Palaeont. afr., 36,43-82 (2000) AFRICAN CHELONIANS FROM THE JURASSIC TO THE PRESENT: PHASES OF DEVELOPMENT AND PRELIMINARY CATALOGUE OF THE FOSSIL RECORD. by France de Lapparent de Broin Museum national d ’histoire naturelle, Laboratoire de Paleontologie -UMR 8569 du CNRS. 8, rue Buffon, 75005 Paris, France. ABSTRACT The five major phases in the palaeontological history of African chelonians are presented: 1) autochthonous development of the north Gondwanan pleurodires from a Pangean source group; 2) littoral expansion of a member of this group (Bothremydidae), accompanied by the arrival of Laurasian marine turtles; 3) in situ development of pleurodires and the immigration of Eurasian cryptodires (Oligo-Miocene) traversing the Tethys in several waves; 4) great diversification and endemism (Pliocene to Holocene); 5) important faunal reduction due to climatic changes at the end of Holocene times (cooling, aridification); elsewhere, great speciation and arrival during the Present of the last European immigrant in the north. Throughout the period under consideration there were several reductions in taxonomic diversity and emigrations from Africa. A preliminary catalogue of the fossil record of African chelonians is given, presented country by country followed by a taxonomic listing. KEYWORDS: Fossil chelonians, Africa, stratigraphy, taxonomy INTRODUCTION This work is based on data collected for a presentation on the settlement of chelonians in Africa which I gave at the PSSA’98 conference in Windhoek, Namibia (Lapparent de Broin 1998). It is not the result of a systematic study of African chelonians and the catalogue presented here does not pretend to be complete but it attempts to include all named taxa. Some data on Pliocene to more recent archaeological sites are not included, in particular those published recently (from 1998), nor are references to chelonians in more general works where descriptions of the forms dealt with are not included. However, the data presented here can reasonably claim to document the spread of chelonians in Africa. The classification adopted here is based on phylogeny (Gaffney & Meylan 1988, emend. Bour& Dubois 1986; Broin 1988a,b, Lapparent de Broin & Murelaga 1999; Lapparent de Broin & Werner 1998) ; it reflects the diversification of taxa as well as their palaeo- biogeographic history (continental drift, geographic barriers, changes in climate) from the Triassic to the Present. Phenetic systematics is rejected in this study; such classifications are still employed by a number of neontologists who establish compilations only for the practical purposes of determination and distribution of extant forms, without the necessity of a historical analysis (e.g. David 1994; Iverson 1992). They include artificial polyphyletic taxa. However, it should be noted that, in the classifications adopted in this study, parts are not fully phylogenetic because several proposed cladograms do not integrate the fossil forms (except for a few genera) (Bour 1985; Gaffney & Meylan 1988; Hirayama 1985), as for example with two important African taxa: Trionychidae and Testudinidae. Step by step, the suppression of artificial genera is in progress, as has happened with the partition of Clemmys, Trionyx (based on extant forms only), Testudo s.s. and Agrionemys (a part of Testudo s.l.) and Podocnemis, but not yet with Geochelone, probably the last artificial taxon (with the remaining part of Testudo s.l.). At family level, the partition of the Pelomedusidae in the several families constituting the hyperfamily Pelomedusoides, is also widely admitted by the scientific community. Environment An important point about the analysis of the African chelonians is recognition of the role of the environment in the geographic spread of taxa. In the catalogue that follows, the environment is specified in the catalogue only when it is littoral or marine: the term ‘littoral’ is assigned to some pleurodire turtles, e.g. the Bothremydidae which lack paddled limbs, and followed coastlines, not crossing wide seaways. The term ‘marine’ is attributed to the cryptodires Chelonioidea and Dermochelyoidea (here separated for better comprehension), which have paddled limbs, are able to cross oceans and to live in deep seas. They are also found, as fossils, in littoral palaeoenvironments. Originally they were continental and then become littoral (as did the Bothremydidae), before they conquered deep seas. The other chelonian taxa are continental, i.e. either freshwater (the majority), sometimes possibly semi aquatic-semi terrestrial, or fully terrestrial, similar to the primitive Triassic chelonians and their pareiasaurid ancestors PALAEONTOLOGIA AFRICANA VOLUME 36 - 2000 D 44 (see Lee 1997). Thus, in Africa, the oldest known form, the very primitive Hettangian South African taxon Australochelys from the Stormberg group of the Karoo, was unquestionably a continental form. In the Testudininei the relatively large forms are also able to cross short seaways, floating in suitable currents (Bour 1985, 1987, 1994, conquest of Indian Ocean islands). Some small freshwater forms, such as Pelomedusa or Pelusios were probably carried on floating debris, by currents during typhoons, or tsunamis, or freshwater floods and brackish water currents and were able to colonize Indian Ocean islands separated by considerable distances from the African mainland. The small freshwater forms have never been known to cross oceans unaided and in order to spread they utilised a network of freshwater rivers, lakes and ponds. The Trionychoidea are freshwater turtles with paddled limbs and they are able to cross a short seaway or to follow a coastline, from the mouth of the home river, being carried by inshore currents to enter other rivers farther down the coast, as observed for example in Trionyx in western Africa (Hughes pers. comm.) and various observations in the Mediterranean and colonization of Figure 1. Countries of Africa, the Arabian Peninsula and the vicinity ofMadagascar, with a record of fossil chelonians. Africa: AL, Algeria; AN, Angola; CVI, Cape Verde Islands; CB, Congo: People’s Republic of (ex Congo- Brazzaville); CH, Chad; CZ, Congo: Democratic Republic of (ex Zaire, Congo-Kinshasa); D, Djibouti: Republic of; EG, Egypt; ET, Ethiopia; G, Ghana; K, Kenya; L, Libya; LO, Lesotho; MA, Mali; MAL, Malta; MAU, Mauritania; MO, Morocco; MW, Malawi; MZ, Mozambique; NA, Nigeria; NI, Niger; NM, Namibia; SA, South Africa; SE, Senegal; SO, Somalia; SU, Sudan; TA, Tanzania; TU, Tunisia; U, Uganda; Z, Zimbabwe. Arabian Peninsula: SAA, Saudi Arabia; AD, Abu Dhabi: Emirate of, United Arab Emirates; O, Oman: Sultanate of. Madagascar area: M, Madagascar and Gloriosa; A, Aldabra and SE Y, Seychelles islands; CO, Comoros islands (Mayotte), MAS, Mascarene islands: La Reunion, Mauritius, Rodrigues. New Guinea and Australia by Trionychidae and Carettochelyidae. PHASES OF DEVELOPMENT Fossil chelonians are known from 32 African- Arabian countries and seven groups of islands (Figure 1). Arabia (the Arabian Peninsula), Madagascar, the surrounding Indian Ocean islands, Malta and Cape Verde Islands are an integral part of the African domain (Figures 7 and 8). The Canary Islands and Mediterranean islands other than Malta have not been integrated into this study, although they share faunas which are similar in part to those from the northern part of Africa. Fossil African chelonians are known from the earliest Jurassic until the Present. The extensive geographic and stratigraphic data now available allow a very close idea of the truth concerning the progressive colonization of the continent by chelonians. Study of the taxa already defined shows that Africa was initially populated by primitive forms, already present during the Pangean period, very soon after the appearance of the first known chelonians (Norian-Keuper). Continental drift then separated the land masses. Until the relatively recent arrival of Eurasian forms during the Oligocene, Africa was isolated during the Cretaceous and Palaeogene as far as continental forms of chelonians are concerned. Africa was still linked to India during Cretaceous times at least and had filtered relations with southwestern Europe during Late Cretaceous-Early Palaeogene times. The Pleurodira (chelonians with a pelvis linked by sutures to the shell and which progressively acquired a neck retracting in an horizontal plane) principally developed in the territories of the fragmented Gondwana (although some forms, e.g. the Dortokidae, evolved in Europe, Lapparent de Broin & Murelagal996,1999; Gheerbrant etal. 2000), while the Cryptodira (which progressively acquired a neck retracting in a vertical plane) were spreading in Laurasia. But Cryptodira progressively immigrated into Africa, in waves, to the point where they now comprise the majority of the African chelonian fauna. First phase: autochthonous development from a Pangean group First chelonians known in the world The earliest known chelonians are Late Triassic (Keuper-Norian) but it is not possible to say which is the oldest taxon (Figure 2). The form that seems the most primitive (taken as a whole, because, apart from its primitive traits, it also has derived characters), is Proganochelys quenstedti Baur 1887, Germany (Fraas 1899; Gaffney 1990; Jaekel 1918), but this form, which is placed in its own infraorder, is not the oldest in the German Triassic. The supposedly related forms, aff. Proganochelys sp., from the Norian of Greenland (Jenkins et al. 1994) and aff. P. ruchae Broin 1985 (Broin et al. 1982), from the Norian of Thailand, are poorly known but they appear a little more derived. They share the character of epiplastral points, only four rather than five in Proganochelys, that are flattened and 45 laterally diverging, rather than rounded at the base and directed forwards. Proter ocher sis robusta Fraas 1913, is from the same German beds as P. quenstedti and, although it is much more derived, it appears earlier in the stratigraphy. It is already in the Pleurodira lineage according to some shell elements, particularly its posterior bifid lobe, its pelvis already derived in the pleurodiran manner, linked to the shell by sutures, much reduced in width with respect to plastral width and with joined thyroid fossae (Broinl985; 1988a; Lapparent de Broin et Fuente 1996; Lapparent de Broin & Murelaga 1999). P. robusta is the oldest known taxon which can be considered at the remote origin of the pleurodiran fauna, including that of Africa. Palaeochersis talampayensis Rougier et al. 1995, from the Norian of Argentina, is more primitive than Proter ocher sis, judging from some characters such as those of the pelvis and plastron but it is difficult to compare them directly because the Proterochersis skull is unknown. However, its affinities with Pleurodira although possible (e.g., beginning of sutural link of the very primitive pelvis to the shell), are difficult to prove. Australochelys africanus Gaffney & Kitching 1994, from the Early Jurassic of Bormansdrift, Orange Free State, Karoo, South Africa (Table 1) is next in chronological sequence. It is the oldest chelonian taxon known from Africa, but is known only from a fragment from the bridge area of the carapace, not described, and a relatively poorly preserved skull in which the sutures are largely obliterated (Gaffney & Kitching 1995). Its relationships with other taxa are still uncertain, even for its authors (see Gaffney 1996). Rougier et al. 1995, consider it to be related to Palaeochersis and do not accept any possible relationship with Proterochersis. However the shared characters of Australochelys and Palaeochersis are weak and these skulls seem too primitive to offer good synapomorphies. As the skull of Proterochersis is not preserved, no comparison is possible with Australochelys. The oldest known African taxon therefore remains mysterious as to its origins and it is impossible to determine wether or not it is related to the infraorder Pleurodira, which developed in Africa before the arrival of Cryptodira. Kayentachelys aprix