prov., Laúca Dam MN846698 MN850711
Panaspis cabindae MUNHAC/MB03-001091 – ANGOLA: Malanje Prov., Laúca Dam MN846699 MN850712
Panaspis ericae sp. nov. MUNHAC/MB03-001525 LMPC 3209 ANGOLA: Namibe Prov. Serra da Neve, Catchi PP810194 PP816726
Panaspis ericae sp. nov. MUNHAC/MB03-001526 LMPC 3210 ANGOLA: Namibe Prov. Serra da Neve, Catchi PP810195 PP816727
Panaspis ericae sp. nov. MUNHAC/MB03-001528 LMPC 3275 ANGOLA: Namibe Prov. Serra da Neve, Catchi, base camp headquarters PP810196 PP816728
Panaspis ericae sp. nov. MUNHAC/MB03-001529 LMPC 3282 ANGOLA: Namibe Prov. Serra da Neve base, 2 km N of Maylowe PP810197 PP816729
Panaspis ericae sp. nov. MUNHAC/MB03-001530 LMPC 3354 ANGOLA: Namibe Prov. Serra da Neve base, 2 km N of Maylowe PP810198 PP816730
Panaspis ericae sp. nov. MUNHAC/MB03-001531 LMPC 3355 ANGOLA: Namibe Prov. Serra da Neve base, 2 km N of Maylowe PP810199 PP816731
Panaspis ericae sp. nov. MUNHAC/MB03-001534 LMPC 3395 ANGOLA: Namibe Prov. Serra da Neve base, 2 km N of Maylowe PP810200 PP816732
Panaspis sp. Katanga 1 UTEP 21175 ELI 295 DRC: Katanga, Mulongo KU236729 KU298685
Panaspis sp. Katanga 1 UTEP 21174 ELI 294 DRC: Katanga, Mulongo KU236730 KU298686
Panaspis sp. Katanga 2 – JHK 26 DRC: Katanga, Kisanfu Camp KU236726 KU298682
Panaspis sp. Katanga 2 PEM R17454 WRB 575 DRC: S Katanga, Kalakundi Copper Mine KU236736 KU298689
Panaspis sp. Katanga 2 – WRBNimb083 ZAMBIA: NW Zambia KU236742 KU298691
Panaspis sp. Limpopo – MCZ-A 27176 SOUTH AFRICA: Limpopo Prov., Hoedspruit KU236743 KU298692
Panaspis maculicollis CAS 234099 MCZF 38733 SOUTH AFRICA: Limpopo Prov., Farm Vrienden KU236720 KU298678
Panaspis maculicollis CAS 234188 MCZF 38848 SOUTH AFRICA: Limpopo Prov., Farm Nooitgedacht KU236728 KU298684
Panaspis maculicollis CAS 234135 MCZF 38790 SOUTH AFRICA: Limpopo Prov., Farm Vrienden KU236747 KU298694
Panaspis maculicollis Uncatalogued MBUR 02843 SOUTH AFRICA: Limpopo Prov., Phalaborwa KU236748 KU298695
Panaspis maculicollis Uncatalogued MBUR 02848 SOUTH AFRICA: Limpopo Prov., Phalaborwa KU236749 KU298696
Panaspis maculicollis PEM R20475 ANG 421 ANGOLA: Cuando Cubango Prov., Benero Campsite, near Jamba KU236770 KU298711
Panaspis mocamedensis MUNHAC/MB03-001532 LMPC 3381 ANGOLA: Namibe Prov. Serra da Neve base, 2 km N of Maylowe PP810201 PP816733
Panaspis mocamedensis MUNHAC/MB03-001533 LMPC 3382 ANGOLA: Namibe Prov. Serra da Neve base, 2 km N of Maylowe PP810202 PP816734
Panaspis mundavambo sp. nov. MUNHAC/MB03-001527 LMPC 3242 ANGOLA: Namibe Prov. Serra da Neve, near Catchi, MPLA cabin PP810203 PP816735
Panaspis sp. Malawi PEM R20247 WRB 568 MALAWI: Mt. Mulanje, Sombani Trail KU236732 KU298687
Panaspis sp. Mozambique 1 – SVN 693 MOZAMBIQUE: Gorongosa National Park KU236754 KU298699
Panaspis sp. Mozambique 1 PEM R20561 WC 1251 MOZAMBIQUE: Sofala Prov, Chemba, Ecofarm KU236764 KU298704
Panaspis sp. Mozambique 1 PEM R20591 WRB 886 MOZAMBIQUE: Tete Prov., Ruoni Hill S KU236769 KU298710
Panaspis sp. Mozambique 4 PEM R20569 WRB 855 MOZAMBIQUE: Cabo Delgado Prov., Balama, Syran graphite mine KU236766 KU298706
Panaspis sp. Mozambique 4 PEM R20576 WRB 856 MOZAMBIQUE: Cabo Delgado Prov., Balama, Syran graphite mine KU236767 KU298707
Panaspis namibiana MCZ R183767 AMB 7634 NAMIBIA: Sesfontein KU236727 KU298683
Panaspis sp. Tanzania 1 – WRB 0026 TANZANIA: Arusha KU236718 KU298676
Panaspis sp. Tanzania 1 – WRB 0021 TANZANIA: Arusha KU236719 KU298677
Panaspis sp. Tanzania 2 PEM R20799 WRB 573 TANZANIA: Serengeti, Klein’s Camp KU236735 KU298688
Panaspis togoensis TCWC 94557 TJH 2629 BENIN: Dogo Forest KU236758 KU298701
Panaspis wahlbergii CAS 234209 MCZF 38868 SOUTH AFRICA: Limpopo Prov. KU236721 KU298679
Panaspis wahlbergii MCZ R184443 AMB 8293 SOUTH AFRICA: Limpopo Prov. KU236722 KU298680
Panaspis wahlbergii CAS 234194 MCZF 38852 SOUTH AFRICA: Limpopo Prov. KU236724 KU298681
Panaspis wahlbergii TM 84299 – SOUTH AFRICA: Limpopo, Groblersdal KU236746 KU298693
Panaspis wahlbergii NMB R10286 SVN 742 MOZAMBIQUE: Sofala Prov., Beira KU236755 KU298700
Panaspis wahlbergii TCWC 95563 TJH 3213 SOUTH AFRICA: Northern Cape, Kimberley KU236759 KU298702
Panaspis wahlbergii TCWC 95588 TJH 3253 SOUTH AFRICA: Northern Cape, Kimberley KU236760 KU298703
Panaspis wahlbergii PEM R21298 WC 2721 SOUTH AFRICA: Mpumalanga, Doornkop Reserve KU236781 KU298712
Panaspis wahlbergii PEM R21297 WC 2723 SOUTH AFRICA: Mpumalanga, Doornkop Reserve KU236782 KU298713
Outgroup
Broadleysaurus major – – – AJ416922 HM161157
Cordylus marunguensis UTEP 20374 EBG 2993 DRC: Katanga, Pepa JQ389803 KU298675
Lacertaspis gemmiventris CAS 207858 RCD 13255 EQUATORIAL GUINEA: Bioko Island KU236792 KU298719
Lacertaspis gemmiventris CAS 207854 RCD 13251 EQUATORIAL GUINEA: Bioko Island KU236793 KU298720
Lacertaspis rohdei ZFMK 75382 – CAMEROON: Mt. Nlonako KU236790 KU298717
Leptosiaphos blochmanni UTEP 21177 EBG 1610 DRC: South Kivu, Bichaka KU236798 KU298722
Leptosiaphos koutoui MNHN 2001.0697 – CAMEROON: Meiganga, Adamaoua Plateau KU236789 KU298716
Leptosiaphos sp. ZFMK 75381 – CAMEROON: Mt. Nlonako KU236791 KU298718
Leptosiaphos sp. ZFMK 69552 – CAMEROON: Mt. Nlonako KU236794 KU298721
Mochlus afer ZFMK 54317 E56.17 KENYA: Kiyawetanga KU705386 KU841442
Plestiodon inexpectatus KU 8232 – – AY217990 AY662632
Trachylepis sulcata ZFMK 66424 – NAMIBIA: Kaokoland Region, Ongongo waterfall KC345403 HQ829808
Typhlosaurus braini CAS 214581 AMB 6340 NAMIBIA: Erongo Region, Rooibank HQ180025 HQ180106
Typhlosaurus braini CAS 214579 AMB 6338 NAMIBIA: Erongo Region, Rooibank HQ180128 HQ180137

