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Pelomedusa subrufa (Bonnaterre, 1789) sensu stricto 1789 Testudo subrufa Bonnaterre —Restricted type locality (Bour 1982): Taolañaro, Madagascar; holotype: Muséum National d’Histoire naturelle, Paris, MNHN 7970 (Fig. 2 in Fritz et al. 2014) 1798 Testudo badia Donndorff (nomen novum) 1798 Testudo rubicunda Suckow (nomen novum) 1935 Pelomedusa galeata damarensis Hewitt —Type locality: Quickborn near Okahandja, Namibia; lectotype (Fritz et al. 2014): Port Elizabeth Museum, PEM R14953 (Fig. 7 top in Fritz et al. 2014) 1937 Pelomedusa subrufa wettsteini Mertens —Type locality: Majunga (Mahajanga), western Madagascar; holotype: Senckenberg-Museum, Frankfurt am Main, SMF 7958 (Fig. 8 in Fritz et al. 2014) Diagnosis: Small to medium-sized helmeted terrapins with a known maximum straight carapacial length of 19.7 cm. Yet, the shell length of most adults is below 14 cm. Pectoral scutes in most specimens with broad contact at plastral midseam; however, terrapins with narrow contact or, exceptionally, with triangular pectorals without midseam contact occur. One large undivided temporal scale on each side of head. Two small barbels under chin. Carapace uniform light to dark brown. Plastron in larger individuals mostly light coloured, in younger specimens often with dark markings. Pelomedusa subrufa sensu stricto differs from all other Pelomedusa species by the presence of cytosine (C) instead of thymine (T) at position 117, by the presence of thymine (T) instead of adenine (A) at position 169, by the presence of thymine (T) instead of adenine (A) or cytosine (C) at position 191, by the presence of guanine (G) instead of adenine (A) at positions 226 and 296, by the presence of adenine (A) instead of cytosine (C) at position 280, and by the presence of cytosine (C) or thymine (T) instead of adenine (A) or guanine (G) at position 289 of the 360-bp-long reference alignment of the 12S rRNA gene (Supporting Information). In addition, P. subrufa sensu stricto differs from all other Pelomedusa species except P. galeata by the presence of cytosine (C) instead of adenine (A) or guanine (G) at position 148, by the presence of guanine (G) instead of adenine (A) at position 159, by the presence of cytosine (C) instead of thymine (T) at position 167, and by the presence of guanine (G) instead of adenine (A) or thymine (T) at position 343. Moreover, P. subrufa sensu stricto differs from all other Pelomedusa species except P. galeata and possibly P. gehafie, P. kobe and candidate species B (in which the respective character states are unknown) by the presence of thymine (T) instead of cytosine (C) at position 26 and by the presence of cytosine (C) instead of thymine (T) at position 38. Distribution: Southern Angola (this study), Botswana, southeastern Democratic Republic of the Congo, Malawi (Vargas-Ramírez et al. 2010), Namibia (Wong et al. 2010; this study), South Africa (Limpopo; this study), and Kilimanjaro region of Tanzania (this study). Introduced to Madagascar (Vargas-Ramírez et al. 2010; Wong et al. 2010). Remarks: Pelomedusa subrufa sensu stricto corresponds to mtDNA lineage VIII of Vargas-Ramírez et al. (2010). Besides P. galeata, it is one of the two southern species of Pelomedusa. In the course of the present study, we recorded P. subrufa sensu stricto for the first time for South Africa, based on one sample from the western border region of the Kruger Park (Hoedspruit, Limpopo, South Africa). To the west and south, the next genetically verified records are approximately 300 km distant and are P. galeata. It seems possible that the ranges of the two species abut or overlap in northeastern South Africa. In the Zoologische Staatssammlung München there are genetically verified specimens of P. subrufa sensu stricto collected