taxonID	type	description	language	source
03B68780CF55102A18C3EEA7FB2FF882.taxon	description	Only when Mertens (1937: p. 139) and Loveridge (1941: p. 470) reinstated the scientific name Pelomedusa subrufa (Lacepède, 1788) for helmeted terrapins, this name combination found wide acceptance. Previously, Pelomedusa galeata (Schoepff, 1792) was used by most authors (see below under Testudo galeata Schoepff, 1792), even though John Edward Gray in his influential works consistently applied the species name subrufa (as Hydraspis subrufa, Gray 1831: p. 39 or Pelomedusa subrufa, e. g., Gray 1856: p. 53; 1863: pp. 99 – 100). The name Testudo subrufa was originally coined by Lacepède (1788) in his “ Histoire Naturelle des Quadrupèdes Ovipares ”, where a specimen from the Royal Cabinet Paris was described under the name “ La Roussâtre ” (p. 173). In the “ Synopsis methodica Quadrupedum oviparorum ” of the same work, a folded table in which binominals were applied to the individual species, Lacepède named this terrapin Testudo subrufa. According to Lacepède (1788: p. 173) the terrapin on which he based his description was obtained from Pierre Sonnerat (1745 – 1814) and came allegedly from “ Inde ”. This specimen, which has to be identified with the holotype of Testudo subrufa Lacepède, 1788, is still present in the collection of the Muséum National d’Histoire naturelle (Bour 1982) and was examined by us; tissue for genetic examination was extracted. The type (MNHN 7970) is a shell of an adult terrapin (straight carapacial length 13.67 cm) with most epidermal scutes missing. Its plastron is broken (Fig. 2).	en	Fritz, Uwe, Petzold, Alice, Kehlmaier, Christian, Kindler, Carolin, Campbell, Patrick, Hofmeyr, Margaretha D., Branch, William R. (2014): Disentangling the Pelomedusa complex using type specimens and historical DNA (Testudines: Pelomedusidae). Zootaxa 3795 (5): 501-522, DOI: http://dx.doi.org/10.11646/zootaxa.3795.5.1
03B68780CF55102A18C3EEA7FB2FF882.taxon	materials_examined	It is obvious that the original type locality (“ Inde ”) of Testudo subrufa is in error. In accordance with the information provided by Lacepède (1788), Mertens (1937: p. 139) and Loveridge (1941: p. 470) believed that the type specimen of Testudo subrufa originates from Sonnerat, who is known to have collected at the Cape of Good Hope. Therefore, they identified the type locality with the Cape. However, based on an extensive discussion of historical details, Bour (1982) concluded that the holotype of Testudo subrufa was most probably not collected by Sonnerat, but by Philibert Commerson (1727 – 1773), who stayed on Madagascar in 1770. Some drawings of Commerson’s Malagasy specimens are still present in the Central Library of the Paris Museum. Bour (1982) identified one of these drawings with the holotype and designated “ Taolañaro (Fort-Dauphin), République Malagasy [Madagascar] ” (p. 535) as type locality of Testudo subrufa. However, probably due to oversight, Bour (1985: p. 56) endorsed later the Cape of Good Hope as type locality of Testudo subrufa again. All of our efforts to generate mtDNA sequences from the holotype resulted only in contaminant fungal sequences. Therefore, in the absence of other evidence, we accept the type locality Taolañaro (Madagascar), as suggested by Bour (1982). Consequently, the name Testudo subrufa Bonnaterre, 1789 refers to lineage VIII of Vargas-Ramírez et al. (2010), which is known to occur in Madagascar, Botswana, the Democratic Republic of the Congo, Malawi and Namibia (Fig. 1). The Malagasy populations are thought to be introduced from continental Africa (Vargas-Ramírez et al. 2010; Wong et al. 2010). If lineage VIII is deemed to be taxonomically distinct, the name Pelomedusa subrufa (Bonnaterre, 1789) will have to be restricted to this taxon.	en	Fritz, Uwe, Petzold, Alice, Kehlmaier, Christian, Kindler, Carolin, Campbell, Patrick, Hofmeyr, Margaretha D., Branch, William R. (2014): Disentangling the Pelomedusa complex using type specimens and historical DNA (Testudines: Pelomedusidae). Zootaxa 3795 (5): 501-522, DOI: http://dx.doi.org/10.11646/zootaxa.3795.5.1