Morphological methods

Specimens were measured with a digital caliper to the nearest 0.1 mm. Lepidosis was examined under a stereo-microscope. Scale nomenclature, scale counts and measurements used in the description follow Broadley (2000), Jacobsen and Broadley (2000), Soares et al. (2018) and Ceríaco et al. (2018, 2020). We measured the following 23 characters: snout–vent length (SVL), from snout to vent; tail length (TL), from anal plate to tip of tail, measured only in specimens with complete original tails; head length (HL), from tip of snout to anterior tympanum border; head height (HH), from the base of the maxilla to top of head; head width (HW), at the widest part of the maxilla; eye-diameter (ED); eye-tympanum distance (ET); eye-snout distance (ES), from the front of the eye to the tip of the snout; length of the arm (LA); length of the forearm (LFA); length of the leg (LL); length of the foreleg (LFL); distance between limbs (LD), minimum distance between the insertion of the anterior and posterior limbs; number of scale rows at midbody (MSR); number of paravertebral scales (SAD), from the nuchal (excluded from count) to base of the tail; number of scales ventrally (SAV), from the mental (excluded from count) to the anal plate (excluded); number of subdigital scales under Finger-IV (LUFF); number of subdigital scales under Toe-IV (LUFT); number of supralabials (SL), with those widened in subocular position indicated between brackets; number of supraciliaries (SC); nature of contact between parietals (CP); nature of contact between frontoparietals (CFP); nature of contact between prefrontals (CPF); nature of contact between prefrontal and preoculars (CPF/PO). Finally, coloration pattern was reported, and high-resolution photographs of preserved specimens taken. These data were compared with relevant literature on the group (Perret 1973, 1975; Jacobsen and Broadley 2000; Ceríaco et al. 2018a, 2020; Soares et al. 2018; Colston et al. 2020) and comparative material of congeners occurring in Angola and Nambia, namely P. cabindae, P. wahlbergii, P. maculicollis, P. mocamedensis and P. namibiana. All new specimens examined in this study are listed in the type and additional material sections of each species’ taxonomic account. Locality data are reported in the form of decimal degrees and use the WGS 84 map datum. Older (non-GPS) records are mostly derived from Marques et al. (2018) and have been georeferenced using GEOLocate web application (https://www.geo-locate.org). Elevations are all reported as meters above sea level.

Species concept

We adopt de Queiroz (1999) General Lineage Concept (GLC) which defines a species as a single, independent metapopulation lineage, diagnosable by multiple lines of evidence/criteria suggesting evolutionary independence. In order to fullfill the criteria of the GLC, we adopted an congruent integrative approach (sensu Padial et al. 2010) combining the results of the above mentioned mtDNA, nuclear DNA and morphological characters.