on 3 May 1937 by Wolfgang Uthmöller (25 October 1904 – 25 August 1951) at the northern slope of the Kilimanjaro, Tanzania (ZSM 320/1937:1–4), and additional genetically verified specimens with the date 5 May 1937 originating from the same collector, but bearing only “Tanganyika” as locality data (ZSM 285/ 1937:1–3). The latter specimens represent another species (P. kobe, see above), suggesting that both taxa occur in close geographical proximity or even sympatry in Tanzania. This is also supported by further genetically verified specimens of P. kobe from the Arusha region, Tanzania, from where also the new species P. neumanni is recorded (see above). The identity of mtDNA sequences of P. subrufa sensu stricto from Tanzania and Madagascar points to Tanzania as geographical source region for the introduced Malagasy helmeted terrapins. Among 94 museum specimens of P. subrufa, two terrapins have triangular pectorals without midseam contact, and 19 terrapins have pectoral scutes with narrow plastral midseam contact, whilst the remaining specimens have pectoral scutes with broad contact at the midseam. Pelomedusa subrufa can cope with extremely arid conditions. For Namibia, there are records from the mouths of temporary streams (so-called riviers) in the Namib Desert (Branch 2008; Boycott & Bourquin 2008; vouchers in the Museum für Naturkunde, Berlin). It is well known that Namibian terrapins burrow underground during arid periods, and if there is no rain, they may evidently survive up to six years burrowed (A. Schleicher, pers. observ., Omaheke, Namibia). The monogenean flatworm Polystomoides chabaudi Euzet & Combes, 1965 was described from Pelomedusa subrufa sensu stricto (collected at Betioky, Madagascar). The same polystomatid species was also reported from helmeted terrapins from the Kampala area, Uganda (Tinsley 1973), that is, probably from P. neumanni., Published as part of Petzold, Alice, Vargas-Ramírez, Mario, Kehlmaier, Christian, Vamberger, Melita, Branch, William R., Preez, Louis Du, Hofmeyr, Margaretha D., Meyer, Leon, Schleicher, Alfred, Široký, Pavel & Fritz, Uwe, 2014, A revision of African helmeted terrapins (Testudines: Pelomedusidae: Pelomedusa), with descriptions of six new species, pp. 523-548 in Zootaxa 3795 (5) on pages 542-543, DOI: 10.11646/zootaxa.3795.5.2, http://zenodo.org/record/4915083, {"references":["Bonnaterre, P. J. (1789) Tableau encyclopedique et methodique des trois regnes de la nature. Erpetologie. Panckoucke, Hotel de Thou, Paris, xxviii + 70 + (1) pp., 35 plates. http: // dx. doi. org / 10.5962 / bhl. title. 59326","Fritz, U., Petzold, A., Kehlmaier, C., Kindler, C., Campbell, P., Hofmeyr, M. D. & Branch, W. R. (2014) Disentangling the Pelomedusa complex using type specimens and historical DNA. Zootaxa, 3795 (5), 501 - 522 http: // dx. doi. org / 10.11646 / zootaxa. 3795.5.1","Vargas-Ramirez, M., Vences, M., Branch, W. R., Daniels, S. R., Glaw, F., Hofmeyr, M. D., Kuchling, G., Maran, J., Papenfuss, T. J., Siroky, P., Vieites, D. R. & Fritz, U. (2010) Deep genealogical lineages in the widely distributed African helmeted terrapin: evidence from mitochondrial and nuclear DNA (Testudines: Pelomedusidae: Pelomedusa subrufa). Molecular Phylogenetics and Evolution, 56, 428 - 440. http: // dx. doi. org / 10.1016 / j. ympev. 