03B68780CF56102418C3EC51FDFFF842.taxon	description	The name Testudo galeata was erected by Schoepff (1792: p. 12) in his famous “ Historia Testudinum iconibus illustrata ”. For a long time Testudo galeata was used for helmeted terrapins, until Mertens (1937) and Loveridge (1941) pointed out that the older name Testudo subrufa should be applied. Schoepff’s original description was based on detailed unpublished notes received from two Swedish naturalists. Originally, Schoepff (1792: pp. 12 – 16) intended to assemble information about another terrapin species, Testudo scabra Linnaeus, 1758, a taxon whose identity puzzled scientists for about 250 years (Rhodin & Carr 2009). Schoepff received from Anders Jahan Retzius (1742 – 1821) of Lund a detailed description and drawings of a “ Testudo scabra ” from “ India orientalis ” [East India], which Retzius had kept alive for two years. Schoepff (1792) published Retzius’ description (pp. 13 – 14) and a coloured etching of the terrapin as figure 1 in plate III. Shown under the name “ Test. scabra Retz. ”, it is clearly a juvenile Pelomedusa subrufa (Fig. 3, top), and “ Prof. D. Swartz ” (probably to be identified with Olof Peter Swartz, 1760 – 1818) of Stockholm wrote to Schoepff that a very similar specimen in spirit is present in the Cabinet of the Royal Academy of Sciences at Stockholm. Schoepff (1792: p. 16) concluded that these two terrapins could represent a new species, for which he proposed the name Testudo galeata. The eponymous character (galeata, Latin = helmeted) of this putative new species is its helmet-like head scalation. The two terrapins from Lund and Stockholm have to be regarded as syntypes of Testudo galeata. In the collection of the Biological Museum of Lund University there is still extant a single alcohol-preserved Pelomedusa specimen, bearing the catalogue number ZMUL 6481 (Fig. 3, bottom). Retzius donated this terrapin as “ Testudo scabra ” in 1811 to the Lund Museum, together with a large collection of other specimens (L. Lundqvist, pers. comm.). We conclude that ZMUL 6481, which originates evidently from Retzius, has to be identified with the syntype described in detail in Schoepff (1792: pp. 13 – 14). The size of ZMUL 6481 (straight carapacial length 5.84 cm, width 4.97 cm) approximately matches the measurements given in Schoepff (1792: p. 13) with 2 ½ and 2 Prussian inches (1 Prussian inch = 2.615 cm), especially when it is considered that Retzius could have measured over the curve of the carapace. However, the shell shape of ZMUL 6481 is a bit too oval compared to Schoepff’s figure (Fig. 3, top). Retzius’ two original drawings of the terrapin and part of his letter have survived among Schoepff’s legacy, and are now kept in the Museum für Naturkunde, Berlin (Fig. 3, top). The original drawings are very rough, but they illustrate a shell shape that agrees better with ZMUL 6481 than with Schoepff’s (1792) published figure. In comparison it appears that Schoepff’s figure is somewhat idealized. The coloration of the syntype is much faded due to the age of the specimen and allows no direct comparison with the description in Schoepff (1792).	en	Fritz, Uwe, Petzold, Alice, Kehlmaier, Christian, Kindler, Carolin, Campbell, Patrick, Hofmeyr, Margaretha D., Branch, William R. (2014): Disentangling the Pelomedusa complex using type specimens and historical DNA (Testudines: Pelomedusidae). Zootaxa 3795 (5): 501-522, DOI: http://dx.doi.org/10.11646/zootaxa.3795.5.1
03B68780CF56102418C3EC51FDFFF842.taxon	discussion	In the Swedish Museum of Natural History, Stockholm, there are two old Pelomedusa specimens catalogued as one lot (NRM 7043), which were once part of the collection of Charles de Geer (1720 – 1778). These specimens were transferred by his widow to the collection of the Royal Swedish Academy of Sciences in 1778. The two alcohol-preserved terrapins are in quite bad condition with many shell scutes missing. They are the only helmeted terrapins which were present in the collection when Swartz corresponded with Schoepff, so that one of these specimens is most likely the second syntype (E. Åhlander, pers. comm.). However, as it cannot be unambiguously identified, we choose to designate hereby the Lund specimen as lectotype of Testudo galeata Schoepff, 1792. Hewitt (1935: p. 326) corrected the type locality “ India orientalis ” of Testudo galeata to the environs of Cape Town (and not Cape Flats as thought by Mertens 1937: p. 139, later repeated by Loveridge 1941: p. 470). This makes sense as the Dutch East India Company (Vereenigde Oost-Indische Compagnie) in 1652 established an outpost near the Cape of Good Hope for provisioning their ships coming from East India. This outpost, later becoming the Cape Colony and finally Cape Town, was a supplier of early natural history specimens for Europe (Wren-Sargent 1999; Bauer & Günther 2013; Bauer & Wahlgren 2013), and such specimens might have been mixed up or confused with materials coming from East India. Gray (1831: p. 40) wrote exactly in this spirit about helmeted terrapins “ I have never seen this tortoise from India, but being brought from the Cape by the Indian ships, they, as well as many other Cape animals, are often called Indian. ” Also the morphology of the Lund type, which has pectoral scutes in broad midline contact, is typical for South African terrapins (Hewitt 1935: p. 325; unpubl. observ.), and is