Results

Serra da Neve Panaspis included in our phylogenetic analysis belong to four mitochondrial and nuclear lineages, of which two are attributable to already known species, while the other two are not (Fig. 1). Serra da Neve representatives of P. cabindae were previously included in the dataset of Ceríaco et al. (2020). Besides P. cabindae, our analysis provides genetic verification for three other species of Panaspis at the Serra da Neve inselberg. Of these additional species, two specimens (MUNHAC/MB03-001532, 001533) from the base of Serra da Neve, near the village of Maylowe (-13.8280, 13.2625) unambiguously represent P. mocamedensis; six other specimens, morphologically very similar to P. cabindae and collected both near Maylowe (MUNHAC/MB03-001529, 001530, 001531, 001534; Fig. 2) and at the top of the inselberg, near Catchi village (MUNHAC/MB03-001525, 001526, 001528) are phylogenetically related to the clade containing P. cabindae and the Gulf of Guinea insular species; and a third form, represented by a single specimen (MUNHAC/MB03-001527) collected in riverine area near Catchi (-13.7618, 13.2514), phylogenetically is sister to P. annettesabinae from Ethiopia. Serra da Neve samples are genetically distinct, displaying respectively 5.3% and 2.9% pairwise 16S sequence divergence from their closest relatives (Table 2). These values fall within the usual recognized species-level divergence for the genus (Ceríaco et al. 2020). The haplotype network of RAG1 shows a significant divergence between the newly described and previously sequenced species. Additionally, while some Panaspis species, namely P. maculicollis, P. wahlbergii, and Panaspis Tanzania 1, share haplotypes, the newly sequenced species do not have any common haplotypes with other species (Fig. 2).

The four taxa of Panaspis occurring in the Serra da Neve Inselberg differ from each other and other Angolan and African species in several consistent morphological characters. Mensural and meristic data for the studied species are presented in Table 3. A combination of molecular and morphological data allows us to recognize the existence of four species of Panaspis in the Serra da Neve inselberg, two of which are here described as new species. Detailed diagnoses for the two new species are given in the taxonomic accounts below.

Figure 1.

Bayesian Inference (BI) phylogenetic tree inferred from the concatenated 16S and RAG1 dataset. Support values above the branches are BI per cent posterior probabilities/ Maximum Likelihood ultrafast bootstraps. Support values > 0.95 for both phylogenetic analyses are indicated by an asterisk. The most divergent external outgroup sequences were deleted for clarity.

Figure 2.

RAG1 haplotype (TCS) network showing the relationships of all Panaspis sequences (see Appendix 1). Circle size is proportional to observed haplotype frequencies, each dash indicates a nucleotide substitution. Coloured circles represent newly sequenced species.

Table 2.
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Uncorrected p-distance of the 16S RNA gene fragment between Panaspis taxa and respective nearest neighbour.

Taxon Mean p-distance (16S) % (nearest neighbour)
Panaspis mocamedensis 2.1 (Panaspis sp. Namibia)
Panaspis ericae sp. nov. 5.3 (Panaspis cabindae)
Panaspis mundavambo sp. nov. 2.9 (Panaspis annettesabinae)
Panaspis africanus 4.5 (Panaspis cabindae)
Panaspis breviceps 6.0 (Panaspis togoensis)
Panaspis cabindae 4.5 (Panaspis africanus)
Panaspis sp. Katanga 1 3.8 (Panaspis sp. Malawi)
Panaspis sp. Katanga 2 3.7 (Panaspis sp. Malawi)
Panaspis sp. Limpopo 1.2 (Panaspis sp. Namibia)
Panaspis maculicollis 3.3 (Panaspis sp. Mozambique 1)
Panaspis sp. Malawi 1.9 (Panaspis sp. Mozambique 4)
Panaspis sp. Mozambique 1 3.3 (Panaspis maculicollis)
Panaspis sp. Mozambique 4 1.9 (Panaspis sp. Malawi)
Panaspis namibiana Namibia 1.2 (Panaspis sp. Limpopo)
Panaspis sp. Tanzania 1 0.7 (Panaspis sp. Tanzania 2)
Panaspis sp. Tanzania 2 0.7 (Panaspis sp. Tanzania 1)
Panaspis togoensis 6.0 (Panaspis breviceps)
Panaspis wahlbergii 2.6 (Panaspis sp. Malawi)
Table 3.
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Comparison between the different species of Panaspis occurring in Angola and neighboring Namibia. Data from Jacobsen and Broadley (2000), Ceríaco et al. (2018, 2020) and this paper.

Panaspis aff. breviceps (n = 3) (data from Ceríaco et al. (2020)) Panaspis cabindae (n = 44) (data from Ceríaco et al. (2020)) Panaspis maculicollis (data from Jacobsen and Broadley 2000 – non-Angolan material) Panaspis wahlbergii (n = 4) (data from Ceríaco et al. (2020)) Panaspis mocamedensis (n = 6) (data from Ceríaco et al. (2020) and the two specimens collected from this study Panaspis namibiana (n = 11) (data from Ceríaco et al. (2018a) – Namibian material) Panaspis mundavambo sp. nov. (n = 1) Panaspis ericae sp. nov. (n = 6)
Maximum SVL 56.9 mm 38.4 mm 43 mm 38.7 mm 36.7 mm 42.6 mm 35.8 35.2
Maximum TL - 51.6 mm - 39 mm 39.3 mm 55.7 mm 51.7 44.8
Condition of the eye Movable lower eyelids Pre-ablepharine Ablepharine Ablepharine Ablepharine Ablepharine Ablepharine Pre-ablepharine
Presence of supranasals Yes Yes No No No No No Yes
Scales across Venter 52–61 53–62 - 56–62 56–61 56–67 54 55–60
Midbody Scale Rows 32–34 23–26 22–28 25–26 23–24 22–26 24 23–26
Scales across Dorsum 56–61 54–62 - 56–61 56–60 57–68 53 53–58
Lamellae under 4th finger 9–10 7–11 - 9–10 9–10 8–12 8 8–10
Lamellae under 4th finger 12–13 13–17 13–17 13–16 14–16 13–16 13 13–15
Frontoparietal scales condition Separated, but in median contact Separated, but in median contact Fused Fused Fused Fused Fused Separated, but in median contact
Presence of a white ventrolateral stripe no no no yes no no no no

Systematic accounts

Panaspis ericae sp. nov.

https://zoobank.org/DF1D1955-945C-44BF-B4FB-5E33548C9EEF
Figs 3, 4, 5, 6, 7, 8

Remarks

This new species belongs to the clade containing both P. cabindae and the Gulf of Guinea Oceanic Island taxa, P. africanus, P. thomensis and P. annobonensis. The phylogenetic relationship between P. cabindae and the island taxa had already been highlighted by Medina et al. (2016), Soares et al. (2018) and Ceríaco et al. (2020). The discovery of an additional taxa belonging to this Central-African clade is interesting, especially in the biogeographic context of the Serra da Neve inselberg. While most of the taxa occurring in the inselberg and its surroundings belong to here called “southern African” clades, the presence of other Central African associated species such as P. cabindae and Bitis gabonica (Marques et al. 2024) denotes a past connectivity between Serra da Neve and the north-south humid/highland corridors that allow taxa from Central Africa to expand to southern regions such as Namibe Province.