Gaffney et al. 1987, from the Early Jurassic of Arizona, USA, is approximately as old as Australochelys but it is a confirmed cryptodire. The skull characters are more derived and it has no obvious relationships with Australochelys. The next oldest chelonians from Africa are from the Middle Jurassic of El Mers, Morocco (Termier et al. 1940). No determination is possible on the poorly preserved material which consists of fragments of some plates. Approximately as old are the cryptodiran chelonians from the Middle Jurassic of China, such as Chengyuchelys baenoides Young & Chow 1953, C. zingongensis Yeh 1982 and Xinjiangchelys junggarensis Yeh 1986a (Yeh 1986b); they cannot be compared with the very poorly preserved African Middle Jurassic material. Besides, the large variety of Pleurodira found in Africa from Early Cretaceous times, Figure 2. Localities of oldest chelonians in Pangea: Triassic: 1, aff. Proganochelys sp., Greenland, Norian; 2, Proterochersis robusta, Proganochelys quenstedti, Germany, upper Keuper; 3, aff. Proganochelys ruchae, Thailand, Norian; 4, Palaeochersis talampayensis, Argentina, Norian. Early Jurassic, Hettangian: 5, Australochelys africanus, South Africa (see Table 1: (5)). Early Jurassic 6, Kayentachelys aprix. Middle Jurassic: 7, Chelonii indet., Morocco (see on Tab. 1:1); 8, Chengyuchelys baenoides, C. zingongensis and Xinjiangchelys junggarensis, China. From Smith & Briden 1977, Rhaetian period. and typical of Africa, indicates that this group was present on the continent much earlier than the Cryptodira. The Pleurodira From the Cretaceous onwards (Figure 3, Table 1), some chelonians are comparable with extant forms. The fossil material recovered from the Early Cretaceous of Cameroon and from the Algoa Basin, South Africa, have not yet been described. Material from the Early Cretaceous of the Anoual Basin, Morocco, although insufficient, shows signs of the presence of northern Gondwanan pleurodires (cf. Taquetochelys Broin 1980). The fossils from the upper part of the ‘Continental Intercalate’ of the Sahara (Kilian 1931 in Furon 1955), in the Early Cretaceous of the northern part of Africa, are well preserved. They are all Pleurodira and belong to the same group as remains found from the Early Cretaceous o f Brazil, South America (Pelomedusoides, see Broin 1988a, b) and its plesion the Araripemydidae. This continent was still linked to Africa during very early Cretaceous times. Pelomedusoides + Araripemydidae are vicariant to the Chelidae, 46 TABLE 1. Mesozoic African Chelonian Localities: (5), Early Jurassic on Figure 2; 1 to 32, Cretaceous, on Figure 3. Littoral and marine taxa underlined. All the taxa are northern Gondwanan elements except the Dermochelyoidea-Chelonioidea, which are Laurasian in origin. In bold face, older representatives ofthe groups. Age Formation Locality Taxa Early JURASSIC, Hettangian Upper Stormberg, Karroo, Elliott F. South Africa. Orange Free State Bormansdrift (5) Australochelys africanus Middle JURASSIC. Bathonian Morocco: 1 El Mers Chelonii indet. c Valanginian Kirkwood Formation South Africa: 2 Algoa Basin ?Chelonii indet K E T Barremian Morocco: 3 Anoual Basin at Oussikis and Ksar Metlili Plesion Pelomedusoides + Chelonii indet A c E Barremian-Aptian Cameroons: 4 Mayo-Rey River, Mayo Djarendi (E Koum Basin) Chelonii indet O U s Late Aptian "Continental Intercalate", lower upper part Niger: 5 Gadoufaoua, Ebrechko, Algeria 6 Aoulef, 7 Timimoun? Araripemydidae, Pelomedusoides including Pelomedusidae Albian- Cenomanian, prior to Early Cenomanian of Baharija "Continental Intercalate”, late upper part Algeria : 7 Timimoun, 8 Gara Samani. 9 Garet Touidjine, 10 Djoua at 120 km E Fort Flatters, In Akhamil, 17 km S Alrar; Tunisia: 11 Touil Dehibat, Rernada, Bir Kamboute, Dehibat, Gara Er Rehi. Guermessa, Er Ronda, Chenini trail; Morocco: 12 Kem-Kem, Hamada of Guir: Niger: 13 In Abagarit; Mali: 14 Tikarkas Araripemydidae, Pelomedusoides: Pelomedusidae, Bothremydidae, Podocnemididae Early Cretaceous Ethiopia: 15 Abay River Basin Araripemydidae, Pelomedusoides as in In Abangarit and Kem Kem (see above) Early Cretaceous Lupata Group Malawi: 16 Mwakasyunguti area. “Nyassaland". NW Lake Malawi Pelomedusoides Albian- Cenomanian South Africa: 19 Umtata mouth, Coast close to Umtafuna & Umzambawi Rivers Chelonioidea or Dermochelyoidea indet. Cenomanian Egypt: 17 Baharija Bothremydidae Cenomanian Madagascar: 18 Betioky Bothremydidae Cenomanian Wadi Milk and Shendi F Sudan: 20 Wadi Abou Hashim and NW Shendi loc.. loc. F1/89 and F 2/89 Pelomedusoides: Podocnemididae, Pelomedusidae; - Pelomedusoides Senoman Niger 21 Ibeceten 1 Pelomedusoides incl. Podocnemididae, Erymnochelys group Senonian Madagascar: 22 Berivotro Bothremydidae ?Late Cretaceous Gokwe F., middle of Calcareous Member Zimbabwe: 23 Gokwe area Pelomedusoides (aff. Platycheloides?) Campanian- Maastrichtian Kababish Formation Sudan: 24 Abyad Basin Chelonii indet. (?Bothremydidae) Maastrichtian Niger: 25 Ibeceten 2 Bothremydidae Senonian probable Maastrichtian Angola: 26 Ambrizette Bothremydidae Nigeria: 27 Sokoto: Wurno, Gada, Kworre, Gilbedi Bothremydidae + indet. Dakhla Formation Egypt: 28 Ammonite Hills Bothremydidae, Chelonioidea Phosphates Morocco: 29 Benguerir, 30 Oued Zem, 31 Oued Erguita Bothremydidae, Chetoniaidea Mali: 32 Tagnout Chaggeret Bothremydidae 47 Figure 3. Cretaceous African localities with chelonians, localities 1 to 32, see Table 1 and early Cretaceous close Brasilian localities: a, Bahia and Reconcavo-Tucano Basins, Barremian-Aptian, primitive Pelomedusoides; b, Chapada do Araripe, Ceard, Early Albian, Araripemydidae, various Pelomedusoides including primitive Pelomedusidae and pre-Podocnemidoidea. All the defined taxa are northern Gondwanan elements except the Chelonioidea, which are Laurasian in origin. From Smith & Briden 1977, Hauterivian period. Pleurodira which developed in southern Gondwana - South America (restricted to Patagonia in Cretaceous times, and probably Antarctica) and Australia. The two extant groups are principally differentiated by the formula of their cervical vertebral joints, which they acquired independently from the formula of primitive amphicoelous vertebrae. From the Cretaceous to the present, the northern Gondwana Pleurodira evolved various continental forms typical of Africa (Figure 3, Table 1, to Figure 6, Table 4) and others typical of South America. These are, firstly, the plesions of the extant Pelomedusidae, a family which developed only in Africa; then the Podocnemididae and Bothremydidae diverged from the Pelomedusidae and they are known in South America and Africa. The Bothremydidae first developed in Africa; they are not known in the very early Cretaceous of South America. Within the Podocnemididae, the Erymnochelyinae evolved in Africa and later a branch emigrated to southern Western Europe. During the Early Cretaceous, the Podocnemididae also developed in South America with their own branch, the Podocnemidinae, initially in the northern part of South America. Later, they migrated south to meet up with the Chelidae, which in turn migrated northwards (Broin 1988a, 1991, Broin & Fuente 1993; Lapparent de Broin & Fuente 1998; Lapparent de Broin et al. 1997). As early as the Late Aptian of Gadoufaoua, Niger, and the Early Albian of Ceara State, Brazil (Broin 1980; Gaffney & Meylan 1991; Lapparent de Broin 1994; Meylan & Gaffney 1991), the species on the two continents and even all the genera, except Araripemys, are different, showing the early break in continental links during Cretaceous times. In India, which at this stage was still linked to Africa, possibly the Bothremydidae and the Schweboemys group, a branch of Podocnemididae, developed during the Cretaceous (Jain 1986; Singh et al. 1998) and in the Palaeocene. This group is known in Africa at least from the Late Eocene of the Fayum, Egypt, and is also found in late-Early Miocene sediments of Egypt and Arabia. From the Cretaceous to Early Palaeogene, other Pleurodira developed in Europe, the Dortokidae, from the same ancient ancestral Triassic Pangean origin. However these arose independently, from a Jurassic branch different from that of the African Pleurodira (Lapparent de Broin & Murelag 1996,1999; Gheerbrant et al. 2000). Undefined marine Cryptodira (Chelonioidea?) During the Mesozoic period, no Laurasian continental migrations are known into Africa. There is only one record of a possibly marine turtle of uncertain Laurasian origin, on the southeastern coast of South Africa (Albian-Cenomanian). It is possibly an Australian Figure 4. Palaeogene African localities with chelonians, 1 to 24, see Table 2. From Smith & Briden 1977, Early Miocene period. 48 TABLE 2. Palaeogene African Chelonian Localities: 1 to 24, Figure 4. In bold face, first mention of continental cryptodires, Testudininei, terrestrial tortoises, Laurasian in origin. Littoral and marine elements underlined, Dermochelyoidea (Dermochelyidae or other families) and Chelonioidea (Cheloniidae or other families), Laurasian in origin. All the others, northern Gondwanan in origin. Age Formation Locality Taxa P A P Landana Cliffs Angola: 1 Cabinda Bothremydidae. Cheloniidae L A 1 a Mali: 2 Cheit Keni, 3 In Farghas Bothremydidae E 0 e 0 Jbel Guersif F. Morocco, E Ouarzazate Basin: 5 llimzi, 6 Hadrar Mgorn Pelomedusoides, Pelomedusidae incl. G E c e Phosphates Morocco: 7 Benguerir, 8 Oued Zem Bothremydidae, Chelonioidea N E n e Saudi Arabia: 9 Jabal Umm Himar Bothremydidae E 0 Phosphates, Ypresian Morocco: 10 Oued Zem, 11 Benguerir Bothremydidae, Chelonioidea c e n Phosphates, Ypresian Tunisia: 12 Gafsa-Metlaoui Bothremydidae, Chelonioidea, Dermochelyidae e Ypresian, Ait Ouarhitane F Morocco. E Ouarzazate Basin: 6 N’Tagourt 2 Chelonii indet. Ypresian, Lutetian Mali: 13 Samit, 4 Tamaguilelt Bothremydidae Early Eocene Senegal: 15 Popenguine Bothremydidae Middle Eocene Nigeria: 16 Ameki, Ombialla district Dermochelyidae Somalia: 20 Las Daban (Berbera) Bothremydidae Eocene, Middle- Late Algeria: 17 El Kohol Fresh-water Chelonii indet. Qasr es Sagha F . Late Eocene Egypt: 14 Fayum at Birket el Kurun, Abusir, Dineh, NW Qasr es Sagha Podocnemididae, Dermochelyidae Cheloniidae Late Eocene + Early Oligocene? Libya: 18 Djebel Coquin, 19 Dor et Talha Pelomedusoides: ?Podocnemididae 0 1 i Qatram F., Early Oligocene Egypt: 21 Fayum at NW Birket el Kurun Podocnemididae,Testudininei g 0 c Libya: 22 Zella Oasis Fresh-water Chelonii indet. e n e Ashawq F., Early Oligocene Oman (Dhofar): 23 Thaytiniti, 24 Taqah Podocnemididae.Testudininei marine cryptodire (see Australian forms in Gaffney 1991; Lapparent de Broin & Molnar, submitted). It co­ incides with the time of the wide spread of marine cryptodires in the world and this record, if its marine nature is confirmed, indicates that the marine environment allows faster spreading than continental environments. Second phase: spreading of Bothremydidae. This phase is partly superposed on the first one. From the end of the Early Cretaceous, the diversification of the Pelomedusoides produced two sister groups, the Podocnemididae and Bothremydidae. Although the family logically appeared earlier, the first confirmed members of the Bothremydidae is a form from the Early Cretaceous of the Tafilalt, Morocco (Lapparent de Broin & Werner 1998), equivalent to the late upper part of the ‘Continental Intercalate’ of the Sahara. The Bothremydidae rapidly developed new larger forms, including giants, first in the Early Cenomanian of Baharija, Egypt, then in the Cenomanian of Madagascar and Israel. The family diversified into a variety of genera belonging to several groups during Late Cretaceous times (Senonian-Maastrichtian). As early as the Cenomanian, the fossils come not only from sediments deposited in freshwater environments, but also from littoral-marine environments. Taking advantage of the opening ofthe Atlantic Ocean and progressing along the coast-lines, the family rapidly reached North and South America, along the northern as well as the southern route (Broin 1988b; Lapparent de Broin & Werner 1998). The