thus in agreement with an origin near Cape Town. Therefore, we accept Hewitt’s type locality designation and identify Testudo galeata with Pelomedusa lineage IX, distributed throughout South Africa (Vargas-Ramírez et al. 2010). Two fresh Pelomedusa samples collected close to the Cape of Good Hope (MTD T 5484, Swellendam District; MTD T 5897, Chelance; Table S 1) can be regarded as topotypes of Testudo galeata. As expected, their DNA sequences (Vargas-Ramírez et al. 2010; this study) are assigned with high support to lineage IX in phylogenetic analyses (Fig. 1). “ Die Skizze der T. scabra welche ein Student gemacht hat, lege ich so fehlerhaft sie auch ist bey, indess aus Nr. 1 forma scutellorum u. aus Nr. 2 die Gestalt des Kopfes und der Hinterfüsse wohl zu ersehen ist. Dass meine Testudo die wahre scabra L. ist, dessen bin ich gewiss, ob er [meant is Linnaeus] gleich sagt [illegible] er hat aber ganz gewiss Nr. 73 u. 74 [illegible] completirt, wie das oft der Fall ist. ― Rezius ” [sic]. English translation: “ I enclose the drawing of the T. scabra made by a student, as flawed as it is, since no. 1 shows well the shape of the scutes and no. 2 the shape of the head and of the hind legs. I am sure that my Testudo is the true scabra L., even though he [meant is Linnaeus] says [illegible] he has surely no. 73 and 74 [illegible] completed, as this is often the case. ― Rezius ” [sic].	en	Fritz, Uwe, Petzold, Alice, Kehlmaier, Christian, Kindler, Carolin, Campbell, Patrick, Hofmeyr, Margaretha D., Branch, William R. (2014): Disentangling the Pelomedusa complex using type specimens and historical DNA (Testudines: Pelomedusidae). Zootaxa 3795 (5): 501-522, DOI: http://dx.doi.org/10.11646/zootaxa.3795.5.1
03B68780CF56102418C3EC51FDFFF842.taxon	materials_examined	Bottom: Dorsal and ventral view of the lectotype of Testudo galeata Schoepff, 1792 (ZMUL 6481, straight carapacial length 5.84 cm). Scale bar, 3 cm. Photos: L. Lundqvist.	en	Fritz, Uwe, Petzold, Alice, Kehlmaier, Christian, Kindler, Carolin, Campbell, Patrick, Hofmeyr, Margaretha D., Branch, William R. (2014): Disentangling the Pelomedusa complex using type specimens and historical DNA (Testudines: Pelomedusidae). Zootaxa 3795 (5): 501-522, DOI: http://dx.doi.org/10.11646/zootaxa.3795.5.1
03B68780CF59102518C3EBA1FD87FCDD.taxon	description	Schweigger (1812: p. 307) described a carapace from the Adanson Collection of the Paris Museum as the new species Emys olivacea. According to the original description, the specimen originates “ in sabulosis Nigritiae ” [in the sands of Senegal] and is still present in the collection of the Muséum National d’Histoire naturelle, Paris (MNHN 7971). This specimen was not available to us. Starting with Loveridge (1941), many authors used the name Pelomedusa subrufa olivacea for helmeted terrapins from the more northern parts of the species’ range, which usually have the pectoral scutes of the plastron separated, even though this character cannot be determined in the holotype, which consists only of a carapace. Moreover, Gasperetti et al. (1993) disputed the reliability of this trait, and since then most authors have treated P. subrufa as monotypic (see above and the review in Boycott & Bourquin 2008). In our phylogenetic analyses (Fig. 1), the concatenated 12 S and cyt b sequences (Wong et al. 2010; this study) of a Senegalese Pelomedusa (ZFMK 17076) are assigned with high support to lineage III of Vargas-Ramírez et al. (2010). The placement of this topotypic terrapin suggests that Emys olivacea has to be identified with lineage III, which is otherwise known to occur in Benin, Burkina Faso, Niger, and Nigeria (Vargas-Ramírez et al. 2010; Wong et al. 2010). The pectoral scutes of ZFMK 17076 are triangular, but the tips just meet in the midline of the plastron. A comparison of 21 helmeted terrapins from Niger, Nigeria, and Senegal from the collections of the natural history museums in Bonn, Frankfurt am Main, London, Port Elizabeth, and Vienna shows that specimens with pectoral scutes in narrow contact, as well as specimens with completely separated pectorals, both occur in lineage III.	en	Fritz, Uwe, Petzold, Alice, Kehlmaier, Christian, Kindler, Carolin, Campbell, Patrick, Hofmeyr, Margaretha D., Branch, William R. (2014): Disentangling the Pelomedusa complex using type specimens and historical DNA (Testudines: Pelomedusidae). Zootaxa 3795 (5): 501-522, DOI: http://dx.doi.org/10.11646/zootaxa.3795.5.1