Type materials

Holotype. MUNHAC/MB03-001525 (field number LMPC 3209; Fig. 3), unsexed adult, from the rock outcrops near Catchi, Serra da Neve (-13.7653, 13.2571, 1645 m), Namibe Province, Republic of Angola, collected by Mariana P. Marques, Diogo Parrinha, Arthur Tiutenko and Luis M.P. Ceríaco on 29 October 2022.

Paratypes. MUNHAC/MB03-001526 (field number LMPC 3210), same data as holotype; MUNHAC/MB03-001528 (field number LMPC 3275) from the basecamp near Catchi (-13.7627, 13.2562, 1597 m), Namibe Province, Republic of Angola, collected on 1 November 2022; MUNHAC/MB03-001529 - 001531 (field numbers LMPC 3282, 3354, 3355; Fig. 4) from 2 km N of Maylowe, near a dry riverbed (-13.8265, 13.2601, 720 m), Namibe Province, Republic of Angola, collected between 3 and 5 November 2022. All specimens were collected by the same collectors as the holotype.

Additional material

2 km N of Maylowe, near dry riverbed [-13.8265, 13.2601, 720 m] (MUNHAC/MB03-001534).

Diagnosis

Panaspis ericae sp. nov. can be distinguished from other member of the genus by the following combination of characteristics: 1) presence of supranasals; 2) pre-ablepharine eye (as defined by Greer 1974); 3) frontoparietals not fused, in broad contact with each other; 4) dorsum coppery-brown, with a dorsolateral light stripe extending to tail, followed by dark brown flanks; 5) absence of dark spots on middorsal region; 5) absence of rows of light spots on the neck; 6) absence of a white ventrolateral stripe; 7) 23 to 26 midbody scales rows.

Description of the holotype

Unsexed adult in good condition. Arrangement and relative size of head, body and tail scalation typical for Panaspis. Robust, cylindrical body with well-developed pentadactyl limbs. Fore- and hind-limbs do not overlap when adpressed against the body. SVL 32.6 mm, TL 44.8 mm. Head length 6.1 mm, with relatively acuminate snout (HL 148% HW). Other relevant measurements are presented in Table 4. Rostral wider than high, and visible from above. Nasals widely separated behind rostral by frontonasal. Frontonasal flat anteriorly, wider than long. Nostrils small, set posteriorly in the nasals bordering the postnasal; supranasals present. Prefrontals separated from each other, contacting loreals, first supraciliary, first supraocular, frontonasal and frontal. Two loreals, subequal in size; two preoculars, inferior slightly larger. Frontal length shorter than distance between anterior tip of frontal and tip of snout; frontal in contact with prefrontals and frontonasal anteriorly, and with first supraoculars and frontoparietals posteriorly. Frontoparietals fused, in contact with each other, the frontal, supraoculars, parietals and interparietal. Frontoparietal plus interparietal length about 1.5 times the length of the frontal. Interparietal diamond shaped, posterior edge more acuminate, with visible parietal foramen in the center; parietals about the same width as that of the frontoparietals and contacting each other behind interparietal. A pair of large, broad nuchals collectively bordered by a total of eight dorsals. Supraoculars three. Supraciliaries four (left side) and five (right side), first higher than broad, last wider than high. Pretemporals two. Tympanum visible, ear opening wider than high, approximately one-fourth the height of the eye. Supralabials seven, the fifth being the subocular. Pre-ablepharine eye. Infralabials six. Postmental bordering seven scales (mental, two primary chin-shields, and two infralabials on each side). Ventral scales smooth. MSR 25, SAD 57, SAV 58. Scales on palms and soles smooth. Relative length of digits of manus III>IV>II>VI>I; relative length of digits of pes IV>III>II=V>I; tips missing on digits III of right pes, II of left pes, and digit 4 of right manus. LUFF 10 (left side); LUFT 14 (right and left side). Tail long, robust and tapering smoothly.

In preservative, dorsal aspect of head, dorsum and tail coppery brown; tail slightly lighter. A light dorsolateral stripe runs from posterior edge of eye through tail (becoming mostly indistinct on distal half of tail), bordered below by a thin black line. Flanks, face and sides of tail uniformly dark to greyish brown, slightly lighter on tail. Labials greyish white, with dark spots. Scales individually stippled with black, especially on top of head; some irregularly scattered dark spots on head and tail, but not on flanks or middorsal region. Ventrum greyish white; underside of tail creamy white.

Variation

Variaton in scalation and body measurements of the type series of Panaspis ericae sp. nov. is reported in Table 4. The majority of the paratypes agree entirely with the holotype, with few minor variations. In MUNHAC/MB03-001528 the prefrontals are in single point contact instead of separated. Paratype MUNHAC/MB03-001529 has an intrusive scale between parietals and nuchals. All paratypes generally agree with the holotype in terms of coloration.