family is found in the Africa-Mediterranean Basin, Western Europe and the two Americas, from the Cretaceous to the Early-Middle Miocene, mostly with 49 littoral forms but sometimes with forms which returned to the fresh-water environment in Europe. The Bothremydidae are well represented in the sediments of the Trans-Saharan Seaway and its restricted gulfs during Cretaceous-Ypresian times (Figure 3, Table 1, to Figure 5, Table 3a). The few preserved limbs do not show any strong adaptation to swimming in deep seas. On the other hand, several types of crushing palates had evolved to eat molluscs or crustaceans. During this phase, cryptodiran marine turtles of non- African origin (Chelonioidea and Dermochelyoidea), are found in the same African littoral sites as those which yield Bothremydidae. Third phase: invasion of continental cryptodiran Eurasiatic chelonians, in several waves. This phase began at the end of the Palaeogene (Figure 3, Table 2) and continued up to the Pliocene (Figure 5, Table 3). During this phase, in situ development of the Pleurodira took place, accompanied by the occasional beaching of marine cryptodiran turtles. First African Testudininei After the breakup of Gondwana, the first immigration of continental Eurasian chelonians was that of cryptodiran tortoises from the eastern part of the Mediterranean Basin as evidenced by fossils of nearly the same age in the earliest Oligocene of Oman and in the Early Oligocene of the Fayum, Egypt (Gigantochersina). Tortoises (Testudininei) existed from the Early Eocene of Laurasia: Early Eocene of USA (Hadrianus majusculus Hay 1904) and Europe (Broin 1977) and Middle Eocene of Asia (Gilmore 1931; Yeh 1963 and others). The lineages were already differentiated in North America, Europe and Asia by the Figure 5. Neogene, Mio-Pliocene African localities with chelonians, 1 to 52, see Table 3a,b,c. From Smith & Briden 1977, Present period. Middle Eocene at the latest (Broin 1977; Hay 1908; Williams 1950,1952). From the North American branch leading to the Gopherus group diverged the Central and South American group (Chelonoidis group). The African forms clearly issued from one of the differentiated genera of Eurasia between the Eocene and Miocene (Eurasiatic Hadrianus, Ergilemys, Cheirogaster, others indeterminate in Asia including ancestors of the living Indian general). Testudininei begin with relatively large forms (at least 40 cm in carapace length) and those arriving in Africa belong to a relatively primitive generalized type ‘Hadrianus’ which still retained a cervical scute. Upon their arrival, these Testudininei developed forms typical of Africa but, after the Early Oligocene, there are no records of continental chelonians in Africa until the Early Miocene. Therefore, African Testudininei had nearly all of the Oligocene in which to diversify before the fossil record resumes in the Early Miocene, by which time they had differentiated relative to the primitive Eurasian forms. It is therefore difficult to determine the precise area of their origin. Besides, there is evidence of a second wave of immigrant chelonians in the Early Miocene (see below) which may also have brought tortoises. During the Early Miocene, fossil remains indicate that the diversification of the extant Ethiopian endemic Kinixys (a tortoise with a hinged dorsal carapace) from Uganda, and Kenya, (ca. 19-20 myr), and another form related to the Ethiopian endemics, Impregnochelys from Kenya (ca. 18 myr) had already occurred. This indicates the minimum age of separation, among African tortoises, of the derived group of Ethiopian endemics from Centrochelys Gray 1872, and Stigmochelys Gray 1873. In this paper these two taxa are separated from the polyphyletic ‘Geochelone' s.l. (see below). Centrochelys may also be represented in the northern part of Africa and Arabia during the Early Miocene, at the same time as Kinixys in Kenya and Uganda. It is surely present in the Middle-Miocene of Arabia and later (Lapparent de Broin & Van Dijk 1999; Wood 1987). Stigmochelys may be represented in forms from the Early Miocene of Karungu, Kenya, and it is well represented in the Pliocene of East Africa (see the catalogue). The Ethiopian endemics have preserved the cervical scute present in the prim itive forms such as G igantochersina , while Centrochelys and Stigmochelys had lost it and they may therefore have had another origin: instead of being related to Gigantochersina of the Fayum, Centrochelys may be related to another form which arrived in a separate wave. Indeed, Centrochelys may be the sister group of the European Cheirogaster, known from the Late Eocene-Pliocene, from which it differs by the narrower xiphiplastra below the anals, and which is already differentiated before the arrival of Gigantochersina in Africa (Lapparent de Broin & Van Dijk 1999; Lapparent de Broin 2000). Among primitive and generalized characters of Testudininei, which give to forms attributed to ‘Geochelone’ an erroneous ■ appearance of close relationship, Cheirogaster, Centrochelys and Stigmochelys share the loss of the cervical (a highly homoplastic character among Testudininei throughout the world) and the associated nuchal notch, as do the two Indian species Geochelone elegans (Schoepff 1795), type species of the genus Geochelone Firtzinger 1835, and G. platynota (Blyth 1863). Relative to Centrochelys and Cheirogaster, the shell of Stigmochelys is different: always higher and often more vaulted, narrower and with a wider dorsal epiplastral lip, rounded instead of flat-concave. Bour (1985) has already demonstrated, on skull characters, the paraphyly of Geochelone including 50 Cylindraspis, which is the sister group of Stigmochelys. Actually, the two latter forms might just as possibly be related to Centrochelys or to the Indian forms as proposed by Bour (1985), or to endemic African forms, an hypothesis which has not been tested because of an arbitrary separation of large forms of ‘Geochelone’ from small forms, true Geochelone s.s. species excepted. These two extant species are relatively small, respectively ca. 25 cm and 26 cm carapace length, and they do not fit a concept of a ‘Geochelone’ being constituted of large forms (carapace length more than 35 cm up to 200 cm).The attribution to the genus Geochelone was done by TA BLE3,a,b,c. Neogene (Mio-PIiocene) African Chelonian Localities: 1 to 52, Figure 5. In bold face and underlined, oldest record of modern pelomedusid genera, 23, Pelusios and 19, Pelomedusa (not the oldest possible record, Pelusios being derived from a Pelomedusa stage), northern Gondwanan in origin. To the Gondwanan fauna, addition of Eurasian elements, oldest records in bold. Marine Laurasian elements (Dermochelyidae and Chelonioidea including Cheloniidae) underlined. Age, Formation Locality (oldest representatives, age in MY) Taxa M 1 0 C E N E Early Late Egypt: - 1 Moghara, 2 Wadi Faregh (ca18) - 3 Suez Canal - 4 Wadi Natrun (ca 6.3) - Carettochelyidae, Podocnemididae, Cyclanorbinae - Cheloniidae - Pelusios, Trionyx, triunguis lineage, Mauremys, Cheloniidae Early Late Libya: - 5 Djebel Zelten (ca16,5) - 6 Sahabi (ca 6,5) - Podocnemididae, Testudininei: cf. Centrochelys - Trionyx, triunguis lineage,Testudininei Early: Dam F. Oman: 7 Ghaba (ca18) Bothremydidae, Podocnemididae, Carettochelyidae, Cyclanorbinae,Testudininei Early to Middle: - Dam F - ?Dam F. - Hofuf F. Saudi Arabia: - 8 As-Sarrar - 9 Chalon -1 0 Al-Jadidah - ?Bothremydidae, Podocnemididae, Carettochelyidae, Cyclanorbinae, Testudininei - Cyclanorbinae - Testudininei Middle, Bayunah F. 11 Abu Dhabi, Western Region (ca 8) Trionychinae, Mauremys,Testudininei: Centrochelys Lower part of Late Miocene Tunisia: - 12 Bled Douarah (ca 11) -13 Djebel Semene (ca 10+) -1 4 Djebel Krechem Malta: 14bis - ?Trionyx - Testudo s.l. {?Testudo) - Testudininei Bothremydidae, Cyclanorbinae Algeria: 15 Bou Hanifia (ca 10,5), Saint-Eugene Chelonii indet. (?Testudo, ?Mauremys) Cheloniidae indet Early Kenya: 16: - Koru, (ca19-20) - Songhor (ca 19-20) - Mteitei area 17: - Rusinga Island, - Mfwangano Island. - Uyoma Peninsula. - Karungu, (ca 18) - Gwasi Peninsula, - Ombo - ?Cyclanorbinae -Testudininei: Kinixys - Chelonii indet. - Pelusios,Testudininei - Chelonii indet. - Chelonii indet. - Erymnochelyinae, Cyclanorbinae, Testudininei - Chelonii indet. - Cyclanorbinae, Chelonii indet. Early Middlle Early Namibia: 18 - Fiskus, Grillental, Elisa bethfeld 19 - Langental (ca 19), - Glastal 20: - Rooilepel (wardi, laini), Karingarab, North of Gypsum Plate Pan, 21 - Arrisdrift 22 - Auchas - Testudininei - Pelomedusa. Testudininei -Testudininei - Testudininei - Erymnochelyinae, Testudininei - Erymnochelyinae, Testudininei Early Uganda: 23: - Napak, (ca 19-20) - Moroto Pelusios. Testudininei: Kinixys Chelonii indet. 51 Age, Division, Formation Locality Taxa M Ngorora F., Mpesida beds, Lukeino F., Kenya: 24: Baringo Basin - Pelusios, Cyclanorbinae, Testudininei, i Miocene; Kaperyon F., Chemeron F., Aterir Chelonii indet. 0 beds, Pliocene; Chemoigut beds, Pleistocene P 25: Kerio River Basin: 1 (1) Late Miocene (1) Lothagam Erymnochelyinae, Cyclanorbinae, Testudininei i (2) Pliocene (2) Kanapoi, Ekora 0 Mio-Pliocene: Uganda, SW and E Lake Albert P (1) Late Miocene, Oluka F. (1) - 27 Kisegi-Nyabusosi area - Cyclanorbinae 1 Early to Late Pliocene: (2) - Nkondo F. (2) - 26 Nkondo-Kaiso - Pelusios. Cyclanorbinae e and (3) - Warwire F. (3) - 26 Nkondo-Kaiso - Pelusios i (4) - Nyakabingo F. (4) - 27 Kisegi-Nyabusosi area - Pelusios - Cyclanorbinae s (5) - Kaiso beds (5) - 26 Kaiso Village - Pelomedusoides indet., t Cyclanorbinae, Testudininei 0 c Congo-Zaire, West Lake Albert: e - Kaiso beds, Pliocene - 28 Lower Semliki River, -Testudininei n - Late Miocene-Pliocene, Ongoliba and - 28 Sinda-Mahori Rivers Region, - Pelomedusoides, Erymnochelyinae, e Sinda beds, Lower Semliki River Carettochelyidae, Cyclanorbinae - Lusso beds, Pliocene - 29 Upper Semliki-Senga Rivers - Pelomedusoides, Pelusios, Cyclanorbinae. Testudininei Tertiary, probable Late Neogene South Africa: 30 Carlisle Bridge Testudininei: first Homopus Age, D ivision, Form ation Loca lity Taxa P I i Pliocene Tunisia: - 34 Hamada Damous - 35 Ichkeul (ca 3,5) - IM aurem ys, ITestudo - Trionyx, ?Mauremys, Testudo s.s., Testudininei 0 c e Pliocene Chad, Koro Toro: 36 Ouadi Derdemi, 37 Bahr el Ghazal Pelusios, Trionyx, Testudininei, Chelonii indet. n e / Pliocene Morocco: 31 Ahl Al Oughlam (ca 2-2,5) Testudo P I i 0 Pliocene Algeria: - 32 Ain B oucherit (ca 2) - 33 Puits Karoubi - M aurem ys leprosa, Testudo, ? Trionyx - Chelonii indet. P I e i s t Pliocene part: - Yellow Sands, Mursi F. - Usno F., Shungura F., - Pliocene, Hadar F. included Ethiopia: 38 Omo River Basin: - ( 1 ) , - (2 ) 39 - Awash Valley, Afar - Pelusios, Cyclanorbinae - Pelusios, cf. Trionyx, Cyclanorbinae, Testudininei - Pelusios, Cyclanorbinae, Testudininei 0 c e Plio-Pleistocene Djibouti: 40 - Annabokoma Chekheyti, - Gobaad plaine - Testudininei - Pelusios n e - Koobi Algi F., Pliocene - Koobi Fora F., Pliocene C - Koobi Fora F., Pleistocene - Homa and Kanam beds, Pliocene - Kanjera beds, Plio-Pleistocene Kenya: 41 East Turkana - (1), -(2 ), -(3) 42 Homa Peninsula, Kanam (1), Kanam (2) - Pelusios - Pelusios, Cyclanorbinae, Testudininei - Trionyx, Cyclanorbinae, Testudininei Chelonii indet. Chelonii indet. - Bed I, II, Pliocene, - Bed IV, Pleistocene Tanzania: 43 - Laeto li area (ca 3,6-3,8) 44 - Olduvai (1) - Olduvai (2) - Testudininei, incl. Stigmochelys - Pelusios, Pelomedusoides, Testudininei - Testudininei Pliocene, Chiwondo beds 45 Malawi Pelusios, Cyclanorbinae Pliocene 46: Varswater Formation, PPM South Africa. -46 Langebaanweg (ca 4-4,5) - 47 Makapansgat, 48 Sterkfontein Member c: - Testudininei ?incl. first Chersina ?Pelomedusa - Testudininei Plio-Pleistocene -49 Taungs -50 Kromdraai A, B, 51 Swartkrans, Member 2 - 52 Drimolen (ca 2 -1,8-1,6) - Pelomedusa - Testudininei Testudininei: Psammobates 52 TABLE 4,a ,b ,c. Quaternary (Pleistocene-Holocene) African Chelonian Localities: 1 to 70, Figure 6. Underlined, marine Laurasian elements. In