03B68780CF59102518C3E8EBFEDAF9D1.taxon	description	Based on all Pelomedusa specimens present in the collection of the Paris Museum upon the time of the species description, Duméril & Bibron (1835: pp. 390 – 394) erected Pentonyx capensis. The series of syntypes comprised helmeted terrapins from the eponymous South African Cape Region and from Madagascar, but also the holotype of Emys olivacea from Senegal, which was considered by Duméril & Bibron to represent merely a juvenile Pentonyx capensis. We examined one of the syntypes, MNHN 9506 originating from “ Le Cap ”, and were allowed to extract tissue for genetic investigation. MNHN 9506 is an adult mounted male in good condition with a straight carapacial length of 23.45 cm (Fig. 4). Mertens (1937: p. 139) restricted the type locality of Pentonyx capensis to the Cape of Good Hope. However, because no lectotype was designated this action was invalid (cf. ICZN 1999: Art. 76). Nevertheless, later authors identified Pentonyx capensis implicitly or explicitly with the nominotypical subspecies of the helmeted terrapin, if the species was treated as polytypic (Mertens 1937; Loveridge 1941; Bour 1986; Ernst & Barbour 1989; Iverson 1992). To stabilize this usage, we designate herewith MNHN 9506 as lectotype of Pentonyx capensis. For the lectotype, mtDNA fragments of all three marker genes could be sequenced (Table 2). In phylogenetic analyses (Fig. 1), the concatenated sequences of MNHN 9506 are embedded within other South African sequences representing lineage IX sensu Vargas-Ramírez et al. (2010). Thus, if lineage IX should be deemed taxonomically distinct, Pentonyx capensis Duméril & Bibron, 1835 would become a subjective junior synonym of Testudo galeata Schoepff, 1792.	en	Fritz, Uwe, Petzold, Alice, Kehlmaier, Christian, Kindler, Carolin, Campbell, Patrick, Hofmeyr, Margaretha D., Branch, William R. (2014): Disentangling the Pelomedusa complex using type specimens and historical DNA (Testudines: Pelomedusidae). Zootaxa 3795 (5): 501-522, DOI: http://dx.doi.org/10.11646/zootaxa.3795.5.1
03B68780CF59102018C3EDFCFBBFFB03.taxon	description	Rüppell (1835: pp. 2 – 3) described Pentonyx gehafie based on terrapins which he found “ häufig in allen fliessenden oder stagnirenden Gewässern auf dem östlichen Abhange der abyssinischen Küstengebirge ” [frequently in all flowing and stagnant waters on the eastern slope of the Abyssinian coastal mountains]. Rüppell diagnosed Pentonyx gehafie from the Pentonyx species occurring at the Cape of Good Hope by the plastral scutation: The pectoral scutes of his new species are triangular and do not reach the median plastral seam, whereas in the Cape species the pectoral scutes are in contact. The character state of the Abyssinian specimens is nicely depicted in Rüppell’s plate 1. Obviously Rüppell collected quite a number of helmeted terrapins, which originally had all syntype status. Mertens (1937: p. 140) mentions that seven of these specimens are in the Senckenberg Museum, Frankfurt am Main. However, there are eight, and not seven, terrapins collected by Rüppell with identical data (SMF 7947 – 7949, SMF 7953, SMF 7960 – 7962, SMF 33054). Whilst SMF 7947 – 7949 and SMF 7960 – 7962 are alcohol-preserved, SMF 7953 and SMF 33054 are dry specimens (Table 2). SMF 7947 was designated by Mertens (1937: p. 140) as lectotype, which is why the remaining terrapins bear today the status of paralectotypes. Further paralectotypes collected by Rüppell are in the Natural History Museum, London (Gray 1856: p. 53; Boulenger 1889: p. 199) and the Muséum National d’Histoire naturelle, Paris (see below under Pelomedusa galeata var. disjuncta Vaillant & Grandidier, 1910). The lectotype is a subadult female of 11.42 cm straight carapacial length (Fig. 5, top). Mertens (1937: p. 140) restricted the type locality of Pentonyx gehafie to “ Massaua ” [also known as Massawa, Missiwa or Mitsiwa, Eritrea], a town within the source region of the type specimens. However, according to Article 76.1 of the International Code of Zoological Nomenclature (ICZN 1999), the type locality “ is the geographical […] place of capture, collection or observation of the name-bearing type. ” Thus, Rüppell’s original type locality has to remain unchanged and is to be identified with the eastern slope of coastal mountains, Eritrea. Boulenger (1880: p. 151) synonymized Pelomedusa gehafie with Pelomedusa galeata (= Pelomedusa subrufa) because he studied specimens with intermediate plastral characters from Madagascar, the Upper Nile Region, and Zanzibar, even though he acknowledged that separated triangular pectorals never occur in South African terrapins and that in Abyssinia the terrapins have consistently widely separated triangular pectorals. Based on similar observations, but drawing a completely different conclusion, Hewitt (1935: p. 325) recognized “ Pelomedusa gehafiae ” [sic] as a valid species and opposed it to his polytypic species Pelomedusa galeata. Hewitt noted that at Mt. Elgon (border region of Uganda and Kenya) helmeted terrapins with separated pectorals are found