Comparison with other Southwestern African Panaspis and related forms

As the molecular data provide evidence of the independence of the P. ericae sp. nov. lineage from all other taxa, we here restrict our morphological comparisons to those named congeners occurring in Namibia and Angola and its sister taxa from the Gulf of Guinea Oceanic Islands (P. africanus, P. thomensis and P. annobonensis). P. ericae sp. nov. can immediately be distinguished from P. maculicollis, P. wahlbergii, P. mocamedensis and P. mundavambo sp. nov. by having its frontoparietal scales not fused and in median contact, while on the latter taxa the frontoparietals are fused. It can also be easily distinguished from P. aff. breviceps by the condition of the eye (pre-ablepharine versus completely movable lower eyelids in P. aff. breviceps) and a considerably lower number of midbody scale rows (23–26 versus 32–34 in P. aff breviceps). Similarly, it can also be easily distinguished from the Gulf of Guinea Oceanic Island species, P. africanus, P. thomensis and P. annobonensis by the eyelid condition (pre-ablepharine in P. ericae sp. nov. versus completely movable lower eyelids in the latter taxa). P. ericae sp. nov. is morphologically very similar to P. cabindae, with which it was originally confused in the field. Due to the extreme morphological conservatism of the genus, the morphological differences between these two species are subtle. However, since the two species are truly cryptic we followed Rheindt et al. (2023) and present a molecular diagnosis of the species. Panaspis ericae sp. nov, differs from its closest relative P. cabindae by a total of 23 fixed single nucleotide substitutions plus one insertion in the 16S rRNA alignment and by 11 fixed single nucleotide substitutions in the RAG1 alignment (see Appendix 2). The main molecular diagnostic feature between both species is the base nucleotide composition at positions 112, 114, 131, 150, 167, 208, 209, 219, 229, 231, 232, 282, 288, 295, 304, 315, 317, 318, 320, 322, 324, 325, 334, and 336 of the 16S rRNA gene alignment and positions 6, 15, 183, 215, 234, 354, 366, 519, 521, 568, and 570 of the RAG1 gene alignment.

Distribution

The newly described species is currently only known from the Serra da Neve Inselberg in northern Namibe Province, southwestern Angola (Fig. 6). Given the isolation of the inselberg and the stark contrast with its surrounding habitat, the newly described species is assumed to be endemic to Serra da Neve.

Habitat and natural history notes

In the highlands of Serra da Neve, at about 1600 m above sea level, specimens were collected under leaf litter in an area dominated by dense Miombo woodlands (MB03-001525, 001526, 001528) (Fig. 7; Grandvaux-Barbosa 1970; Huntley 2023). On the other hand, the specimens collected in the lowlands of the inselberg, at about 720 m above sea level, were found near a dry riverbed in an area dominated by dense mopane woodland in sandy soils with abundant leaf litter (MUNHAC/MB03-001529, 001530, 001531, 001534) (Fig. 8) The species, as most Panaspis, seems to be a leaf-litter dweller. Its occurrence in both the base and top of Serra da Neve in two different habitats (miombo in the more humid highlands and mopane in the arid lowlands) indicates some ecological adaptability. It occurs in sympatry with Panaspis mocamedensis near Maylowe, preferring denser woodlands with more shade and leaf litter, especially associated with riparian areas, while P. mocamedensis occurs in the more open and arid woodlands. Paratypes MUNHAC/MB03-001530 and 001528 were gravid. Paratypes MUNHAC/MB03-001530 and 001531 were collected on a pitfall trap.

Etymology

The specific epithet “ericae” is formed in the genitive singular and is feminine. It is given in honor of Erica Tavares (1997–), an Angolan biologist and conservationist. Through her work in the Angolan environmental platform “Eco Angola” (of which Erica is a co-founder), Erica has revolutionized the Angolan conservation and ecological discourse, providing opportunities for members of the Angolan civil society, students, and young researchers to learn, debate and contribute to environmental causes. We suggest “Erica’s Snake-Eyed Skink” and “Lagartixa da Manta-Morta de Erica” as the English and Portuguese common names, respectively, for this species.

Figure 3.

Live photo of the holotype of Panaspis ericae sp. nov. (MUNHAC/MB03-001525) from vicinity of Catchi, Serra da Neve. Photo by Arthur Tiutenko.

Figure 4.

Live photo of the paratype of Panaspis ericae sp. nov. (MUNHAC/MB03-001531) from vicinity of Maylowe, Serra da Neve. Photo by Arthur Tiutenko.

Figure 5.

Preserved holotype of Panaspis ericae sp. nov. (MUNHAC/MB03-001525). Photos by Diogo Parrinha.

Figure 6.

Topographic map of the Serra da Neve Inselberg with collection localities of Panaspis species in the region. Stars represent the type localities of the newly described species.

Figure 7.

Miombo woodlands near the type locality of P. ericae sp. nov., in Catchi, Serra da Neve. Photo by Arthur Tiutenko.

Figure 8.

Dry riverbed in mopane woodlands and its associated leaflitter in the base of Serra da Neve, near Maylowe, where some of the paratypes of Panaspis ericae sp. nov., were collected. Photo by Arthur Tiutenko.

Panaspis mundavambo sp. nov.

https://zoobank.org/BFE63A66-529B-421D-8617-E78DDF44E9EE
Figs 6, 9, 10

Holotype

MUNHAC/MB03-1527 (field number LMPC 3242), unsexed adult, from the MPLA post near Catchi, Serra da Neve (-13.7618, 13.2514, 1614 m), Namibe Province, Republic of Angola, collected by Mariana P. Marques, Diogo Parrinha, Arthur Tiutenko and Luis M.P. Ceríaco on 31 October 2022 (Fig. 9).

Diagnosis

Panaspis mundavambo sp. nov. can be distinguished from other members of the genus by the following combination of characters: 1) absence of supranasals; 2) ablepharine eye (as defined by Greer 1974); 3) frontoparietals fused; 4) dorsum coppery-brown, with a thin dorsolateral light stripe extending from above eye to base of tail, most distinct on anterior third of its length; flanks dark brown; 5) absence of rows of light spots on the neck; 6) presence of series of black spots on middorsal region; 7) absence of a white ventrolateral stripe; 8) 24 midbody scales rows.