bold face, oldest records of tortoises from Madagascar area. Table 4a, locality ‘HaaskraaF, South Africa, Late Holocene: unlocalized on Figure 6, see Sampson, 1998. Age Locality Taxa Pleistocene-Holocene Morocco: Occidental Morocco 1 - Kenitra 2 - Coast from Rabat to Temara (a), Rabat 8, 9,10, - Coast from Rabat to Temara (b), Rabat 6, - Carriere Thomas I, Ain Bahya, - Dar Es Soltane, - Bouknadel, - Doukkala II, - Mehdia, Toulkine-Bou Ben Adam 3 - Jebel Irhoud, Oualidia, - Ain Rohr, El Khenzira Oriental Morocco: 4 - Taforalt 5 - Rhafas Cave, El Heriga, Abri Rhirane, Keneg Kenadsa - Abri Bou Guennouna - Oued el Haij Terrace, Jorf el Anngra - Testudo - Testudo - Trionyx - Testudo - Mauremys, Testudo, - Testudinidae - Mauremys, Testudo, - Testudo - Testudo - Testudinidae - Mauremys, Testudo - Testudinidae - Testudo, Testudinidae - Testudinidae Pleistocene-Holocene Algeria: 6 -Tighenif - Archaelogical sites Oran province, North of Aures 7 - Mansourah - Archaelogical sites, Constantine province Tunisia: 7bis, Archaelogical sites Malta: 7ter - Mauremys, Testudo - Testudo, Mauremys? - Mauremys -Testudo, Mauremys? Testudo, Mauremys? Testudininei, Mauremys Pleistocene 8 Cape Verde islands: Pedra de Lume Testudininei Pleistocene-Holocene Chad: 9 - Djourab, El Djour, Goz Kerki Recent, Bochianga, Neo- Bochianga, -1 0 Low Lands of Chad, 9 Borkou and UTibesti, -11 Puits Tirenno (Tibesti) - Pelusios, Trionyx, Cyclanorbinae, Testudininei - Trionyx - Pelusios, Cyclanorbinae Pleistocene Sudan: 12 Wadi Haifa Chelonii indet. Pleistocene Saudi Arabia: 13 An Nafud Testudininei Pleistocene Kenya: 14 Rawi Testudininei Pleistocene Tanzania: 15 Lake Eyasi - Mumba Cave Pelusios - Testudininei Pleistocene-Holocene Mozambique: 16 Zambezi tributary Lesotho: 17 Holocene sites Cyclanorbinae Testudininei, Chelonii indet. Pleistocene-Holocene Historic South Africa: 18 (N to S), Leliehoek Shelter; Oakleigh; Edgehill-Welgeluk; 19 Hantam Mountains; 20 (N to S) Elands Bay Cave, Klipfonteinrand Cave.Hopefield, Die Kelders 1, Byneskranskop Cave 1 - (Haaskraal) 21 Brandberg Chelonii indet.; Testudininei; Pelomedusa, Stigmochelys, Chersina, Homopus; Homopus, Chersina; - (Pelomedusa, Stigmochelys, Psammobates, Homopus) Stigmochelys reference to the large African species Testudo sulcata (up to 90 cm) which was attributed by Fitzinger (1836) to Geochelone before the erection of the genus Centrochelys Gray 1872, for that species. The mistake of many zoologists was then to consider as monophyletic a special group of large forms, when it was nothing more than a regrouping of large forms with the practical aim of determination (Auffenberg 1974; Wermuth & Mertens 1961; Iverson 1992). Cladograms have therefore been proposed for the large forms : - for ‘Geochelone’ by Crumly (1982); - for the ‘phylum’ (sic) Chelonoidina Gray, approximately corresponding to the extant ‘ Geochelone' s.l. by Bour (1985). Another cladogram has been proposed for all the group of tortoises, called the ‘Testudinidae’, by Gaffney & Meylan (1988), also separating the large ‘Geochelone’ forms from the others. All these studies are principally based on skull characters without any accurate study of the shells and they do not include the fossil elements of the various radiations from their appearance in Eocene times, in each continent. Even though all the fossil African forms have not yet been determined, it is possible that there were two or three waves of immigration, besides which there are three lineages of Testudininei in Africa, which deserve separate names, namely: Centrochelys', the Ethiopian 53 Age Locality Taxa Holocene - Historic t Holocene - Historic | Present Madagascar: 57 Gloriosa, 58 Mahajanga area, 59 Ampasambazimba, 60 Antsirabe, western-southern area between 61 Ambato (Morondava) and 64 Andrahomana (Taolanaro) including 62 Etsere and 63 Ambolisatra, - 61 Ambato, 61 Ankevo, 60 Antsirabe, 63 Ambolisatra, 64 Andrahomana, Antinosi D ipsoche lys - A s troche lys Holocene - Historic Present, in part t Aldabra area: 65 Granitic Seychelles islands, 66 Seychelles atolls including Aldabra Dipsochelys Historic Comoros islands, Mayotte island: 67 Dembeni 1 (introduced from Madagascar) Erymnochelys, Pyxis, Astrochelys, Cbefortia Holocene - Historic Present Mascarene islands : 68 La Reunion, 69 Mauritius, 70 Rodrigues 69 Mauritius (introduced with other taxa) - C ylindrasp is - Dipsochelys Age Loca lity Taxa Holocene Algeria: 22 Ti-n Hanakaten Pelomedusa, Testudininei Holocene Mauritania: 23 Chami Testudininei Protohistoric Senegal: 24 Sintiou Bara, 25 Tulel-Fobo - Faboura Cyclanorbinae - Cheloniidae. Trionyx Holocene Mali: Taoudenni Basin: 26 - Araouan, Guir, Djouf 27 - Hassi el Abiod 28 - Erg Ine Sakane 29 - Kobadi - Trionyx, Cyclanorbinae - Pelusios, Trionyx, Cyclanorbinae - Pelusios, Cyclanorbinae - Trionyx Holocene N ige r: Tenere: 30 - Adrar Bous - 31 Tin Ouaffadene, 32 - Bilma, Kaouar 33 - Azaouak Valley : - In Aruinat - Ikawaten - Takane Barva - Pr Baumhauer site - Pelusios - Testudininei - Pelusios - Pelusios, Cyclanorbinae - Cyclanorbinae - Trionyx - Pelusios, Trionyx Protohistoric Chad: 34 - Sao de Mdaga 35 - Koyom - Pelusios - Pelusios, Cyclanorbinae Protohistoric Ghana: 36 Mole National park Cyclanorbinae Holocene Nigeria: 37 N Maiduguri, Chad basin Pelusios Protohistoric Cameroons: 38 - Sou, - Lake Chad - Pelusios, Cyclanorbinae - Pelusios Holocene Libya: 39 - Djebel Zelten, - ?S CyrenaTca, Libyan desert - Cyclanorbinae - Pelusios Holocene Egypt: 40 - Toukh, 41 Fayum at Birket el Kurun, 42 Adai'ma 43 - Berenike 44 - Abu Balias - Trionyx - Cbeionia, Testudininei - Testudininei Holocene-Protohistoric Sudan: Nile Valley at 45 - Saggai and Geili, - Umm Marihi 46 - El Kadada 47 - Debbat Bangdit 48 - Debt El Eheima. Atbara River Valley at: 49 - Khashm el Girba 50 - Jebel Shaqadud W est to Nile Sudan at: 51 - Burg et Tuyur 52 - Wadi Howar 80/73 - Pelusios, Trionyx, Cyclanorbinae - Pelusios, Testudininei - Pelomedusa, Trionyx, Cyclanorbinae - Pelusios, Trionyx, Cyclanorbinae - Pelusios, Cyclanorbinae - ?Pelusios, Trionyx, Cyclanorbinae, Testudininei Pelusios, Testudininei - Testudininei - Pelusios Holocene Delta Ethiopia: 53 Awash Valley, Afar Testudininei Holocene Congo-Brazza: 54 Ntadi Yomba Testudininei Holocene Congo-Zai're: 55 - Matupi Cave 56 - Malemba-Nkulu, Sanga - Testudininei - Pelusios ■ endemics; and Stigmochelys. The lineages separated at ca. 20 myr, each one having developed several species since then. Furthermore, it is no longer possible to attribute to ‘Geochelone s.l.’ large forms which have not been accurately studied and compared, and which are actually included in constituted lineages of different continents, themselves separated from Early Eocene times and in Africa at least from the Early Miocene. There is no diagnosis o f Geochelone which corresponds to all the taxa that are attributed to this artificial genus and its definition is confused in the literature (Auffenberg 1974; Loveridge & Williams 1957; and others). Meylan & Auffenberg (1986), acknowledge that the ‘genus’ is not monophyletic and employ it simply for convenience. Because there is no consensus on the definition of Geochelone, the name must therefore be restricted to the extant Indo-Asiatic type species, G. elegans, and possibly to G. platynota, which is regarded as close by some workers, and their possible fossil relatives which are not yet recognized in the fossil forms from Asia. Arrival o f first Trionychoidea: The oldest Trionychoidea, including Carettochelyidae and Trionychidae, are known from the Late Jurassic- W ealden o f China. The Carettochelyidae are represented in Arabia-northern part of Africa at the base of the Miocene. This family is known from the base of the Early Eocene in North America and in Europe (MP 7-8) (Broin 1977). However, the African-Arabian form has shell characters of the Eurasiatic subfamily Carettochelyinae, with an anterior carapace border more similar to that of Allaeochelys (the European form) and more primitive than that of the extant Figure 6 . Quaternary, Pleistocene-Holocene and Historic African localities with chelonians, 1 to 70, see Table 4a,b,c. Missing locality: ‘Haaskraal’, South Africa, late Holocene (unlocalized; see Sampson 1998). From Smith & Briden 1977, Present period. Carettochelys of New Guinea-northern Australia. The Carettochelyinae of the Indian subcontinent could also be an ancestor (Broin 1987), but they are too poorly preserved for meaningful comparison. Trionychidae Cyclanorbinae must have originated from an undifferentiated Asiatic trionychid. Cyclanorbinae are unknown before their arrival in Africa, first appearing in the fossil record at the same time as the Carettochelyidae in Arabia and the northern part of Africa. The two taxa might have arrived together, crossing eastern Tethys by following the coastline. Cyclanorbinae are also recorded farther south at Karungu, Kenya (ca. 18 myr), and an undefined trionychid was already present at Koru (ca. 20 myr), which might indicate that Cyclanorbinae were present a little earlier than Carettochelyidae. The Cyclanorbinae developed forms typical of Africa, Cycloderma and Cyclanorbis, while Lissemys developed only in India. As well as the primitive African Testudininei, the primitive African Cyclanorbinae are not yet well defined because of insufficient material. The origin of the Cyclanorbinae is completely unknown but probably should be sought in the Indian subcontinent: once India contacted Asia, as early as the Eocene in Pakistan, trionychid remains are found which might be related to the Cyclanorbinae (Broin 1987), but the sub-family itself is known in this subcontinent only from the Late Miocene (Siwaliks of the Potwar Plateau, Pakistan, and of Ramnagar, India, in the Chinji Formation, ca. 10,5 and 13-14 myr respectively), long after their record in Africa. While Cyclanorbinae still survive today, the Carettochelyidae disappeared during the Early Pliocene, the last known remains coming from the Lower Semliki in the Democratic Republic of Congo. Arrival o f the extant Palaearctic fauna A new wave of Eurasiatic immigrants occurred during the Middle-Miocene which brought in Trionyx, Mauremys and Testudo s.s. (Figure 5, Table 3a, 3c). The first references to these taxa are of dubious value (lower part of the Late Miocene, Algeria, Tunisia). However, a true Trionychinae, although not Trionyx s.s., and a Mauremys, primitive or related to the extant M. caspica from the eastern Mediterranean Basin (including the Arabian Peninsula on the Arabian Gulf border) are confirmed in the Baynunah Formation of the Emirate of Abu Dhabi (ca. 8 myr), probably coming to northern Africa from the east of the Mediterranean Basin. During the Pliocene, the presence of Trionyx s.s. (lineage of Trionyx triunguis), Mauremys and Testudo s.s. is confirmed in the northern part of Africa (Sahabi, Wadi Natrun and later the Maghreb, with extant lineages). The first Trionyx triunguis is attested in Kenya (Koobi Fora Formation, upper members) only during the beginning of the Pleistocene. On the other hand, Mauremys and Testudo did not reach the southern Nile valley and become permanently established in northern Africa. Trionyx disappeared from this area during the Late Pleistocene, after the conquest of all the northern mid-part of Africa and a part of the south­ western margin. The origin of the Palaearctic chelonian fauna of Africa is probably from the Mediterranean Basin because Trionyx s.s. belongs to a group represented in Europe from the Palaeocene, although the lineage of the extant T. triunguis truly diversified only during the Late Miocene in northern Africa. The same is true for Mauremys, represented in Europe from the Oligocene, with possible earlier plesions (Palaeochelys s.s.): however the lineage of the extant Mauremys is confirmed only from the Pliocene. The earliest confirmed occurrence of the extant Maghrebian and Franco-Spanish form, M. leprosa, is from the Pliocene of Ain Boucherit, Algeria (ca. 2 myr) although it had most probably already differentiated before that (fide some of the material