together with terrapins having the pectorals in contact. Finally, based on the occurrence of morphologically intermediate terrapins in some regions, Parker (1936: p. 609) and Mertens (1937: p. 140) recognized Pentonyx gehafie as a valid subspecies of the helmeted terrapin and used the names Pelomedusa galeata gehafie and Pelomedusa subrufa gehafie, respectively. In a similar vein, Loveridge (1941: p. 480) accepted the validity of a subspecies characterized by separated triangular pectoral scutes, but placed Pentonyx gehafie in the synonymy of Pelomedusa subrufa olivacea (Schweigger, 1812). Summarizing his own observations and those of previous authors, Loveridge believed that this taxon occurs in “ the drier regions of a belt extending from Senegal to Eritrea, intergrading with the typical form in the Anglo-Egyptian Sudan, Ethiopia, Somaliland, northern Kenya and Uganda ” (p. 481). Later authors identified Pelomedusa subrufa olivacea generally with the more northern populations of the helmeted terrapin, including the populations of the Arabian Peninsula (cf. Ernst & Barbour 1989; Iverson 1992; Ernst et al. 2000; Boycott & Bourquin 2008), until the usage of subspecies was abandoned after Gasperetti et al. (1993).	en	Fritz, Uwe, Petzold, Alice, Kehlmaier, Christian, Kindler, Carolin, Campbell, Patrick, Hofmeyr, Margaretha D., Branch, William R. (2014): Disentangling the Pelomedusa complex using type specimens and historical DNA (Testudines: Pelomedusidae). Zootaxa 3795 (5): 501-522, DOI: http://dx.doi.org/10.11646/zootaxa.3795.5.1
03B68780CF59102018C3EDFCFBBFFB03.taxon	materials_examined	For the present investigation, we studied the lectotype and the nine paralectotypes of Pentonyx gehafie in Frankfurt am Main and London; all specimens conform well to the description by Rüppell (1835) in having widely separated triangular pectoral scutes (Fig. 5). Tissue for genetic examination was extracted from the eight Frankfurt specimens, but DNA sequences could be generated only for three terrapins, among them the lectotype (Table 2).	en	Fritz, Uwe, Petzold, Alice, Kehlmaier, Christian, Kindler, Carolin, Campbell, Patrick, Hofmeyr, Margaretha D., Branch, William R. (2014): Disentangling the Pelomedusa complex using type specimens and historical DNA (Testudines: Pelomedusidae). Zootaxa 3795 (5): 501-522, DOI: http://dx.doi.org/10.11646/zootaxa.3795.5.1
03B68780CF59102018C3EDFCFBBFFB03.taxon	discussion	In phylogenetic analyses (Fig. 1), the sequences of the type specimens constitute a deeply divergent clade being sister to Kenyan terrapins representing lineage V sensu Vargas-Ramírez et al. (2010). We examined 18 Pelomedusa from Kenya morphologically (7 live terrapins and 11 specimens in the collections of the natural history museums of Berlin, Bonn, London, and Vienna) and these differ consistently from the type series of Pentonyx gehafie in that their pectoral scutes are in wide contact in the midline, suggesting that Pentonyx gehafie could be a distinct taxon. At present, we refer to the genetic lineage of the type specimens of Pentonyx gehafie as lineage X, and do not relate it to any of the lineages identified by Vargas-Ramírez et al. (2010).	en	Fritz, Uwe, Petzold, Alice, Kehlmaier, Christian, Kindler, Carolin, Campbell, Patrick, Hofmeyr, Margaretha D., Branch, William R. (2014): Disentangling the Pelomedusa complex using type specimens and historical DNA (Testudines: Pelomedusidae). Zootaxa 3795 (5): 501-522, DOI: http://dx.doi.org/10.11646/zootaxa.3795.5.1
03B68780CF5C102018C3EFD0FE60F95B.taxon	description	Cornalia (1849: pp. 312 – 313) described a specimen allegedly from “ Novaeboracum ” [= Novum Eboracum, New York] under this name. After its description, Pentonix americana has never been regarded as a valid taxon. It is traditionally identified with the helmeted terrapin (e. g., Boulenger 1889: p. 198; Mertens 1937: p. 139; Loveridge 1941: p. 473; Wermuth & Mertens 1961: p. 284, 1977: p. 115; Fritz & Havaš 2007: p. 345; van Dijk et al. 2012: p. 000.295), even though Strauch (1865: p. 111) doubted that it could be reliably assigned to any species. The type specimen was in the collection of the natural history museum of Milan (Museo Civico di Storia Naturale di Milano) and was destroyed, together with most of the herpetological collection, by British air raids in 1943 (S. Scali, pers. comm.).	en	Fritz, Uwe, Petzold, Alice, Kehlmaier, Christian, Kindler, Carolin, Campbell, Patrick, Hofmeyr, Margaretha D., Branch, William R. (2014): Disentangling the Pelomedusa complex using type specimens and historical DNA (Testudines: Pelomedusidae). Zootaxa 3795 (5): 501-522, DOI: http://dx.doi.org/10.11646/zootaxa.3795.5.1