Description of the holotype

Unsexed adult in good condition, but the tail missing the tip. Arrangement and relative size of head, body and tail scalation typical for Panaspis. Robust, cylindrical body with well-developed pentadactyl limbs. Fore- and hind-limbs do not overlap when adpressed against the body. SVL 35.6 mm; tail incomplete, measuring 51.7 mm. Head length 6.6 mm, with relatively short snout (HL 129% HW). Other relevant measurements are presented in Table 4. Rostral wider than high, and visible from above. Nasals widely separated behind rostral by frontonasal. Frontonasal rounded anteriorly, wider than long. Nostrils small, set posteriorly in the nasals bordering the postnasal; supranasals absent. Prefrontals separated from each other, contacting loreals, first supraocular, first supraciliary, frontal and frontonasal. Two loreals, anterior one higher than broad, posterior one subquadrangular; preocular divided diagonally into two subtriangular scales on right side, and horizontally on left side into a small superior scale and a larger inferior one. Frontal length shorter than distance between anterior tip of frontal and tip of snout; frontal in contact with frontonasal and prefrontals anteriorly, and first supraocular posteriorly. Frontoparietals fused, in contact supraoculars; parietals and interparietal. Frontoparietal plus interparietal length twice the length of the frontal. Interparietal subtriangular, anterior margin wider and rounded, slightly acuminate posteriorly, with visible parietal foramen in the center; parietals about the same width as fused frontoparietals, in broad contact with each other behind interparietal. A pair of large, broad nuchals collectively bordered by a total of eight dorsals. Supraoculars three. Supraciliaries four, first higher than broad and last wider than high. Pretemporals two. Tympanum visible, ear opening wider than high, approximately one-fifth the height of the eye. Supralabials seven, the fifth being the subocular. Ablepharine eye. Infralabials six. Postmental bordering seven scales (mental, two primary chin-shields, and two infralabials on each side). Ventral scales smooth. MSR 24, SAD 53, SAV 54. Scales on palms and soles smooth. Relative length of digits of manus III>IV>II=V>I; relative length of digits of pes IV>III>II=V>I; toe V of right pes missing. LUFF scales eight; LUFT scales 13 on right side and 12 on left side. Tail long, robust and tapering smoothly.

In preservative, ground color of dorsum and upper side of head coppery brown; tail lighter, golden brown. A thin, light dorsolateral stripe extends from supraocular area to base of tail, being most distinct on anterior third of its length and faint posteriorly; below, a dark brown lateral band extends from snout to base of tail, darker on face, neck and axilla, lighter and less distinct from middorsal coloration posteriorly. Dorsal scales individually stippled with black, especially on dorsal aspect of head; labials greyish white, with individual black spots; pale and black spots scattered on lateral sides of neck. Middorsal region with black spots, forming somewhat longitudinal series. Venter uniformly greyish to blueish white; underside of tail slightly orange; palms and soles brownish.

Comparison with other Southwestern African Panaspis and related forms

As the molecular data provide evidence of the independence of the P. mundavambo sp. nov. lineage from all other taxa, we here restrict our morphological comparisons to those named congeners occurring in Namibia and Angola and its sister taxon P. annettesabinae. P. mundavambo sp. nov. can immediately be distinguished from P. cabindae. P. ericae sp. nov. and P. aff. breviceps, by not having supranasals (versus present in P. cabindae, P. ericae sp. nov. and P. aff. breviceps), by having an ablepharine eye (versus pre-ablepharine eye in P. cabindae and P. ericae sp. nov., and completely movable lower eyelids in P. aff. breviceps), and by having the frontoparietals fused (versus divided in P. cabindae, P. ericae sp. nov. and i P. aff. breviceps). In comparison with P. wahlbergii, P. mundavambo sp. nov. can be distinguished by its coloration, namely by the presence of longitudinal series of dark spots on the middorsal region and the lack of a ventrolateral white stripe (versus the absence of middorsal dark spots and presence of white ventrolateral stripe in P. wahlbergii), and by having an overall lower number of scales across venter (54 versus 56–62 in P. wahlbergii), midbody scales rows (24 versus 25–26 in P. wahlbergii), scales across dorsum (53 versus 56–61 in P. wahlbergii) and lamellae under the fourth finger (8 versus 9–10 in P. wahlbergii). Panaspis mundavambo sp. nov. can be distinguished from P. maculicollis by the presence of longitudinal series of dark spots on middorsal region (versus no dark spots on middorsal region in P. maculicollis), and a lower number of lamellae under fourth finger (8 versus 13–17). P. mundavambo sp. nov. can be distinguished from P. namibiana by having a lower count of scales across venter (54 in P. versus 56–67 in P. namibiana) and dorsum (53 versus 57–68 in P. namibiana). P. mundavambo sp. nov. can be distinguished from P. mocamedensis by the presence of series of black spots on middorsal region (versus black spots absent in P. mocamedensis), and by having a smaller count of scales across venter (54 in P. versus 56–61 in P. mocamedensis), across dorsum (53 versus 56–60 in P. mocamedensis) and smaller number of lamellae under the fourth finger (8 versus 9–10 in P. mocamedensis). Comparing P. mundavambo sp. nov. with its sister taxon, P. annettesabinae from Ethiopia, the newly described species has prefrontal scales separated from each other (versus in contact in P. annettesabinae), a smaller count of scales across venter (54 versus 68 in P. annettesabinae), smaller number of scales across dorsum (53 versus 62 in P. annettesabinae).

Distribution

The newly described species is currently only known from single locality on the Serra da Neve Inselberg, southwestern Angola (Fig. 6). Given the isolation of the inselberg and the stark contrast with its surrounding habitat, the newly described species is assume to be locally restricted, although it may occur elsewhere in Angola (W. Conradie, pers. comm.).

Habitat and natural history notes

The holotype was collected under a log in an area dominated by woodlands at about 1600 m above sea level (Grandvaux-Barbosa 1970; Huntley 2023). Very little is known about its ecology and natural history. It seems to be associated with riparian areas (Fig. 10). It occurs close to P. ericae sp. nov.. The single collected specimen was found in denser woodlands near a riparian gallery, while P. ericae sp. nov. was found in slightly more open miombo that dominates the landscape.

Etymology

The specific epithet “mundavambo” refers to the Mukwando (local tribe) name for Serra da Neve Inselberg (Opunda Mundavambo), to which the species is endemic, and is applied here as a substantive in apposition. We propose the English vernacular name “Serra da Neve Snake-Eyed Skink” and the Portuguese vernacular name of “Lagartixa da Manta-Morta da Serra da Neve”.