from the Maghreb, most of which regrettably is lost). As far as Testudo s.s. is concerned, a genus characterized by the presence of a hypo-xiphiplastral hinge in both males and females, poorly preserved Maghrebian forms attributed to the genus Testudo s.l. are known from the lower part of the Late Miocene, but the definitive presence of Testudo s.s. in Africa is only from the Pliocene of Ahl Al Oughlam, Morocco (ca. 2 to 2,5 myr), and possibly from Ichkeul, Tunisia (ca. 3,5 myr). The first mention of a true Testudo s.s. in Europe is only from the Late Miocene of Pikermi and Saloniki, MN 12-13, Turolian (Gmira 1995), but ancestral forms of Testudo s.l. existed from the Oligocene. Its precise point of differentiation remains unknown. During all of the Late Miocene and Pliocene in the northern part of Africa, large and giant forms of tortoises attributed to ‘Geochelone’ s.l. or to ‘Testudo’, coexisted with Testudo s.s.: their phyletic relationships and their origin are unknown, although a relationship with Centrochelys is possible or even probable in some localities. Continuation o f in situ development o f Pelo­ medusoides during theTertiary The Erymnochelys group, characterized by its gulars linked behind the intergular, is first recorded from Early Cretaceous sediments of Niger (Ibeceten 1), and it continued its development in the northern part of Africa (Egypt, Arabia) during the Late Eocene and Early Miocene (no data available from the Palaeogene up to Fayum times), with its principal collaterals, Stereogenys and Schweboemys, known from the Late Eocene of Fayum. The oldest known occurrence of the latter is from the Cretaceous of the Indian subcontinent. Later, the group migrated to Pakistan (Early Miocene) and to Burma (Plio-Pleistocene) but there are no post-Early Miocene records from Africa-Arabia. The Erymnochelys group first reached East Africa and Namibia during the Early Miocene, and continued its development during the Late Miocene-Pliocene only in East Africa. This Erymnochelys group is no longer known from Africa from the Early-Mid Pliocene; Erymnochelys survives today as a refugee in Madagascar. Its disappearance is comparable to that of C arettochely idae. After their record from the Early-Mid Cretaceous of Sahara and Thanetian of Morocco, the Pelomedusidae disappear from the record throughout the Palaeogene. They reappear in the Early Miocene of Langental, Namibia (ca. 19 myr), with a new species of Pelomedusa discovered recently (1998). This genus is a fragile form which is rarely fossilized (Wood 1973b). There is no further record until the Pliocene of Langebaanweg, South Africa, and then it is known from only a few citations up to the Present. On the other hand, Pelusios, a box-turtle and a form robust and more derived than Pelomedusa, is currently found from the Miocene of Uganda (ca. 19-20 myr) and Kenya (ca. 18 myr) and it still survives today. Marine cryptodiran forms There are very few records of marine Cryptodira in Africa during the Late Palaeogene-Neogene; some localities in Egypt (Fayum, Suez, Wadi Natrun) record their passage between the Mediterranean Basin and the northern Atlantic up to the North American coast. Emigration from Africa Other than the littoral Bothremydidae, only the Erymnochelyinae seem to have emigrated from Africa during the Tertiary. Neochelys, a genus also possibly represented in the Late Eocene of the Fayum, is known in western Europe from the earliest part of the Early Eocene (MP 7; Broin 1988a). It may have arrived earlier, but it is not yet recorded in the rare European Palaeocene localities. Erymnochelys eremberti Broin 1977 (with the intergulars linked behind the gular, the curved premaxillary and mandibular hook, the prolongated temporal roof and the absence of lateral cheek emargination, characters typical of the genus) is known from the Mid-Eocene of France. The members of the Schweboemys group present in the Late Tertiary - Pleistocene of Pakistan and Burma may also be immigrants from the north of Africa-Arabia. Phase of diversification and endemism From the Pliocene to the Holocene (Figures 5 and 6, Tables. 3b-c and 4), localities with continental chelonians are more abundant, first in the Maghreb, the East African Rift Valley and in South-Africa, then in the northern part of Africa. The communicating basins of the Nile, Chad, Niger and Senegal constituted a favourable area for the spread of turtles. All the extant genera of turtles then diversified and no new taxon entered Africa during that time. However, the diversity of species increased locally. Several independent waves of chelonians, tortoises and turtles, entered Madagascar and the surrounding Indian Ocean islands from Africa at an uncertain date (data from 125 000 years at Gloriosa) (Figure 8) and they diversified widely. The oldest known tortoise, D ipsochelys, is probably related to "1 Astrochelys (itself possibly linked to the Ethiopian endemics); Cylindraspis is probably related to Stigmochelys (Bour 1984a,b, 1985, 1987); Pyxis (no known fossil record, but sub-fossil historical data) is linked to the African Ethiopian endemics. Dipsochelys and Cylindraspis were exterminated by humans, the last citations being from ca. 150 years ago, except for D. e. elephantina, which is still present in Aldabra and recently introduced to other islands. Gerlach & Canning (1998) consider that some individuals living in captivity on granitic Seychelles belong to the extinct D. hololissa and to D. arnoldi. However, such a sensationally novel suggestion has not yet gained acceptance among the scientific community, partly because the morphological data are unclear and not fully in accord with the original descriptions of the species, and partly because the published genetic data are inconclusive. Without deliberately rejecting these results, it seems premature to give them full confidence, and we prefer to wait for further, more decisive data (Bour pers. comm.). The freshwater Pelomedusidae also entered Madagascar and Seychelles, at an uncertain date (Figure 7) but sufficiently long ago to allow for their known diversification. Regressive phase (end of Holocene-Present) The figures showing extant turtle distribution (Iverson 1992; Figures 7 & 8) compared with figures of fossil data (Figures 5 & 6) during the Miocene-Holocene time, show the important faunal reduction due to climatic change at the end of the Holocene. Climatic alternations of cooling and/or drying periods with intercalations of 56 Figure 7. Present approximate limit of distribution in Africa, Arabian Peninsula and Madagascar area, of extant Pleurodira: Pelomedusidae, 1, Pelomedusa, 2, Pelusios', Podocnemididae, 3, Erymnochelys (Madagascar only). From Smith & Briden 1977, Present period. Figure 8 . Present approximate limit of distribution, in Europe, Africa, Arabian Peninsula and Madagascar area of extant continental Cryptodira: 1, Palaearctic fauna, freshwater Emys and Mauremys, tortoise Testudo s.l.; 2, African tortoises, Testudininei, endemics and ‘Geochelone ’ i.e. Stigmochelys and Centrochelys', freshwater turtles, outside ofNorth Africa: 3, Cycloderma (left, C. aubryi, right, C.frenatum), 4, Trionyx, 5, Cyclanorbis (both left and right, C. elegans and C. senegalensis)\ 6 , Indian Ocean and Madagascar islands, Testudininei. From Smith & Briden 1977, humid periods, eliminated Trionyx and the large tortoises from the Maghreb, leaving only the extant palaearctic elements, Testudo and Mauremys. The aridification of the Sahara pushed back the other elements of the African endemic chelonian fauna to the limits ofthe Sahel (Broin 1983, fig. 50; R osetta /. 1990, fig. 5) and in the reduced area of the Sahel, extant records are few and isolated. Even the desert tortoise, Centrochelys sulcata, no longer occupies all of its potential distribution area after so many Holocene climatic fluctuations. Emys orbicularis (unknown as a fossil in Africa) arrived in the Maghreb from the Mediterranean Basin and it is the last known immigrant of the Palaearctic fauna into Africa. It is a recent genus, known from the Late Miocene of Ukraine, Pliocene of Poland and Slovakia and the Pleistocene-Holocene of southern Europe. It has the most northerly distribution of the extant European fauna (Iverson 1992; Figure 8) and its adaptation to Africa is most probably linked to climatic changes. 57 PRELIMINARY CATALOGUE OF THE FOSSIL CHELONIANS OF AFRICA Stratigraphic and geographic distribution of the taxa The countries are presented in alphabetical order and for each country, the order of information supplied is: stratigraphic division: locality, area, geological formation (when known), age, taxa (new and first determination and references). When possible, an appropriate determination is proposed for the reviewed material, following recent taxonomies which divide the families s.l. or genera s.l. (for example ‘pelomedusid’, ‘trionychid’ or Geochelone, Testudo, Podocnemis) into monophyletically more precise elements. The original definition is given (quotation marks, brackets). The determinations may differ from those proposed in earlier works of mine. The present geographic distribution of the taxa is given in Iverson’s (1992) figures. Some general stratigraphic and geographic data on fossil localities are given in Cooke (1978) (all Africa), Pickford et al. (1993) and in references given by the authors of taxa. The position of the land masses from the Jurassic to Present are from Smith & Briden (1977) (see also Smith etal. 1994). The museum repository of the undescribed or reviewed material is given. The references to localities are not exhaustive, including only those with the first mention of chelonian taxa and their further taxonomic modification and at least one for the stratigraphic and geographic position of the localities. Lists of fossil and living chelonians are principally given by Kuhn (1964), Wermuth & Mertens (1961, 1977) and determinations of the extant African chelonians are principally from Loveridge (1941) and Loveridge & Williams (1957); general ecological information is presented in Pritchard (1979). It is impossible in this catalogue to cite all the references relating to extant African chelonians but additional references may be found in the following older works: Baur (1888a, b); Boulenger (1889); Cope (1868); Dumeril (1855-1856); Dumeril & Bibron (1835); Fitzinger (1826, 1835, 1836, 1843); Gray (1825, 1855-1870, 1872, 1873); Hewitt (1914,1927); Schweigger(1812); Siebenrock(1902); Smith (1838-1849); Wagler (1830); or in more recent such as: Bour (1981, 1983, 1986); Broadley (1981a,b, 1983, 1993, 1997a,b); Gerlach (1998, 1999); Laurent (1956, 1964, 1965) etc. The given MN (‘Mammal Neogene’) ages are from Mein (1990), the given MP (‘Mammal Palaeogene’) ages are from Escarguel etal. (1997). AFRICA ALGERIA Cretaceous: ‘Continental Intercalaire ’ o f Sahara, of Kilian (1931) (see Furon 1955; Lapparent 1960), upper part, Early Cretaceous MNHN (Broin det.): Late Aptian, A.F. de Lapparent coll., Aoulef, Tidikelt, Late Aptian, Araripemys sp.: Fuente & Lapparent de Broin (1997). Timimoun, Gourara (Foggara Amerhai'er), Late Aptian?, Araripemys sp., Albian-Cenomanian prior to Cenomanian age of Baharija and to marine Cenomanian transgression, Timimoun, Gourara (Foggara Amerhai'er), ?Araripemys sp.: Fuente & Lapparent de Broin (1997), first given as ‘Primitive trionychoids’: Broin (1977); Pelomedusoides indet.; A.F. de Lapparent coll. Gara Samani, between El Golea and Timimoun (Broin et al. 1971), Araripemys sp.: Fuente & Lapparent de Broin (1997), first given as ‘Primitive trionychoids’: Broin (1977); Fuente & Lapparent de Broin (1997); Pelomedusoides indet. including Podocnemidoidea (?Podocnemididae, Bothremydidae). Oued Boudjihane area, close to Atlas (Ksour), E of Ain Sefra, (Bassoulet & Iliou 1967), Iliou coll. and MNHN (coll. Bassoulet): a - Garet Touidjine, Araripemys sp.: Fuente & Lapparent de Broin (1997); b - Gouret Tin (high level, East), Pelomedusoides indet. Djoua (El Djoua), type locality of the ‘Continental Intercalate’ of Kilian (1931, see