03B68780CF5C102118C3EC72FEE6FD26.taxon	description	Based on three syntypes from Natal, Gray (1863) described the new species Pelomedusa nigra. Boulenger (1889) lists these specimens as k, l and m in his “ Catalogue of the Chelonians, Rhynchocephalians, and Crocodiles of the British Museum (Natural History) ”. Today, these mounted dry specimens are catalogued under the numbers BMNH 1849.1.30.27 and BMNH 1862.12.4.4 – 5 (the latter re-registered as BMNH 1947.3.5.80 – 81). Pelomedusa nigra was soon synonymized with Pelomedusa galeata (Strauch 1865: p. 111; Boulenger 1880: p. 151, 1889: p. 198) or Pelomedusa subrufa (Mertens 1937: p. 139; Loveridge 1941: p. 474), until Bour (1986: p. 37) resurrected it in a footnote as a subspecies of Pelomedusa subrufa. However, following Gasperetti et al. (1993) most authors abandoned the usage of any subspecies of Pelomedusa subrufa (cf. Boycott & Bourquin 2008). The three syntypes of Pelomedusa nigra were examined for the present study and tissue for genetic investigation was extracted. We designate hereby the oldest specimen, BMNH 1849.1. 30.27 (adult male, straight carapacial length 16.19 cm; Fig. 6, top), as lectotype of this taxon. A fragment of the 12 S rRNA gene could be sequenced from all three type specimens. In phylogenetic analyses (Fig. 1), these sequences cluster with high support within lineage IX from South Africa. Moreover, the three type sequences are identical with a 12 S sequence of a fresh sample from the same geographical source region (MTD T 5509, Ndumo, KwaZulu-Natal). Since we identify Testudo galeata Schoepff, 1792 with lineage IX, Pelomedusa nigra Gray, 1863 would become a subjective junior synonym of that name, if lineage IX should be considered taxonomically distinct.	en	Fritz, Uwe, Petzold, Alice, Kehlmaier, Christian, Kindler, Carolin, Campbell, Patrick, Hofmeyr, Margaretha D., Branch, William R. (2014): Disentangling the Pelomedusa complex using type specimens and historical DNA (Testudines: Pelomedusidae). Zootaxa 3795 (5): 501-522, DOI: http://dx.doi.org/10.11646/zootaxa.3795.5.1
03B68780CF5D102118C3E9E1FEE6FAF0.taxon	description	Rochebrune (1884: pp. 25 – 26; plate 1: figs 1 – 2) described helmeted terrapins from several sites in Senegal (“ Dagana, Saidé, lac de N’Guer, marigot des Maringouins ” [the latter meaning mosquito oxbows of N’Gor lake, Dakar, Senegal]) as his new species Pelomedusa gasconi, which was later synonymized by Loveridge (1941: p. 480) with Pelomedusa subrufa olivacea. In accordance with this assignment, Rochebrune’s (1884) plate 1 shows a helmeted terrapin with widely separated pectoral scutes, although Rochbrune explained that this character may be variable (pp. 26 – 27). According to Angel (in Loveridge 1941: p. 480), there was no type specimen of Pelomedusa gasconi preserved. Loveridge (1941: p. 480) restricted the type locality to Dagana, Senegal, which is invalid without lectotype or neotype designation (cf. ICZN 1999: Art. 76). The sites of the composed type locality are up to 400 km distant from one another. To restrict the type locality and to associate the name Pelomedusa gasconi Rochebrune, 1884 unambiguously with a genetic lineage of Pelomedusa, we designate in accordance with Article 75 (ICZN 1999) a topotypic specimen from Dakar as its neotype (ZFMK 17076, a subadult female of 11.18 cm straight carapacial length in alcohol; Fig. 6, bottom). For the neotype, mtDNA sequences are available (see also above under Emys olivacea Schweigger, 1812). Thus, the name Pelomedusa gasconi is to be identified with the same genetic lineage as Emys olivacea (lineage III of Vargas- Ramírez et al. 2010; Fig. 1) and becomes a subjective junior synonym of the latter name, if lineage III is deemed taxonomically distinct.	en	Fritz, Uwe, Petzold, Alice, Kehlmaier, Christian, Kindler, Carolin, Campbell, Patrick, Hofmeyr, Margaretha D., Branch, William R. (2014): Disentangling the Pelomedusa complex using type specimens and historical DNA (Testudines: Pelomedusidae). Zootaxa 3795 (5): 501-522, DOI: http://dx.doi.org/10.11646/zootaxa.3795.5.1
03B68780CF5D102218C3EE9FFF8EFD6D.taxon	description	This taxon was treated by Loveridge (1941: p. 480) as a junior synonym of Pelomedusa subrufa olivacea. It was originally described by Vaillant & Grandidier (1910: pp. 55 – 56) as a new variety. These authors compared helmeted terrapins from Madagascar and the Cape of Good Hope with specimens of “ Pentonyx gehafie ” from the collection of the Muséum National d’Histoire naturelle, highlighting that the latter differ from Malagasy and South African individuals by separated pectoral scutes. Vaillant & Grandidier (1910: p. 56) assigned the terrapins with the separated pectorals to their new variety Pelomedusa galeata var. disjuncta and mention expressis verbis that specimens of Rüppell’s “ Pentonyx gehafie ” fitting this