2010.03.019","Wong, R. A., Fong, J. J. & Papenfuss, T. J. (2010) Phylogeography of the African helmeted terrapin, Pelomedusa subrufa: genetic structure, dispersal, and human introduction. Proceedings of the California Academy of Sciences, 61, 575 - 585.","Branch, B. (2008) Tortoises, Terrapins & Turtles of Africa. Struik Publishers, Cape Town, 128 pp.","Boycott, R. C. & Bourquin, O. (2008) Pelomedusa subrufa (Lacepede 1788), helmeted turtle, helmeted terrapin. In: Rhodin, A. G. J., Pritchard, P. C. H., van Dijk, P. P., Saumure, R. A., Buhlmann, K. A. & Iverson, J. B. (Eds.), Conservation Biology of Freshwater Turtles and Tortoises: A Compilation Project of the IUCN / SSC Tortoise and Freshwater Turtle Specialist Group. Chelonian Research Monographs, No. 5. Chelonian Research Foundation, Lunenburg, MA, pp. 007.1 - 007.6. http: // dx. doi. org / 10.3854 / crm. 5.007. subrufa. v 1.2008","Euzet, L. & Combes, C. (1965) Parasites des cheloniens malgaches. Polystomoides chabaudi n. sp. (Monogenea) chez la tortue d'eau douce Pelomedusa subrufa Lacepede 1788. Annales de Parasitologie Humaine et Comparee, 40, 455 - 450.","Tinsley, R. C. (1973) Observations on Polystomatidae (Monogenoidea) from East Africa with a description of Polystoma makereri n. sp. Zeitschrift fur Parasitenkunde, 42, 251 - 263. http: // dx. doi. org / 10.1007 / bf 00328887"]}

Pelomedusa barbata sp. nov. Diagnosis: Medium-sized, light-coloured helmeted terrapins with a known maximum straight carapacial length of 21.6 cm (Gasperetti et al. 1993). Pectorals with narrow to broad midseam contact. Two small temporal scales (rarely only one large scale) on each side of head. Two or three large to very large barbels under chin. In adults, carapace light coloured, plastron entirely yellow. Pelomedusa barbata differs from all other Pelomedusa species by a gap instead of adenine (A), cytosine (C) or thymine (T) at position 122 and by the presence of guanine (G) instead of adenine (A) at position 330 of the 360-bp-long reference alignment of the 12S rRNA gene (Supporting Information). Holotype: Museum für Tierkunde, Senckenberg Dresden (MTD D 24637, male, Zinjibar, Abyan, Yemen, N13°7.75 E45°22.81; leg. W. Wranik, 3 June 1985; Fig. 3 top). Description of the holotype: Straight carapacial length 12.3 cm, plastral length 10.4 cm. Pectoral scutes triangular and in narrow contact at the midline. Temporal scale divided on left side, undivided on right side of head. Two large barbels below chin. Carapace horn-coloured, plastron yellow with sparse grey mottling. Dorsal side of soft parts grey to horn-coloured, ventral side cream-coloured with a yellow tinge. Tail long. Paratypes: Muséum d’Histoire Naturelle Genève (MNHG 2310.062, adult male, Al Sukhna, Yemen; MNHG 2455.073 – 075, adult male and two juveniles, Wadi Warazan, Lahij, Yemen); Museum für Tierkunde, Senckenberg Dresden (MTD D 24638, adult female, Lahij, Yemen); The Natural History Museum, London (BMNH 1985.1475, mummified adult, Al Kadan, Tihama, Al Hudaydah, Yemen; BMNH 1985.1478 – 1479, adult male and female, Amanat Al Asimah, Sana’a Area, Yemen); Zoologisches Forschungsmuseum Alexander Koenig, Bonn (ZFMK 87122, juvenile, Asir Region, Saudi Arabia). Derivatio nominis: The scientific name (Latin: bearded) refers to the large barbels beneath the chin of the new species. Distribution: Southwestern Arabian Peninsula (Saudi Arabia, Yemen; Gasperetti et al. 1993; Vargas-Ramírez et al. 2010; Wong et al. 2010). Remarks: Pelomedusa barbata corresponds to mtDNA lineage VII of Vargas-Ramírez et al. (2010). According to our analyses of mtDNA, P. barbata belongs to the northern species group of Pelomedusa and is most closely related to P. somalica (Fig. 1). Gasperetti et al. (1993) mention that they have studied helmeted terrapins from southwestern Arabia that “more or less encompass all of the plastral variations” described for Pelomedusa and “all of the colour descriptions, from the lightest tan colour to almost black”. However, their figures show only terrapins with pectoral scutes in contact, matching our observations. We never encountered dark coloured Arabian terrapins. Pelomedusa barbata is the only chelonian species endemic to the Arabian Peninsula. Due to the paucity of local freshwater habitats, P. barbata should be regarded as an endangered species.