Furon 1955), 120 km E Fort Flatters, In Akhamil and 17 km S Alrar, E Algeria close to Libyan border; several taxa of Pelomedusoides indet., including Bothremydidae (Djoua) and at least one Podocnemididae (cervical vertebra at In Akhamil); coll. Nougarede and A.F. de Lapparent. Eocene: El Kohol (El Kohel), S Oran Province, near Brezina, between Early Eocene and Late Eocene, ?Pelomedusoides: ‘Paludine turtle’: Bergounioux (1954-1955) = fresh-water indet., unknown localization; Mahboubi etal. (1986). Late Miocene: Saint-Eugene, ‘Carriere des chaux et ciments’, Oran Province, Sahelian, Late Miocene (Tortonian, see Pomel references in Bleicher 1875), Cheloniidae indet., Dr L. Geslin coll., MNHN (Broin det.). Bou Hanifia (Oued El Hammam), Oran Province, Late Miocene, Vallesian, MN 9+, ca. 10,5 myr, Chelonii indet.: ‘Testudo': Arambourg (1952b, and 1954) ‘Emys’: Arambourg (1958) (possibly a Testudo s.l.? sp. and Mauremys sp., unverified presence, unknown localization). Pliocene: Puits Karoubi, ca. 2 km SW Eckmtihl, Oran, Chelonii indet.,‘Argiles du niveau b a Tortue’ (Clays, level b with chelonians): Arambourg (1950) (unknown localization) (Pomel 1878). Ain Boucherit (area of A'in Hanech and El Eulma, ex Saint- Amaud), Constantine Province, towards Setif, Pliocene, Ruscinian, MN 14-15, ca. 2 myr. ITrionyx sp. (‘Trionyx’: Arambourg 1953, unverified presence, unknown localization); Testudo s.l. indet. (possible s.s.) sp., Mauremys leprosa (Schweigger 1812), MNHN (Broin det.). Pleistocene: Tighenif (Temifine, Palikao), 20 km E Mascara, Oran Province, Early Middle or Middle Pleistocene, ca. 400 000 to 700 000 yr BP (Geraads et al. 1986: Hublin 1985), Testudo s. 1. (probable s.s.) sp., Mauremys leprosa, MNHN. Mansourah, Constantine Province, Pleistocene, Mauremys sp., probable M. leprosa (‘Emys, close to the extant Emys sigriz’, Thomas 1878, 1880, i.e. Mauremys leprosa)', MNHN (Broin det.). Holocene: Epipalaeolithic, Capsian (see references in Roubet 1966, localizations fig. 2,1979), Cubitus, near Tiaret, A'in Keda, near Tiaret, Abri Alain, Oran, Eckmtihl quarries, Testudo g. graeca : Roubet (1966), ‘Tortue de Mauretanie’, T. ibera\ Various localities, Mauremys leprosa? Vaufrey (1955). Neolithic (see references in Roubet 1966, localizations fig. 2, 1979) from East to West, Abri du Relilai', Col des Kifene, Damous el Amar (3770- 3450 years BC), Capeletti cave, Khanguet Si Mohamed Tahar (between 3950±150 and 2390±200 years BC), Djebel Fartas, Djebel Marhsel, Hyenes Cave, Bou Zabaouine Cave, Ours Cave, Hadrar Gueldaman, Columnata, Rhar Oum el Fernan, Cascades Cave, Oued Saida, Troglodytes Cave, Polygone Cave, Cuartel Cave, Ciel Ouvert Cave, Foret Cave, Chabet Sardi Cave, El Bachir Cave, Corales fireplaces -Escargots cemetery, Ain Guedara upper cave, Dahar Mendjel, Testudo g. graeca : Roubet (1966), ‘Tortue de Mauretanie’, T. ibera’. South: 58 Hassi Mouilah (Ouargia), Testudo g. graeca: Roubet (1966), ‘Tortue de Mauretanie’, ‘T.ibera Ti-n Hanakaten, NE Hoggar Djanet Province, around 7000 yr BP (Chai'd-Saoudi 1987): Pelomedusa cf. subrufa (Lacepede 1788); Centrochelys sulcata (Miller 1779): coll. G. Aumassip (Broin det.). ANGOLA Senonian (littoral): Ambrizette, 1 km S Ambrizette, N’Zeto, lat. around 7° 15', N Ambriz, Late Senonian (ca. Campanian), Bothremydidae indet., MNHN (Broin det.). Palaeocene (littoral and marine): Landana cliffs, Cabinda, Montian, Taphrosphys congolensis (Dollo 1913), (Bantuchelys Dollo 1924 ex parte)', ?Toxochelyidae indet. (Bantuchelys Dollo 1924, ex parte) (see: Antunes & Broin 1988; Dollo 1924; Lapparent de Broin & Werner 1998; Wood 1973a, 1975). CAPE VERDE ISLANDS Pleistocene: Pedra de Lume crater, Sal Island, ar 16° N, 24° W, Middle Quaternary, Centrochelys sulcata: Chevalier et al. (1935) (unknown localization). CAMEROONS Early Cretaceous: Koum Basin, 150 km SE Garoua, Mayo Djarendi, loc. KB6 , Mayo-Rey River, North Cameroons, Barremian-Aptian, Chelonii indet.: ‘Chelonia’: Brunet etal. (1990) (not seen). Protohistoric: Sou, Lake Chad, 75 km S Middle South shore, (Lebeuf; W. Van Neer pers. comm., Broin det.), 7th-19th Century; Pelusios adansonii (Schweigger 1812), Cyclanorbis senegalensis (Dumeril & Bibron 1835). Lake Chad, S shore, probably close to Sou, Middle Age (S. Becouch & H. Thomas, pers. comm, from Lebeuf?, Broin det.), Pelusios adansonii. CHAD Pliocene to Recent: Koro-Toro: Ouadi Derdemi (Ouadi Derdemy), close to Goz Kerki fossil bank, ca. 46 km E Koro-Toro, NE Fort-Lamy, ca. 3-3,2 myr, Pliocene (Coppens 1962a, 1965, 1967), Pelusios sinuatus (A. Smith 1838), cf. Trionyx sp., Centrochelys cf. sulcata (Miller 1779): Broin (1969; Chad coll., deposited MNHN. Bahr el Ghazal, E Koro-Toro, KT 12 site, ‘Abel’ site, 16°00’21" N, 18°52’34" E, Chelonii indet.: ''Trionyx sp., Geochelone sp.’, (provisional determinations, unverified); other sites, Pliocene, ca. 3-3,5 myr, Chelonii indet.: Brunet etal. (1996). Other Chad localities, Plio-Pleistocene-Holocene (Broin det.): El Djour, Centrochelys sp., Pelusios cf. sinuatus, likely prior to Holocene, Pliocene as Ouadi Derdemi; Yayo, Trionyx sp., Pelusios cf. sinuatus', Goz Kerki, Koula, Trionyx triunguis (Forsk&l 1775), Centrochelys sp.; Bochianga (diatomite, fauna 2), Pleistocene-Holocene, Cyclanorbis senegalensis (Dumeril & Bibron 1835) (small form); Neo-Bochianga (Stone Age with harpoons), Trionyx triunguis; (Coppens 1962a,b, 1965a,b, 1967a,b, 1968; see references not given here in Broin 1969); Puits Tirenno, 21°36' N, 17°25' E, Tibesti, Pleistocene- Holocene, Pelusios cf. castaneus (Schweigger 1812), Cyclanorbis senegalensis, (small form), cf. Centrochelys sp.; Y. Coppens coll.; Low Lands of Chad, Eguei', Kanem, Djourab, Borkou and Tibesti, Pliocene?, Pleistocene- Holocene, old mentions (unknown localization of fossil chelonians), Trionyx sp.: Priem (1914) (and Arambourg 1934; Joleaud 1934, 1936; Roman 1935). Protohistoric: Koyom, 9°48' N, 15°52' E, Middle Logone, S. Chad, ca. 19th Century, Pelusios adansonii (Schweigger 1812); Cyclanorbis senegalensis', Cycloderma aubryi; Trionychidae indet. (Trionyx sp.?); Kinixys belliana Gray 1831: Broin and Van Neer in Rivallain& Van Neer (1983,1984). Sao deMdaga, 12° 12’45" N, 15°03’30" E, ca. 12 km N to N’Djamena on Massakori road, Lake Chad basin, from 425 BC to 1780 years AD: Pelusios sp.: Thomas (1980) (Broin det.). CONGO, PEOPLES REPUBLIC OF (EX CONGO-BRAZZAVILLE): Holocene, around 7090-6890 yr BP: Ntadi Yomba, W Brazzaville, 13°46' E, 4° 15' S, Tshitolian Abri, Testudininei indet. including ?Kinixys sp.: Van Neer & Lafranchi (1985) (Broin det.). CONGO, DEMOCRATIC REPUBLIC OF (EX ZAIRE, CONGO-KINSHASA) Late Miocene - Pliocene o f West Lake Albert (Makinouchi et al. 1992): Sinda-Mohari Rivers Region, Lower Semliki River; Late Miocene, Sinda beds; Ongoliba Horizon, lower member, Late Miocene, site 5, Pelomedusidae indet., site 11, aff. Erymnochelys sp.; Mio-Pliocene or Pliocene?: sites 3,7, middle member, 15, upper member, aff. Erymnochelys sp.; site 3, middle member, Carettochelyidae indet.; sites 1, 3, middle member, 15, upper member, Trionychidae, probable Cyclanorbinae, apparently including Cyclanorbis senegalensis (Dumeril & Bibron 1835) group (‘Trionychidae’: Hirayama 1992, pi. 5). The remaining presence of Carettochelyidae and aff. Erymnochelys should be more favourable to an oldest possible age during the Pliocene. Lower Semliki River, Kaiso beds (old coll.), Pliocene, Testudininei indet. (giant terrestrial form: Leriche 1939). Upper Semliki-Senga Rivers, Lusso beds, Senga 5 and other localities, Pliocene, ca. 2,4-2,5 myr (Boaz 1990; Verniers & Heinzelin 1990), Pelomedusoides indet., either Erymnochelyinae or Pelomedusidae (‘Pelomedusidae indet.’), cf. Pelusios sinuatus, cf. Cycloderma sp., Testudininei indet., Ethiopian endemics group? (‘Testudinidae indet.’): Meylan (1990). Holocene-Protohistoric: Matupi Cave, northeastern Zaire, just east of Lake Albert shore: 1) Late Stone Age, between 22 000 and 2000 yr BP, Kinixys erosa, Testudininei indet. ( ‘Testudo’ sp., ‘Testudinidae’), 2) Iron Age, before 2000 yr BP, Kinixys erosa, Testudininei indet. (‘Testudinidae’) (Van Neer 1984, 1989). Malemba-Nkulu, Sanga, Upemba depression, ca. 26° to 21° E, ± 8 ° S, Iron Age, Pelusios sp.: Van Neer (1978) (including Pelusios nanus Laurent 1965, at Malemba-Nkulu, Broin det.). DJIBOUTI, REPUBLIC OF Plio-Pleistocene (Broin det.): Annabokoma-Chekheyti site, loc. NW 100, Gobaad plaine, 1,2-1,8 myr, ? cf. Centrochelys sp.: ‘Geochelone’ sp.: Thomas etal. 1984). Undefined site of Gobaad plaine: Pelusios cf. sinuatus (A. Smith 1838) (Gasse & Rognon 1973: ‘chelonians’). EGYPT Cretaceous: Baharija, SW Cairo, Western Desert, ca. 28° N, 29° E, Early Cenomanian, Apertotemporalis baharijensis Stromer 1934; (Antunes & Broin 1988; Broin 1988a; Dacque 1912; Lapparent de Broin & Werner 1998; Stromer 1934); material destroyed. 59 Idfu, just N of Idfu, Nile Valley, a 25° N, 33° E, Turonian-Early Senonian, Chelonii indet.: Dacque (1912) (unverified). Ammonite Hills, NW Dakhla Oasis, ca. 26° N, 26° E, SSW Cairo, Western desert, Dakhla Formation, Maastrichtian (littoral and marine): Arenila krebsi Lapparent de Broin & Werner 1998; Zolhafah bella Lapparent de Broin & Wemer 1998 and Bothremydidae indet. of Bothremys group, Taphrosphys cf. sulcatus, cf. Taphrosphys sp., aff. Tasbacka sp. (see Barthel 1980; Barthel & Herrmann-Degen 1981; Quaas 1902 in Lapparent de Broin & Wemer 1998; Dacque 1912; Wanner 1902); TUB. Eocene (mixed, continental and marine forms): Fayum, S Cairo area, Qasr El Sagha Formation, at: Birket el Kurun (Andrews 1901, 1903), Abusir, Dineh (Dimeh), N of West border of Birket el Kurun, Qasr es Saga (Reinach 1903, Dacque 1912); upper Mokattam, Late Eocene, Stereogenys cromeri Andrews 1901; Schweboemys antiqua (Andrews 1903) (see Andrews 1906, pi. 25, fig. 1; Wood 1970) (= ‘Podocnemis stromeri Reinach, 1903’ and ‘Podocnemis stromeri var. major, Reinach 1903’); genus indet., ‘Stereogenys’ podocnemoides Reinach 1903 (a Neochelys group member, n.g.?); Pelomedusoides indet. (NHM); Egyptemys Wood etal. 1996: Psephophorus eocaenus Andrews 1901; genus indet., 'Thalassochelys’ libyca Andrews 1901 [a cheloniid: ‘Thalassochelys (Chelone?) libyca Andrews’; ‘Thalassochelys Boul. (Caretta Raf.)’ in Dacque 1912]; Cheloniidae indet.: ^Trachyaspis: Dacque (1912) (= ‘Trachyaspis cf. aegyptiaca Lydekker 1889’ in Reinach 1903); material MNHB, NHM and SMNS only observed (other material in CM or destroyed). Oligocene: Fayum, NW Birket el Kurun, Dimeh, Tamieh, ‘Schweinfurthplateau’ (Dacque 1912), S Cairo area, Qatrani Formation, Early Oligocene: Stereogenys libyca Andrews 1903; aff. Erymnochelys fajumensis (Andrews 1903) = Dacquemys palaeomorpha Williams, 1954b: ? [‘Podocnemis fajumensis Andrews 1903’, ‘Podocnemis blanckenhorni Reinach 1903’, ‘Podocnemis blanckenhorni var. ovatum Reinach, 1903’, ‘Pelomedusa progaleata Reinach 1903’, ‘Dacquemys palaeomorpha Williams 1954b = Erymnochelys fajumensis’ (Andrews 1903): ? in Williams 1954c] (Dacque 1912; Broin 1988a); Gigantochersina ammon (Andrews andBeadnell 1903) (‘Testudo ammon Andrews & Beadnell 1903’ = T. is is, Andrews 1906’? = T. beadnelli Andrews 1906’?) (Andrews 1904; Chkhhikvadze 1989; Lapparent de Broin & Van Dijk 1999); NHM and SMNS material only observed, other in CM. Mio-Pliocene (marine): Suez Canal, Tertiary indet., probable Miocene, Trachyaspis aegyptiaca Lydekker 1889 (see Weems 1980, erroneous attribution to Syllomus Cope 1896, instead of Trachyaspis Meyer 1843); NHM. Early Miocene: Moghara, ca. 30° N 29° E, Early Miocene, Burdigalian, Orleanian, MN 3+, ca. 18 myr, aff. Erymnochelys aegyptiaca (Andrews 1900), ‘Erymnochelys aegyptiaca (Andrews 1900)’: Williams (1954c); genus indet., ‘Podocnemis’ bramlyi Fourtau 1920 (= ?Schweboemys, rather than a Bothremydidae?); genus indet., ‘Sternothaerus ’ blanckenhorni Dacque, 1912 (= a member of the Schweboemys group?) (‘a precursor of Peltocephalus: Williams 1954c’); Erymnochelyidae indet. (‘Podocnemide’: Williams, 1954c); aff. Allaeochelys sp.? and Cyclanorbinae ( ‘Trionyx senckenbergianus Reinach 1903’, pi. 17, figs. 2, 5 and 