character are present in the holdings of the Paris Museum. Consequently, it is clear that the description of Pelomedusa galeata var. disjuncta was based on several syntypes. In addition to Rüppell’s specimens, Vaillant & Grandidier (1910: p. 56) mention another terrapin collected by Vicomte Pierre Marie Robert Du Bourg de Bozas (1871 – 1902) as representing their new variety. Based on this information and locality data from the Paris Museum provided by Fernand Angel, Loveridge (1941: p. 480) restricted the type locality of Pelomedusa galeata var. disjuncta to the “ shore of Lake Abaya, Sidamo, Ethiopia ”, where the specimen of Du Bourg de Bozas had been collected. However, this type locality restriction is invalid without lectotype designation (cf. ICZN 1999: Art. 76). The specimen in question is still present in the collection of the Muséum National d’Histoire naturelle (MNHN 1902.346, “ Bord du lac Abbay, Sidamo, Ethiopie ”). In addition, there are also two specimens labelled as syntypes 1 of Pentonyx gehafie (MNHN 7870, 9398, “ Abyssinie ”), which bear also syntype status for Pelomedusa galeata var. disjuncta. Consequently, the latter taxon was based on three syntypes and the type locality encompasses distinct localities, the shore of Lake Abaya, Sidamo, Ethiopia, and Abyssinia. Abyssinia can be more exactly identified with the eastern slope of the coastal mountains in present-day Eritrea, where Rüppell (1835: p. 3) discovered his specimens (see also above under Pentonyx gehafie Rüppell, 1835). The syntypes of Pelomedusa galeata var. disjuncta were not available for study. However, since mtDNA sequences could be produced for other type specimens of Pentonyx gehafie (see above), the identity of terrapins from the coastal mountain region of Eritrea could be clarified genetically. Therefore, we designate hereby the topotypic specimen MNHN 7870 from “ Abyssinie ”, being simultaneously a paralectotype of Pentonyx gehafie Rüppell, 1835, as lectotype of Pelomedusa galeata var. disjuncta Vaillant & Grandidier, 1910. By this action, we associate the latter name unambiguously with a defined genetic lineage of Pelomedusa. Thus, if the distinct genetic lineage matching the name-bearing types of Pentonyx gehafie and Pelomedusa galeata var. disjuncta should be deemed taxonomically distinct, the latter name would be automatically a subjective junior synonym of Pentonyx gehafie.	en	Fritz, Uwe, Petzold, Alice, Kehlmaier, Christian, Kindler, Carolin, Campbell, Patrick, Hofmeyr, Margaretha D., Branch, William R. (2014): Disentangling the Pelomedusa complex using type specimens and historical DNA (Testudines: Pelomedusidae). Zootaxa 3795 (5): 501-522, DOI: http://dx.doi.org/10.11646/zootaxa.3795.5.1
03B68780CF5E102218C3E81AFBDFFA61.taxon	description	This subspecies was based on two syntypes (shells) from “ Quickborn, near Okahandja, South West Africa ” [Namibia] (Hewitt 1935: p. 338). Actually, Quickborn is a farm approximately halfway between Okahandja and Otjiwarongo, Namibia. Following Mertens (1937: p. 139) and Loveridge (1941: p. 474), Pelomedusa galeata damarensis was later synonymized with the nominotypical subspecies of the helmeted terrapin, if the species was regarded as polytypic. Herrmann & Branch (2013: p. 102) suggested that Pelomedusa galeata damarensis is morphologically distinctive and may deserve specific recognition. The two syntypes of Pelomedusa galeata damarensis are now in the collection of the Port Elizabeth Museum (PEM R 14953 – R 14954) and were examined for the present study. Whilst PEM R 14953 is perfectly preserved, parts of the epidermal scutes on the right carapacial margin of PEM R 14954 are missing. We designate herewith the better preserved specimen PEM R 14953 (Fig. 7, top) as lectotype of Pelomedusa galeata damarensis. Tissue for genetic investigation was extracted from both type specimens. From PEM R 14953, 252 bp of the 12 S rRNA gene could be successfully sequenced, whereas only 35 bp of the 12 S rRNA gene and 319 bp of the cyt b gene could be generated for PEM R 14954 (Table 2). In phylogenetic analyses (Fig. 1), the sequences of both type specimens are embedded with high support within lineage VIII of Vargas-Ramírez et al. (2010). The oldest available name for lineage VIII is Testudo subrufa Bonnaterre, 1789 (see above), rendering Pelomedusa galeata damarensis Hewitt, 1935 a subjective junior synonym of the previous name, if lineage VIII should be regarded as taxonomically distinct.	en	Fritz, Uwe, Petzold, Alice, Kehlmaier, Christian, Kindler, Carolin, Campbell, Patrick, Hofmeyr, Margaretha D., Branch, William R. (2014): Disentangling the Pelomedusa complex using type specimens and historical DNA (Testudines: Pelomedusidae). Zootaxa 3795 (5): 501-522, DOI: http://dx.doi.org/10.11646/zootaxa.3795.5.1