Table 4.
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Measurements (in mm) and scale counts of the holotype and paratypes of Panaspis ericae sp. nov. and Panaspis mundavambo sp. nov. Abbreviations are the same as those described in Materials and methods.

Taxon Panaspis ericae sp. nov. Panaspis mundavambo sp. nov.
Specimen ID. MUNHAC/MB03-1525 (Holotype) MUNHAC/MB03-1526 (Paratype) MUNHAC/MB03-1528 (Paratype) MUNHAC/MB03-1529 (Paratype) MUNHAC/MB03-1530 (Paratype) MUNHAC/MB03-1531 (Paratype) MUNHAC/MB03-1527 (Holotype)
SEX Unsexed Unsexed Female Unsexed Female Unsexed Unsexed
SVL 32.6 35.2 34.7 31.0 34.4 29.3 35.6
TL 44.8 _ _ 43.8 36.2 _ 51.7 (missing tail tip)
HL 6.1 6.0 5.7 5.5 5.5 5.2 6.6
HH 2.6 2.5 2.8 2.4 2.2 2.5 4.2
HW 4.1 4.2 3.8 4.0 3.9 3.6 5.1
LA 2.5 2.5 2.3 2.4 2.2 2.1 2.8
LFA 2.0 1.7 1.7 2 1.9 1.7 2.5
LL 3.6 3.5 2.9 2.8 3.0 2.9 4.0
LFL 2.9 3.0 2.3 2.2 2.4 2.1 3.4
LD 17.3 20.8 20.3 17.8 19.9 16.4 19.4
ED 1.1 1.1 1.0 1.0 0.9 1.1 0.9
ET 2.2 2.0 2.0 2.3 2.1 2.2 2.3
ES 2.5 2.5 2.1 2.4 2.9 2.1 2.9
MSR 25 25 24 25 26 23 24
SAV 58 57 57 57 60 55 54
SAD 57 55 56 53 58 54 53
LUFT 14 15 13 15 15 13 13
LUFF 10 9 9 19 9 8 8
SC 4 5 4 5 5 4 5
SL(SO) 7(5) 6(5) 7(5) 7(5) 7(5) 7(5) 7(5)
CP Contact Contact Contact Contact Contact Contact Contact
CFP Contact Contact Contact Contact Contact Contact Fused
CPF Separated Separated Single Point Contact Single Point Contact Separated Separated Separated
Figure 9.

Preserved holotype of Panaspis mundavambo sp. nov. (MUNHAC/MB03-1527). Photos by Diogo Parrinha.

Figure 10.

Type locality of P. mundavambo sp. nov. from a riparian gallery near Catchi, Serra da Neve. Photo by Luis M.P. Ceríaco.

Discussion

The discovery of two new species of Panaspis on Serra da Neve is unexpected, as is the inselberg's capacity for harboring a total of four species of such cryptic andecologically conservative group of animals. Panaspis, also known as leaf-litter skinks, are generally ground dwellers who inhabit areas of leaf-litter and forest floors preying on termites and other small invertebrates (MPM pres. Obs.). While our data are extremely limited to make any kind of robust ecological inference, it appears that the type of soil and habitat may ecologically partition the niche of the four taxa: P. mundavambo sp. nov. may be restricted to high elevation riparian galleries, while P. cabindae occurs in lowland humid and forested areas. P ericae sp. nov. was found both in miombo and mopane habitats, but in the latter, it was found only in dry riverbeds, while P. mocamedensis in leaf-litter outside these. This ecological partition needs attention and Serra da Neve Panaspis may become an interesting case study for anyone wanting to focus on these types of ecological questions.

Besides this surprising intrageneric diversity in such a small area, the discovery of these two new species recovers recurring patterns regarding the Serra da Neve herpetofauna. In just the last five years, one toad (Ceríaco et al. 2018b) and four reptile strictly endemic species (Marques et al. 2019, 2020, 2023a; Branch et al. 2021), one strictly endemic gecko in the process of description (Parrinha et al. in prep.), and four other lizard species (Parrinha et al. 2021; Marques et al. 2022, 2023b; Ceríaco et al. 2024) have been described from the inselberg and its surroundings. These discoveries demonstrate that Serra da Neve is a biodiversity hotspot, particularly interesting for the large proportion of endemic species per area. Therefore, the description of these two additional species of Panaspis fits into the recent history of discovery in the inselberg, which is likely to continue yielding new taxonomic discoveries. Biogeographically, these two new species also share common trends with other recently described species. It has been shown that some of the Serra da Neve endemic taxa have closely related species in the Eastern African highlands, as is the case for Acontias mukwando and Lygodactylus baptistai (Marques et al. 2020, 2023b), a pattern also seen in the newly described P. mundavambo sp. nov. On the other hand, P. ericae sp. nov. belongs to a more Central African lineage, composed of species such as P. cabindae and the Gulf of Guinea Oceanic Islands taxa. The presence of P. mocamedensis at the base of Serra da Neve ties the Panaspis fauna to the more southwestern Angolan endemic fauna. Serra da Neve is thus an interesting biogeographic melting pot, which ultimately contributes to its rich and unique biodiversity. Interestingly though, P. wahlbergii has not been recorded in the inselberg, despite being common in the neighboring Huila escarpment and plateau (Ceríaco et al. 2020).

This new addition to the known endemic fauna of Serra da Neve is another example of the conservation importance of this inselberg, a fact that has already been noted by several authors (Pinto et al. 2023; Marques et al. 2024). As the two new species appear to be endemic to Serra da Neve, and therefore their areas of occurrence do not exceed 630 km2 (assuming that both of the species are homogeneously distributed across the inselberg), both are of conservation interest. Nevertheless, the currently available data do not allow us to conduct a full-scale conservation assessment and therefore we suggest that a status of Data Deficient (DD) is warranted for both species (IUCN Standards and Petitions Committee 2019). More work is needed to uncover the true biodiversity of Serra da Neve and ensure its protection for future generations.