Pelomedusa olivacea (Schweigger, 1812) 1812 Emys olivacea Schweigger —Restricted type locality: Senegal; holotype: Muséum National d’Histoire naturelle, Paris, MNHN 7971 1884 Pelomedusa gasconi Rochebrune —Restriced type locality (by neotype designation, Fritz et al. 2014): Dagana, Senegal; neotype (Fritz et al. 2014): Zoologisches Forschungsmuseum Alexander Koenig, Bonn, ZFMK 17076 (Fig. 6 bottom in Fritz et al. 2014) Diagnosis: Small to medium-sized helmeted terrapins with a known maximum straight carapacial length of 16.8 cm. Pectoral scutes triangular and either just meeting at their tips along the plastral midseam or more or less widely separated. One large undivided temporal scale on each side of head. Two small barbels under chin. Carapace and plastron light coloured. Soft parts dorsally darker than ventrally. Pelomedusa olivacea differs from all other Pelomedusa species by the presence of guanine (G) instead of adenine (A) or cytosine (C) at position 123 and by the presence of adenine (A) instead of guanine (G) at position 271 of the 360-bp-long reference alignment of the 12S rRNA gene (Supporting Information). Distribution: Genetically verified records for P. olivacea are known from Benin, Burkina Faso, Niger, Nigeria, and Senegal (Vargas-Ramírez et al. 2010; Wong et al. 2010; Fritz et al. 2014). Remarks: Pelomedusa olivacea corresponds to mtDNA lineage III of Vargas-Ramírez et al. (2010). Phylogenetically, P. olivacea belongs to the northern clade of Pelomedusa and is most closely related to P. variabilis from Ghana and the Ivory Coast (Fig. 1). In both species, the pectoral scutes may be triangular with or without midseam contact. Among 21 museum specimens of P. olivacea, 18 have completely divided pectoral scutes and three have triangular pectoral scutes just meeting at the plastral midline. This matches the description for Senegalese P. olivacea given in Rochebrune (1884). The monogenean flatworm Polystomoides nabedei Kulo, 1980 was described from helmeted terrapins from Siborototi, Togo (most likely Pelomedusa olivacea or P. variabilis). This parasite was described as a distinct species based on morphological differences from Polystomoides chabaudi, which is known from Pelomedusa subrufa and probably P. neumanni (see respective species accounts)., Published as part of Petzold, Alice, Vargas-Ramírez, Mario, Kehlmaier, Christian, Vamberger, Melita, Branch, William R., Preez, Louis Du, Hofmeyr, Margaretha D., Meyer, Leon, Schleicher, Alfred, Široký, Pavel & Fritz, Uwe, 2014, A revision of African helmeted terrapins (Testudines: Pelomedusidae: Pelomedusa), with descriptions of six new species, pp. 523-548 in Zootaxa 3795 (5) on page 539, DOI: 10.11646/zootaxa.3795.5.2, http://zenodo.org/record/4915083, {"references":["Fritz, U., Petzold, A., Kehlmaier, C., Kindler, C., Campbell, P., Hofmeyr, M. D. & Branch, W. R. (2014) Disentangling the Pelomedusa complex using type specimens and historical DNA. Zootaxa, 3795 (5), 501 - 522 http: // dx. doi. org / 10.11646 / zootaxa. 3795.5.1","Vargas-Ramirez, M., Vences, M., Branch, W. R., Daniels, S. R., Glaw, F., Hofmeyr, M. D., Kuchling, G., Maran, J., Papenfuss, T. J., Siroky, P., Vieites, D. R. & Fritz, U. (2010) Deep genealogical lineages in the widely distributed African helmeted terrapin: evidence from mitochondrial and nuclear DNA (Testudines: Pelomedusidae: Pelomedusa subrufa). Molecular Phylogenetics and Evolution, 56, 428 - 440. http: // dx. doi. org / 10.1016 / j. ympev. 