03B68780CF5E102218C3ED32FA5AF80D.taxon	description	The description of this subspecies was based on a single shell from “ Besondermeid, Steinkopf, Namaqualand, C. P. ” [Cape Province, now Northern Cape, South Africa] (Hewitt 1935: p. 337). Following Mertens (1937: p. 139) and Loveridge (1941: p. 474), Pelomedusa galeata devilliersi was later synonymized with the nominotypical subspecies of the helmeted terrapin, if the species was regarded as polytypic. The holotype of Pelomedusa galeata devilliersi is now in the collection of the Port Elizabeth Museum (PEM R 14962) and was examined for this study (Fig. 7, bottom). Tissue for genetic investigation was extracted and mtDNA fragments of all three marker genes could be sequenced (Table 2). In phylogenetic analyses (Fig. 1), the concatenated sequences are associated with high support with other South African samples representing lineage IX sensu Vargas-Ramírez et al. (2010), and thus Pelomedusa galeata devilliersi Hewitt, 1935 would become a junior synonym of Testudo galeata Schoepff, 1792, if lineage IX should be regarded as taxonomically distinct.	en	Fritz, Uwe, Petzold, Alice, Kehlmaier, Christian, Kindler, Carolin, Campbell, Patrick, Hofmeyr, Margaretha D., Branch, William R. (2014): Disentangling the Pelomedusa complex using type specimens and historical DNA (Testudines: Pelomedusidae). Zootaxa 3795 (5): 501-522, DOI: http://dx.doi.org/10.11646/zootaxa.3795.5.1
03B68780CF41103D18C3EE57FDA9F849.taxon	description	Hewitt (1935: pp. 332 – 335) based the description of his new subspecies Pelomedusa galeata orangensis on several specimens from distinct collecting sites in South Africa. However, on page 333 he explicitly named a male shown in his plate XXXII, fig. 4, “ the type ”, so that it is clear that this specimen represents in accordance with Article 73.1 of the International Code of Zoological Nomenclature (ICZN 1999) the holotype of Pelomedusa galeata orangensis. This terrapin was in the Kimberley Museum (now McGregor Museum, Kimberley) and originates “ presumably from the Kimberley neighbourhood ” (Hewitt 1935: p. 333). Following Mertens (1937: p. 139) and Loveridge (1941: p. 474), Pelomedusa galeata orangensis was later synonymized with the nominotypical subspecies of the helmeted terrapin, if the species was regarded as polytypic. The holotype of Pelomedusa galeata orangensis could not be located in the McGregor Museum (B. Wilson, pers. comm.), and must be considered lost. Efforts to sequence mtDNA of two paratypes from the collection of the Port Elizabeth Museum (PEM R 9404, Warrenton, Northern Cape; PEM R 9408, Thaba ‘ Nchu, Free State) failed. However, genetic data were available for several South African localities encircling the type locality Kimberley (Vargas-Ramírez et al. 2010; this study), and all South African sequences correspond to lineage IX sensu Vargas- Ramírez et al. (2010). Consequently, we identify Pelomedusa galeata orangensis with this lineage. If lineage IX should be deemed taxonomically distinct, Pelomedusa galeata orangensis Hewitt, 1935 becomes a subjective junior synonym of Testudo galeata Schoepff, 1792.	en	Fritz, Uwe, Petzold, Alice, Kehlmaier, Christian, Kindler, Carolin, Campbell, Patrick, Hofmeyr, Margaretha D., Branch, William R. (2014): Disentangling the Pelomedusa complex using type specimens and historical DNA (Testudines: Pelomedusidae). Zootaxa 3795 (5): 501-522, DOI: http://dx.doi.org/10.11646/zootaxa.3795.5.1
03B68780CF42103E18C3EBA1FD66FD95.taxon	description	Based on five Malagasy specimens in the collection of the Senckenberg Museum, Mertens (1937: p. 141) described the subspecies Pelomedusa subrufa wettsteini. The holotype (SMF 7958) is an alcohol-preserved male from “ Majunga, West-Madagaskar ” [Mahajanga, western Madagascar] with a straight carapacial length of 12.26 cm (Fig. 8). The taxon was soon synonymized with the nominotypical subspecies of the helmeted terrapin (Loveridge 1941: p. 474). The type specimens were examined for this study, but tissue was not extracted because many DNA sequence data are available for Malagasy Pelomedusa (Vargas-Ramírez et al. 2010; Wong et al. 2010; Fig. 1), which represent lineage VIII sensu Vargas-Ramírez et al. (2010). Thus, Pelomedusa subrufa wettsteini Mertens, 1937 would become a subjective junior synonym of Testudo subrufa Bonnaterre, 1789, if lineage VIII should be considered taxonomically distinct.	en	Fritz, Uwe, Petzold, Alice, Kehlmaier, Christian, Kindler, Carolin, Campbell, Patrick, Hofmeyr, Margaretha D., Branch, William R. (2014): Disentangling the Pelomedusa complex using type specimens and historical DNA (Testudines: Pelomedusidae). Zootaxa 3795 (5): 501-522, DOI: http://dx.doi.org/10.11646/zootaxa.3795.5.1