Acknowledgments

The present work is a result of the ongoing collaboration between the Instituto Nacional de Biodiversidade e Áreas de Conservação (INBAC) from the Ministry of Environment of Angola and its international partners. Angolan specimens were collected and exported under permits issued by INBAC (65/INBAC.MINAMB/2022). We also thank the provincial and local authorities for their support and cooperation during our fieldwork. We thank Adam Ferguson, Ben Marks and Daryl Coldren for their support during field work. Special thanks to Álvaro (“Varito”) Baptista and his team from Omauha Lodge, for all the assistance, great support, and friendship during the field work. This work was funded by the National Geographic Society Explorer Grant (NGS-73084R-20) to LMPC. Fundação para a Ciência e Tecnologia (FCT) supported DP under grant (2021.05238.BD), MLL under contract (2020.03608.CEECIND) and MPM under grants (SFRH/BD/129924/2017, COVID/BD/152155/2022). Work supported by the European Union’s Horizon 2020 Research and Innovation Programme under the Grant Agreement Number 857251. AMB and LMPC were supported by grant DEB 2146654 from the National Science Foundation of the United States.

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Appendix 1

Table A1.
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RAG1 haplotype (TCS) network showing the relationships of all Panaspis sequences.

SPECIES COLLECTION NUMBER RAG1 HAPLOTYPE
Panaspis ericae sp. nov. LMPC-3209 1
Panaspis ericae sp. nov. LMPC-3210 1
Panaspis ericae sp. nov. LMPC-3275 1
Panaspis ericae sp. nov. LMPC-3282 1
Panaspis ericae sp. nov. LMPC-3354 1
Panaspis ericae sp. nov. LMPC-3355 1
Panaspis ericae sp. nov. LMPC-3395 1
Panaspis mundavambo sp. nov. LMPC-3242 9
Panaspis mocamedensis LMPC-3381 17
Panaspis mocamedensis LMPC-3382 17
Panaspis africanus – 8
Panaspis breviceps ZFMK 87662 2
Panaspis breviceps ZFMK 87663 2
Panaspis cabindae PEM R19467 5
Panaspis cabindae Uncatalogued 5
Panaspis cabindae Uncatalogued 5
Panaspis cabindae PEM R21594 6
Panaspis cabindae PEM R20256 5
Panaspis cabindae AMB 10315 3
Panaspis cabindae FLMNH 187242 4
Panaspis cabindae MB03-001088 5
Panaspis cabindae MB03-001091 7
Panaspis sp. Katanga 1 UTEP 21175 26
Panaspis sp. Katanga 1 UTEP 21174 26
Panaspis sp. Katanga 2 – 27
Panaspis sp. Katanga 2 PEM R17454 29
Panaspis sp. Katanga 2 – 28
Panaspis sp. Limpopo – 18
Panaspis maculicollis CAS 234099 19
Panaspis maculicollis CAS 234188 13
Panaspis maculicollis CAS 234135 13
Panaspis maculicollis Uncatalogued 14
Panaspis maculicollis Uncatalogued 13
Panaspis maculicollis PEM R20475 15
Panaspis sp. Malawi PEM R20247 30
Panaspis sp. Mozambique 1 – 10
Panaspis sp. Mozambique 1 PEM R20561 11
Panaspis sp. Mozambique 1 PEM R20591 12
Panaspis sp. Mozambique 4 PEM R20569 31
Panaspis sp. Mozambique 4 PEM R20576 31
Panaspis namibiana MCZ R183767 16
Panaspis sp. Tanzania 1 – 22
Panaspis sp. Tanzania 1 – 19
Panaspis sp. Tanzania 2 PEM R20799 20
Panaspis togoensis TCWC 94557 18
Panaspis wahlbergii CAS 234209 24
Panaspis wahlbergii MCZR 184443 13
Panaspis wahlbergii CAS 234194 19
Panaspis wahlbergii TM 84299 24
Panaspis wahlbergii NMB R10286 23
Panaspis wahlbergii TCWC 95563 21
Panaspis wahlbergii TCWC 95588 19
Panaspis wahlbergii PEM R21298 25
Panaspis wahlbergii PEM R21297 23
Broadleysaurus major – –
Cordylus marunguensis UTEP 20374 –
Lacertaspis gemmiventris CAS 207858 –
Lacertaspis gemmiventris CAS 207854 –
Lacertaspis rohdei ZFMK 75382 –
Leptosiaphos blochmanni UTEP 21177 –
Leptosiaphos koutoui MNHN 2001.0697 –
Leptosiaphos sp. ZFMK 75381 –
Leptosiaphos sp. ZFMK 69552 –
Mochlus afer ZFMK 54317 –
Plestiodon inexpectatus – –
Trachylepis sulcata – –
Typhlosaurus braini CAS 214581 –
Typhlosaurus braini CAS 214579 –

Appendix 2

Fixed nucleotide differences of 16S rRNA and RAG1 gene sequences of P. ericae sp. nov. compared to the se­quence alignment of P. cabindae (del = deletion; ins = in­sertion mutation).

Panaspis ericae sp. nov.

16S rRNA: 112C, 114G, 131C, 150T, 167A, 208T, 209 ins T, 219A, 229C, 231C, 232T, 282A, 288A, 295T, 304C, 315G, 317T, 318G, 320T, 322C, 324C, 325C, 334C, 336T.

RAG1 : 6T, 15G, 183A, 215A, 234T, 354A, 366A, 519T, 521C, 568C, 570G.

Panaspis cabindae

16S rRNA: 112T, 114A, 131T, 150C, 167G, 208A, 209 del, 219G, 229A/G, 231A, 232C, 282G, 288C, 295A, 304A, 315T/A, 317A, 318C, 320C, 322G, 324A, 325T, 334A, 336G.

RAG1 : 6G, 15C, 183G, 215G, 234C, 354G, 366G, 519C, 521T, 568A, 570A.

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