2010.03.019","Wong, R. A., Fong, J. J. & Papenfuss, T. J. (2010) Phylogeography of the African helmeted terrapin, Pelomedusa subrufa: genetic structure, dispersal, and human introduction. Proceedings of the California Academy of Sciences, 61, 575 - 585.","Rochebrune, A. - T. de (1884) Faune de la Senegambie. Reptiles. Octave Doin, Paris, 41 + (1) pp., 8 plates.","Kulo, S. - D. (1980) Parasites de cheloniens en Afrique intertropicale. Polystomoides nabedei n. sp. (Monogenea, Polystomatidae), parasite de la vessie urinaire de la tortue d'eau douce Pelomedusa subrufa Lacepede, 1788 (Chelonien, Pelomedusidae). Annales de Parasitologie Humaine et Comparee, 55, 367 - 377."]}

Pelomedusa schweinfurthi sp. nov. Diagnosis: Small to medium-sized, dark-coloured helmeted terrapins with a known maximum straight carapacial length of 15.7 cm. Pectoral scutes rectangular with wide midseam contact or triangular with narrow midseam contact. Temporal head scales large, mostly undivided. Two small barbels under chin. In adults, carapace and plastron rather dark; soft parts ventrally lighter than dorsally. Pelomedusa schweinfurthi differs from all other Pelomedusa species by the presence of thymine (T) instead of cytosine (C) at position 279, by the presence of guanine (G) instead of adenine (A) or a gap at position 297, and by the presence of adenine (A) instead of guanine (G) at position 305 of the 360-bp-long reference alignment of the 12S rRNA gene (Supporting Information). Holotype: Senckenberg-Museum, Frankfurt am Main (SMF 56161, female, Liria, Central Equatoria, South Sudan, N4°38.66 E32°4.83; leg. C. Scherpner, 3 March 1955; Fig. 4 bottom). Description of the holotype: Straight carapacial length 12.3 cm, plastral length 10.5 cm. Pectoral scutes rectangular with wide midseam contact. Temporal scale divided on left side, undivided on right side of head. Two small barbels under chin. Carapace and plastron dark brown. Dorsal side of soft parts and entire forelegs dark grey, ventral side of hind legs light grey. Tail short. Paratypes: Senckenberg-Museum, Frankfurt am Main (SMF 56160, 65162, males, same data as holotype); Museum für Naturkunde, Berlin (ZMB 15697, juvenile, Djur River W Wau, Western Bahr el Ghazal, South Sudan). Derivatio nominis: We dedicate this species to Georg August Schweinfurth (29 December 1836 – 19 September 1925) who was the last of the great explorers of Africa. Schweinfurth collected the oldest of the paratypes (ZMB 15697) during his famous third Africa expedition (1868–1871) “inside the heart of Africa” (Schweinfurth 1874) in present-day South Sudan. Distribution: Central African Republic (Vargas-Ramírez et al. 2010) and South Sudan (this study). Remarks: Pelomedusa schweinfurthi corresponds to mtDNA lineage IV of Vargas-Ramírez et al. (2010), which was previously only recorded from the Central African Republic. Our new 12S sequences of the type specimens provide evidence for the occurrence in South Sudan. The temporal scale is divided on the left side of the head in the holotype, but undivided on the right side, as it is in all paratypes. The largest specimen (SMF 56162) has a straight carapacial length of 15.7 cm. Unlike many other Pelomedusa species, P. schweinfurthi is overall quite dark coloured, with a predominantly brown plastron. Pelomedusa schweinfurthi belongs to the northern species group and is with high support sister to candidate species B from Sudan (Fig. 1).

Petzold, Alice, Vargas-Ramírez, Mario, Kehlmaier, Christian, Vamberger, Melita, Branch, William R., Preez, Louis Du, Hofmeyr, Margaretha D., Meyer, Leon, Schleicher, Alfred, Široký, Pavel, Fritz, Uwe (2014): A revision of African helmeted terrapins (Testudines: Pelomedusidae: Pelomedusa), with descriptions of six new species. Zootaxa 3795 (5): 523-548, DOI: http://dx.doi.org/10.11646/zootaxa.3795.5.2

Pelomedusa candidate species A This candidate species from Cameroon corresponds to mtDNA lineage I of Vargas-Ramírez et al. (2010). It is genetically clearly distinct and differs in 12S and cyt b sequences by uncorrected p distances resembling the divergences among other Pelomedusa species (Tables 1 and 3). Since there is just one genetically verified voucher specimen available (Zoologisches Forschungsmuseum Alexander Koenig, Bonn, ZFMK 15171, Mokolo, Margui- Wandala, Extreme North Province, Cameroon, coll. Wolfgang Böhme, 14–17 February 1974), we refrain from describing this species formally. ZFMK 15171 is a dark-coloured subadult terrapin of 9.3 cm straight carapacial length, resembling in gross morphology P. schweinfurthi or dark-coloured P. variabilis. However, unlike P. schweinfurthi, the pectoral scutes of ZFMK 15171 are triangular and do not reach the plastral midseam. Another terrapin from the same source region was only sampled, but not collected (Museum of Zoology, Senckenberg Dresden, Tissue Collection, MTD T 5183, Maroua, Extreme North Province, Cameroon) and is genetically very similar. Our phylogenetic analyses of mtDNA sequences place candidate species A in the well-supported northern clade of Pelomedusa (Fig. 1). Therein, it is best understood as having a basal position. Pelomedusa candidate species A differs from all other Pelomedusa species by the presence of guanine (G) instead of cytosine (C) at position 212, by the presence of thymine (T) instead of cytosine (C) at position 222, by the presence of cytosine (C) instead of thymine (T) or adenine (A) at position 302, by the presence of cytosine (C) instead of adenine (A) or guanine (G) at position 325, by the presence of thymine (T) instead of adenine (A) or guanine (G) at position 343, and by the presence of adenine (A) instead of cytosine (C) or thymine (T) at position 345 of the 360-bp-long reference alignment of the 12S rRNA gene (Supporting Information). In addition, Pelomedusa candidate species A differs from all species except possibly P. gehafie and P. kobe by the presence of thymine (T) instead of cytosine (C) at position 50 and by the presence of thymine (T) instead of adenine (A) or cytosine (C) at position 65 (character states in P. gehafie and P. kobe unknown)., Published as part of Petzold, Alice, Vargas-Ramírez, Mario, Kehlmaier, Christian, Vamberger, Melita, Branch, William R., Preez, Louis Du, Hofmeyr, Margaretha D., Meyer, Leon, Schleicher, Alfred, Široký, Pavel & Fritz, Uwe, 2014, A revision of African helmeted terrapins (Testudines: Pelomedusidae: Pelomedusa), with descriptions of six new species, pp. 523-548 in Zootaxa 3795 (5) on pages 543-544, DOI: 10.11646/zootaxa.3795.5.2, http://zenodo.org/record/4915083, {"references":["Vargas-Ramirez, M., Vences, M., Branch, W. R., Daniels, S. R., Glaw, F., Hofmeyr, M. D., Kuchling, G., Maran, J., Papenfuss, T. J., Siroky, P., Vieites, D. R. & Fritz, U. (2010) Deep genealogical lineages in the widely distributed African helmeted terrapin: evidence from mitochondrial and nuclear DNA (Testudines: Pelomedusidae: Pelomedusa subrufa). Molecular Phylogenetics and Evolution, 56, 428 - 440. http: // dx. doi. org / 10.1016 / j. ympev. 2010.03.019"]}