6 ) (Broin 1983; Lapparent de Broin & Gmira 1994; Wood 1970). Wadi Faregh, ca. 30° N, 30° E, Early Miocene, Burdigalian,Orleanian, MN 3+, ca. 18 myr, aff. Allaeochelys sp., (“ICyclanorbis n. sp.’: Dacque 1912; ‘Trionyx senckenbergianus Reinach 1903’ ex parte); ?Cyclanorbinae ( ‘Trionyx senckenbergianus Reinach 1903 ’ ex parte; Trionyx sp.: Dacque 1912) (Broin 1983; Lapparent de Broin & Gmira 1994; Wood 1970). El Arag area, ca. 28°40' N, 26°30' E, N of El Bahrein, Th. Monod coll., Egypt, Neogene, Burdigalian? Cf. or aff. Erymnochelys, Chelonii indet. (large Testudininei indet.?); MNHN. Late Miocene: Wadi Natrun (Natrontal) (Djebel El Muluk, Garet El Muluk, Der Baramus), Late Miocene, Late Turolian, MN 13+ (ca 6,5 myr), Pelusios sinuatus (A. Smith 1838) ( ‘Sternothaerus dewitzianus Reinach 1903’: Dacque 1912'; = ‘Pelomedusa pliocaenica Reinach 1903 ’: Dacque (1912) (Broin 1969, 1983). Trionyx sp.: ‘Trionyx pliocaenicus Reinach 1903’, ’Trionyx sp. = Trionyx pliocaenicus Reinach 1903', ‘protriunguis serie, Reinach 1903’: Dacque (1912) (Broin 1983; Lapparent de Broin & Gmira 1994). Mauremys sp. (‘Ocadia n. sp. ind.’: Dacque 1912) (Lapparent de Broin & Van Dijkl999). Trachyaspis cf. aegyptiaca Lydekker 1889 (‘indet’: Dacque 1912); material partly destroyed. Holocene: Abu Balias, 200 km SW Dakhla Oasis, Mudpans, site 83/89, ca. 8300 yr BP, site 85/50-1, ca. 6800 yr BP, site 85/51-3, ca. 6800, Testudininei; Van Neer & Uerpmann (1989). Toukh, close to Negadah, 30 km N Louxor, Neolithic (Amratian and Gerzean), Trionyx triunguis (Forskal 1775): Joleaud(1936). Anteopolis (Siout, Asyut), Nile Valley ca. 400 km S Cairo, Pleistocene-Holocene tomb, ICyclanorbis senegalensis: ‘Emyda sivalensis ’ Lyd. : Parona (1918) = ‘Emyda sivalensis’ Lydekker 1885, a junior synonym of Lissemys punctata (Lacepede 1788), the Indian form which has a similar decoration to that of this African form C. senegalensis; ?Trionyx sp.: Parona (1918); (unknown localization) (Gautier 1984; Joleaud 1936). Fayum, Birket el Kurun, Pleistocene-Holocene; Trionyx triunguis (Forskal 1775), TUB; ‘Chelonians’: Andrews (1906). Adaima, between Luxor and Esna, pre-dynastic site, Neolithic, Trionyx triunguis: Midant-Reynes et al. (1993). Berenike, Egyptian Red Sea Coast, Roman period, 2000-1500 yr BP, Chelonia mydas; ?’Centrochelys sulcata (?’ Geochelone sulcata'): Van Neer & Ervynck (1998) (Van Neer & Lentacker 1996). ETHIOPIA Cretaceous: Abay River Basin (Blue Nile Basin), North of Addis-Ababa, Mugher Mudstone, Early Cretaceous, aff. Araripemys, Podocnemidoidea indet.: fauna comparable to that of the upper part ofthe ‘Continental Intercalate’ of Sahara, late upper part (Kem Kem of Morocco, In Abangarit of Niger, Albian-Early Cenomanian prior to marine transgression) (Schmidt & Wemer 1998; Wemer 1995, 1996); TUB (Broin det.). Plio-Pleistocene o f Omo River Basin (Arambourg 1947; Arambourg et al. 1967, 1969; Bonnefille etal. 1973a,b; Brown et al. 1985), NME, MNHN (Broin det.): Yellow Sands, Mursi Formation, older than 4 myr, Pliocene, Pelusios cf. sinuatus (A. Smith 1838), Cyclanorbis cf. elegans (Gray 1869). Shungura Formation, Plio-Pleistocene, chelonians seen from bed A l, ca. 3,79 to beds H sup., K 11, ca. 1,6 myr: Pelusios sinuatus (including ‘Sternothaerus rudolphi' Arambourg 1947) and P. cf. sinuatus (beds Al to G13), P. adansoni (Schweigger 1812) (beds G4-13, Omo 75, H Upper Kalam 11); cf. Trionyx sp. (bed A3), Cyclanorbis elegans (bed B9-10); Cycloderma frenatum Peters 1854 (beds A l, B2, E, G27); Testudininei indet., large sp., Stigmochelyspardalis (Bell 1828) group (beds B9-12, C5 to C9, D3): Arambourg (1947) (Broin 1979). Pliocene-Holocene o f Awash Valley (NME) (Broin det.): Pliocene o f Central Afar (Johanson 1996; Taieb et al. 1976), PALAEONTOLOGIA AFRICANA VOLUME 36 2000 I 60 Hadar Formation, Pliocene; Sidi Hakoma Member, ca. 3,40 to 3,28 myr, Pelusios gabonensis (Dumeril 1856), Pelusios sp., Centrochelys sulcata (Miller 1779) (SH-SH3), large Stigmochelys sp. (SH-SH3; SH1-SH2); Denen-Dora Member, <3,18 myr, Pelusios gabonensis, large (DD1- DD3), gigantic aff. Stigmochelys sp. (DD1-DD3), indet. round eggs (Chelonii? Testudininei or Trionychidae; not revised) (DD3=KH 1-KH2). Central Ledi Basin, Pelusios cf. gabonensis, P. sp., Cycloderma frenatum. West of Central Ledi Basin, Pelusios cf. gabonensis, C. frenatum. Amado Basin?, Geraru Basin, cf. Cycloderma sp. Holocene, Delta, Loc. AL 42-1, ?cf. Centrochelys sp., large form. GHANA Protohistoric: Mole National park, Nyanga camp, Mole River, 9°32' N, 1°57' W, Cyclanorbis elegans (Gray 1869), C. senegalensis (Dumeril & Bibron 1835): Hughes (1979). KENYA Early Miocene o f Koru-Songhor-Muhoroni area: Koru, ca. 19-20 myr (Pickford et al. 1986b), ?Cyclanorbinae: Lapparent de Broin & Gmira (1994); NHM. Songbor, ca. 19-20 myr (Pickford et al. 1986b), Kinixys erosa (Schweigger 1812): Meylan & Auffenberg (1986); NHM. Mteitei area, Chelonii indet.: Pickford (1986). Mio-Pleistocene o f Lake Victoria area: Rusinga Island, NE Lake Victoria, Upper Katwanga series, Early Miocene, ca. 18 myr (Pickford et al. 1986b), Pelusios rusingae Williams 1954a; lmpregnochelyspachytectis Meylan & Auffenberg 1986; other testudinineine (BM, NH, R 6422)?; NHM. Formations Wayondo, Hiwegi and Kulu, Chelonii indet.: mentions in Pickford (1986) (‘Testudinidae’, ‘Pelomedusidae’, i. e. undecorated chelonians such as Testudinidae, Pelomedusidae, Podocnemididae etc.; and ‘Trionychidae’, i.e. decorated forms, possible Cyclanorbinae, not probable Carettochelyidae and Trionychinae at that time). Mfwangano Island, Walangani and Higeni Formations, Early Miocene, ca. 18 myr (Pickford 1986), Chelonii indet. (‘Testudinidae’: Pickford 1986, i.e. undecorated chelonians; possible Testudinidae but also Pelomedusidae, Podocnemididae or other indet.). Uyoma Peninsula, Early Miocene, as Rusinga, Chelonii indet., possible Cyclanorbinae (‘Trionyx’, ‘Cycloderma’, Pikford 1986). Karungu, NE Lake Victoria, Kachuku beds, Early Miocene, ca. 18 myr (Pickford et al. 1986b), aff. Erymnochelys sp., young ( ‘Podocnemis aegyptiaca': Andrews 1914), bed 22; aff. Cycloderma victoriae (Andrews 1914) (‘Cycloderma victoriae Andrews 1914’), bed 31; Testudininei indet., ? Stigmochelys group?: ‘Geochelone crass a (Andrews 1914)’: Meylan & Auffenberg (1986), bed 31; NHM. Gwasi Peninsula, Simenya, Early Miocene, Chelonii indet.: Pickford (1986). Ombo, Maboko Formation, Early Middle Miocene, ca. 15-16 myr (Pickford et al. 1986b), Cyclanorbinae indet.; Lapparent de Broin & Gmira (1994); NHM; Chelonii indet. ( ‘Pelomedusidae’, ‘Trionychidae’: Pickford 1986, i.e. undecorated Chelonii, Testudininei and/or Pelomedusoides, and probable Cyclanorbinae). Homa Peninsula: Kanam (1), Homa and Kanam beds, Pliocene, ca. 4 myr, Chelonii indet. (‘Pelomedusidae’, ‘Trionychidae’, Pickford 1986, i.e. undecorated Chelonii, Testudininei and/or Pelomedusoides, and Trionychoidea). Kanam (2), Kanjera beds, Plio-Pleistocene, ca. 1-2 myr, Chelonii indet. ( ‘Pelomedusidae’, Pickford 1986, i.e. undecorated Chelonii, Testudininei and/or Pelomedusoides). Rawi (3), upper Kanjera beds, Pleistocene, ca. 1 myr, ?aff. Stigmochelys sp. { ‘G. aff. pardalis’: Broin 1979: possible Stigmochelys group), NHM. Mio-Pliocene o f Lake Baringo Basin, Baringo district (Bishop & Chapman 1970; Bishop & Pickford 1975; Bishop et al. 1971; Pickford et al. 1986b, 1993; Wood 1973b): Ngorora Formation, Middle Miocene, ca. 11-12 myr, Pelomedusoides including Pelusios sinuatus (‘Pelomedusidae’, ‘Pelusios cf. sinuatus Smith’), Cyclanorbinae indet. ( ‘Trionychidae’, ‘Trionyx’ sp.’), Testudininei indet. (‘Testudinidae, Testudo sp.’) (Bishop & Pickford 1975; Bishop & Chapman 1970; Bishop et al. 1971). Giant Testudininei indet., loc. 2/106, Member D, ca. 11,7 myr and Chelonii indet., a giant freshwater chelonian ca. 2 m long, at Ngeringuerwa, ca. 10 myr, Baringo Basin, Miocene (pers. comm. M. Pickford). Mpesida beds, Late Miocene, ca. 6,2 myr, Trionychoidea indet. (?Carettochelyidae, ?Cyclanorbinae: ‘Trionychidae indet’), Testudininei indet. (‘Testudinidae’: Bishop et al. 1971). Lukeino Formation, Late Miocene, ca. 5,8-6 myr, Pelomedusoides indet. ( ‘Pelomedusidae indet.’ i.e. ?Pelomedusidae ± Podocnemididae), Trionychoidea indet. (?Carettochelyidae, ?Cyclanorbinae: ‘Trionychidae indet.’), Testudininei indet. (‘Testudinidae’) (Pickford 1975; Bishop et al. 1971). Kaperyon Formation, Pliocene, ca. 5 myr, Trionychoidea indet. (?Carettochelyidae, ?Cyclanorbinae: ‘Trionychidae’, Bishop et al. 1971). Chemeron Formation, Pliocene, <2,2 myr, Cyclanorbis sp. ( ‘Cycloderma sp’. in Meylan 1990) (Bishop etal, 1971; Lapparent de Broin & Gmira 1994). Aterir beds, Pliocene (> 4 to ca. 2,2 myr), Pelomedusoides indet. (possible Pelomedusidae and still possible Podocnemididae at that time: ‘Pelomedusidae’; Bishop etal. 1971). Chemoigut beds, Pleistocene, 1,2 myr, Pelomedusoides indet. (‘Pelomedusidae’, i.e. probable Pelusios), Cyclanorbinae indet. (‘Trionychidae’) (Bishop et al. 1971, 1975). Lake Turkana, (Brown et al. 1985): Mio-Pliocene o f Kerio River Basin, SW Lake Turkana, (Patterson et al. 1970; Behrensmeyer 1976) NMK (Broin partly observed), Lothagam Hill, Late Miocene, above 8,5 myr, around 6 myr at Lothagam 1, to Pliocene, older than ca. 3,8 myr at Lothagam 3: Lothagam 1, aff. Erymnochelys sp. A ( Podocnemis sp. A’ of Patterson et al. 1970; Witmer 1990?), Kenyemys williamsi Wood 1983 ( ‘Podocnemis sp. B’ of Patterson et al. 1970); aff. Cycloderma debroinae (Meylan et al. 1990), lowest horizon; Testudininei, indet. group ( ‘Geochelone’ sp.); Lothagam Unit 2, Chelonii indet. (‘turtles’), Lothagam 3, aff. Erymnochelys sp. A (‘Podocnemis sp. A: Witmer 1990?), ?Cyclanorbinae (‘Trionychidae indet.’), Testudininei indet. group (‘Geochelone’ sp.); undefined horizon (‘Pliocene’), Cycloderma frenatum Peters, 1854: Meylan et al. (1990). Pelomedusoides indet. (abnormal Pelusios?) (‘Chelonii indet.’: Wood 1976). Kanapoi, 50 km S Lothagam, Pliocene, from < 4 myr? to < 2,6 myr. Aff. Erymnochelys sp. A { ‘Podocnemis sp. A’), Cyclanorbinae (‘Trionychidae indet.’, Patterson et al. 1970), including ‘Cyclanorbini indet.’ in Meylan et al. (1990) and Cyclanorbis turkanensis, Meylan et al. 1990, bed E; Testudininei indet. {Stigmochelys group?) {Geochelone crassa Andrews, 1914 in Meylan & Auffenberg 1986). Ekora, ca. 23 km NE Kanapoi, Pliocene (just above 4 myr), Aff. Erymnochely sp. A ( ‘Podocnemis sp. A’ of, Patterson et al. 1970), Testudininei indet. group (‘ Geochelone’ sp.). Plio-Pleistocene o f East Lake Turkana, Koobi Fora beds, NMK (Broin 1979, 1983; Harris 1978; Harris et al. 1988; Lapparent de Broin & Van Dijk 1999; Meylan & Auffenberg 1986; Meylan etal. 1990; Wood 1979), Koobi Algi Formation, areas 201, 202, 204, Early Pliocene, ca. 3,9-4,5 myr, Pelusios sinuatus (A. Smith 1838). Koobi Fora Formation (Brown et al. 1985): Lower member, Pliocene: area 116, above Tulu Bor 61 tuff, (ca 3,35 myr), below KBS tuff (ca 1,88 myr), Pelusios sinuatus. Lower-upper member limit, Late Pliocene: - 25m below tuff KBS (ca 1,88 myr) area 102,130-1, P. sinuatus, area 102, Cyclanorbis elegans: Meylan et al. (1990); 20 m below KBS, area 105, Cycloderma frenatum and Cyclanorbis senegalensis (Dumeril & Bibron 1835), Trionychidae indet.: Meylan et al. (1990); - 20 m below KBS, unknown area, C. elegans (Gray 1869) or C. turkanensis : Meylan etal. (1990); - below to above KBS, KFII, Pelusios sp.; - just above KBS, area 130-1, P. sinuatus; - upper member, above KBS, ca. 1,88 myr (Pliocene), around ‘Okote tuff (ca 1,57 myr, age of J7 in Shungura F. of Omo, Pleistocene) and below Chari tuff (ca 1,39 myr), area 104-5, 104-A, Trionyx triunguis (Forskal 1775): Broin (1983), Harris (1978), Meylan et al. (1990); (Trionyx cf. T. triunguis: Wood 1979); area 103, 104-A, Cycloderma cf. frenatum', area 104, Testudininei indet. (large sp.); - undefined horizon and area: Trionychinae indet, Cyclanorbis elegans: Meylan et al. (1990); gigantic Testudininei indet. (Harris pers. comm.). LESOTHO Holocene: Ntloana Tsoana, north-western Lesotho, 8780 ±80, 12110 ±120 yr BP, Chelonii indet.: Mitchel (1993, in Branch et al. 1995), Tloutle, north-western Lesotho, 6140 ±100 yr BP, Testudininei indet.; see Carter Mitchell & Winnicombe (1988, in Branch et al. 1995), Sehonghong Rockshelter, Qacha’s Nek District, western Lesotho, 1400±50 yr BP, Testudininei indet.; see Carter, Mitchell & Winnicombe (1988, in Branch et al. 1995), Hololo Crossing, 28°44’S; 28°, 27’E, 330-260 yr BP, Testudininei indet.; see Mitchell, Parkington & Yates (1994, in Branched al. 1995). LIBYA Eocene-Early Oligocene: Dor et Talha, E Fezzan, S Syrta Major, 25°45' N, between 17°50' and 19° 15' E: - Dor et Talha, 15 and 80 km E of Oriental border of Djebel Harouj el Assoued, Podocnemidoidea indet., probable Podocnemididae: ‘Chelonian plates’: Bellair et al. (1954), the same layer as Djebel Coquin: Arambourg (1963); Late Eocene; MNHN, coll. Lefranc; - = Djebel Coquin, 25°45' N, between 17°50' and 19° 15' E, Pelomedusoides indet., probable Podocnemididae: ‘palustral tortoises’: Arambourg (1963); ‘palustral turtles’: Arambourg & Magnier (1961), Late Eocene, Priabonian (collected? unknown localization); - = Dor et Talha, 25°45' N, between 17°45' E and 19°05' E, Evaporite Unit, Late Eocene, Chelonii indet.; Idam Unit, ?Early Oligocene, Pelomedusoides indet., probable Podocnemididae: ‘Pelomedusidae’: Savage (1969) (Wight 1980). Oligocene: Zella Oasis, S Syrta Major (Sirte Desert), 28°30' N, 17°37' E, Chelonii indet.: ‘paludal turtles’: Arambourg (1963); ‘palustral turtles’: Arambourg & Magnier (1961) (collected? unknown localization). Miocene: Djebel Zelten, SE Syrta Major, 28°45' N, 19°30' E, Early Miocene, Burdigalian, Early Orleanian, MN 4+, ca. 16,5 myr; Podocnemididae indet. (including Stereogenys?: an epiplastron): ‘palustral turtles’: Arambourg (1963), Arambourg & Magnier (1961), cf. Centrochelys sp., large (a femur conform to C. sulcata', see Lapparent de Broin & Van Dijk 1999) and very large (a plate); MNHN (Broin det.). Sahabi: Bir Guetin, Gara el Beda, Cyrena'fca, SSE to Benghazi, Late Miocene, Late Turolian, MN 13+, ca. 6,5 myr (Geraads 1989), Trionyx triunguis (Forskal 1775): ‘ Trionyx sp.’: D’Erasmo (1934); ‘Trionyx cf. triunguis’’: Wood (1987); Centrochelys aff. sulcata (Miller 1779): ‘cf. Geochelone’: Wood (1987) (Lapparent de Broin & Van Dijk 1999). Holocene: Djebel Zelten, surface coll. Magnier, Cyclanorbis senegalensis (Dumeril & Bibron 1835), small form; MNHN (Broin det.). ?Cyrenai'ca, Libyan desert, unprecised, surface coll. Magnier? (Coll. Arambourg) (or from more western part of Africa), Pelusios sp., sub-group P. castaneus (Schweigger 1812); MNHN (Broin det.). MALAWI Cretaceous: ‘Nyasaland’, NW Lake Malawi, Dinosaur beds of the Mwakasyunguti area, Siwe Valley, Karonga district, Early Cretaceous, Lupata group, upper member (new finds, Jacobs et al. 1996); Platycheloides nyasae Haughton 1928; SAM. Pliocene: Chiwondo beds, Pliocene, 2,5-4,8 myr, Pelusios sinuatus (A. Smith 1838): pers. comm. Wood (1971) {Pelusios: Wood 1973b), Cycloderma frenatum: Meylan et al. (1990), Wood (1979). MALI ‘Continental Intercalate’ o f Sahara, late upper part, Early Cretaceous, Albian-Cenomanian - prior to Cenomanian transgression (see Lapparent 1960): Tikarkas, 4 km S, 115 km NW Tessalit (Bellion et al. 1992); Bothremydidae indet.: ‘Eusarkia’ sp.: Bergounioux & Crouzel 1968; not a Taphrosphys, contra Lapparent de Broin & Wemer (1998); MHNT. Maastrichtian (littoral): Tagnout Chaggeret, Erg Ine Sakane, MK 42 loc. (Broin 1983), Bothremydidae indet., aff. Arenila sp., Niger emys group: Lapparent de Broin & Wemer (1998); MNHN. In Afarag, E balise 560, S Tanesrouft, Bothremydidae indet., coll. A.F. de Lapparent. Palaeocene-Eocene o f the Tilemsi valley (Lavocat & Radierl953; Buffetautl980) (littoral, marine) MNHN: In Farghas, Cheit Keni, Palaeocene, Bothremydidae indet., Taphrosphys sp. (Mali C, Lapparent de Broin & Wemer 1998). Samit, Ypresian, Bothremydidae indet. (Mali B, Lapparent de Broin & Wemer 1998). Tamaguilelt, Lutetian, Bothremydidae indet. (Broin det.). Holocene: Taoudenni Basin, North Mali (MMB; MNHN), Araouan (Arawan, Araouan(e) and Guir (Gir) area, Djouf, unspecified Holocene, old coll. (see data in Broin 1983: Gallay 1966; Joleaud 1934, 1936 - Capitaine Poggi coll., 50 km NNW Araouan-; Monod 1958; Roman,1935); Trionyx triunguis (Forskal 1775), figured in Monod (1958), 45 km NW Arawan, and in Gallay (1966), from Outeidat, as Trionyx triunguis, pro parte, figs. 9, 15, and as indetermined vertebrate, figs. 10, 44; Cyclanorbis senegalensis (Dumeril & Bibron 1835), small form (‘Clarias’ pro parte, fig. 2, and ‘Trionyx, T. triunguis size’, fig. 6 , in Roman, 1935, pi. 4, 10 km S Guir; figured as Trionyx triunguis pro parte, 16-18, in Gallay, 1966, Outeidat, fig. 9). Hassi el Abiod, 19° 10' N, 3°50' W, 70 km NW Araouane, 6970+130 yr BP, Pelusios adansoni (Schweigger 1812), P. castaneus (Schweigger 1812), Trionyx triunguis, Cyclanorbis senegalensis, small form: Broin (1983), and new coll. Petit-Maire et al. 1983. Erg Ine Sakane, 21°10' N, 0°40' W, 9500-6400 yr BP, Pelusios castaneus, Cyclanorbis senegalensis, small form: Broin (1983). South Mali, Kobadi, KBD 84, E Nampala, 15°21’30" N, 5°29’30" W, Peul country-Mauritania frontier (pers. comm. M. Raimbault), Trionyx cf. triunguis (Broin det.). 1 62 MALTA Miocene: Bothremydidae genus indet., 'Podocnemis’ lata Ristori, 1894: Lapparent de Broin & Wemer (1998), Aff. Cycloderma melitensis (Lydekker 1891) ( ‘Trionyx’ melitensis Lydekker 1891): Lapparent de Broin &Van Dijk (1999). Pleistocene: Mauremys leprosa (Schweigger 1812) (‘Lutremys EuropceaT in Leith-Adams 1877), from Zebbug cavern; Testudininei genus indet., 'Testudo’ robusta Leith-Adams 1877; = ‘ Testudo Spratti Leith-Adams, 1877’, a giant tortoise from Benghisa Gap, Mnaidra Gap and Zebbug, ossiferous caverns. MAURITANIA Holocene: Chami, (well of) or Nouaferd, 25 km E Cape Tafarit, Neolithic, ca. 2100 to 3500 yr BP, Centrochelys cf. sulcata (Miller 1779): Broin (1983) (Petit-Maire 1979); MNHN deposit. MOROCCO Middle Jurassic: El Mers, Middle Atlas, 100 km S Fes, Bathonian, Chelonii indet.; (Termier et al. 1940) one fragment in MNHN. Early Cretaceous o f High Oriental Atlas: Anoual area, Oussikis ans Ksar Metlili, Barremian, Chelonii indet., ?Pelomedusoides indet., cf. Taquetochelys sp.: Gmira (1995) ( Sigogneau-Russell etal. 1988,1990); MNHN deposit. ‘Continental In tercalate’ o f Sahara, late upper part, Early Cretaceous, Albian-Cenomanian prior to Early Cenomanian of Baharija and to Cenomanian transgression (see Lapparent 1960; Lavocat, 1954), MNHN; part in CMN: Hamada of Guir, Kem-Kem, E and S to Tafilalt, S Maroc, Araripemys sp., Pelomedusoides indet., Bothremydidae indet., Podocnemididae indet, Hammadachelys escuillei Tong & Buffetaut 1996; (Fuente & Lapparent de Broin 1997; Gmira 1995; Lapparent de Broin & Wemer 1998; Russell 1996). Phosphates (marine, littoral) (Arambourg 1952a): Maastrichtian, Oued Erguita, N Taroudant, Oued Sous tributary, Chelonii indet. (‘chelonians indet’.: Ambroggi & Arambourg, 1951) (unknown localization). Benguerir, Ganntour Basin, aff. Euclastes sp. (‘Aff. Rhetechelys sp .’: Gmira 1995), G. Termier coll.; Chelonioidea indet: Moody (1976). Oued Zem, Bed III, E Ouled Abdoun Basin, Chelonioidea indet., new undescribed large form (private coll.). Between Kouribga and Oued Zem, Ouled Abdoun Basin, Chelonioidea indet., giant pre-cheloniid, SMNS. Palaeocene, Benguerir, Ganntour Basin, aff. Taphrophys sp. (‘close to Podocnemis', Pelomedusidae indet.: Moody 1976) (Antunes & Broin 1988; Lapparent de Broin & Wemer 1998); Musee du Ministere de PEnergie et des Mines, Rabat. Palaeocene- Ypresian, Oued Zem, E Ouled Abdoun Basin, Bothremydidae indet., Osteopygidae indet. (sold osteopygid skulls artificially linked to pleural discs of Taphrosphys), new undescribed possible Dermochelyidae (private coll.). Paleogene o f Ouarzazate Basin, N of Oriental border, Anti Atlas (Gheerbrant 1987; Gheerbrant et al., 1993): Palaeocene, Several localities with chelonians, Jbel Guersif Formation, Thanetian, including: Ilimzi, aff. Pelomedusa sp., Broin det. (in Nicolas 1984, see Gmira 1995); Pelomedusoides indet., MNHN deposit. Adrar Mgorn, Pelomedusoides indet., MNHN deposit. Ypresian, N’Tagourt 2, Ait Ouarhitane Formation, Chelonii indet. Pliocene: Ahl Al Oughlam, carriere Deprez, Casablanca, Occidental Morocco, Pliocene, ca. 2 to 2,5 myr, ?cf. Centrochelys: ( ‘Geochelone s.l. sp’.: Raynal et al. 1990); Testudo aff. kenitrensis (Gmira et al. in prep.); INSAP. Pleistocene-Holocene (in Gmira 1995: see Ennouchi 1949, 1954, 1969, 1976, Michel 1988, 1990): Occidental Morocco, Kenitra, Middle Pleistocene, Inter Amirian-Tensiftian, Testudo kenitrensis Gmira 1993a (Gmira 1993b, 1995); MNHN et FSR. Carriere Thomas I, (Thomas Quarry I), Late Pleistocene, Tensiftian, ca. 400 000 yr BP, Hublin, 1985, Testudo graeca Linnaeus, 1758, FSR. Gmira (1995). Rabat 8 , 9, 10, coast from Rabat to Temara a, small dune (Choubert & Mar9ais 1947), Tensiftian, lower part of Late Pleistocene, Testudo g. graeca, Testudo sp., FSR. Rabat 6 , coast from Rabat to Temara b, pink sandstones, Temara Formation, Late Pleistocene, Tyrrhenian, Trionyx sp., FSR. A'in Rohr, Late Pleistocene, Early Soltanian, Testudinidae indet. { ‘Testudo': Ennouchi 1949). Jebel Irhoud, Late Pleistocene, Early Soltanian, Testudo graeca: Gmira {in Amani & Geraads 1993; Gmira 1995); FSR. Ain Bahya, Late Pleistocene, Soltanian, Testudininei indet.: Gmira (1995) (‘Testudo g. graeca': Michel, 1988, 1990); INSAP. Oualidia, Late Pleistocene, Soltanian, Testudo g. graeca', FSR. El Khenzira, near El-Jadida,Cap Blanc, Cave 1, bed C, Late Pleistocene-Holocene, Epipalaeolithic, Testudinidae indet. (‘Testudo sp. ‘: Ruhlmann 1936). Dar Es Soltane, Late Pleistocene-Holocene, Soltanian-Rharbian, Mauremys sp, Testudo cf. graeca', FSR. Bouknadel, Middle-Late Pleistocene, Testudinidae indet. (Michel 1990); INSAP. Doukkala II, Late Pleistocene-Holocene, Soltanian- Rharbian, Mauremys leprosa (Schweigger 1812), Testudo g. graeca, INSAP. Mehdia, Holocene, Neolithic probable, Testudo cf. graeca', FSR. Toulkine-Bou Ben Adam, Neolithic, Testudo cf. graeca; FSR. Gmira (1995). Oriental Morocco, Taforalt, 55 km NW Oujda (Roche 1953, 1963), Late Pleistocene, Aterian, Mauremys leprosa, Testudo g. graeca; MNHN (Broin det. and Roubet 1966). Rhafas Cave, Late Pleistocene, Soltanian and Middle Holocene, Testudinidae indet., El Heriga, Late Pleistocene, Soltanian, and Holocene, Testudinidae indet., Abri Rhirane, Late Pleistocene, Soltanian, and Holocene, Testudinidae indet., Oued el Haij Terrace, Jorf el Anngra, Holocene, Testudinidae indet.: ‘ Testudo graeca, T. g. graeca’: Michel (1990), INSAP. Abri Bou Guennouna, Holocene, Neolithic, Testudo cf. graeca, Testudinidae indet.; INSAP. Kheneg Kenadsa, Tendrara, Neolithic, Testudinidae indet.: ‘Testudo g. graeca': Jodin (1956). MOZAMBIQUE Pleistocene-Holocene o f Zambezi (NHM): Tributary stream of the Zambezi about 8 miles below Mazzaro, right bank, 40 miles from the present coast-line. Collected during the first Livingstone Expedition, 1858, by Sir John Kirk. Pleistocene-Holocene. Due to the association of various extant mammals, including a Cape buffalo, etc., and the presence of human activity (pottery) in the alluvions collected by the stream, possibly Holocene, Neolithic; but this activity is not clearly established as contemporaneous with the fossils (Kirk 1864; Murchison 1864a,b): cf. Cycloderma frenatum. NAMIBIA Miocene, South-West to Namib Desert, Sperrgebiet = Diamond area, SW Namibia (MSGN) (Broin det.): 63 Western part, from North to South, Early Miocene, Fiskus, ca. 19-20 myr, Testudininei n.g. a (large form), sp. Grillental, ca. 19-20 myr, Testudininei n.g. a (large form), sp. Elisabethfeld, ca. 19-20 myr, Testudininei n.g. a (large form), sp. namaquensis (Stromer 1926) = 'Testudo namaquensis Stromer 1926’ (material destroyed in Munich); [‘Geochelone namaquensis (Stromer 1926)’ and ‘G. stromeri Meylan & Auffenberg 1986]. Langental, ca. 19 myr, Pelomedusa n. sp., Testudininei indet., n.g. a (large form), n. sp. (also AMNH) and n.g. b, n. sp. (small form, Stigmochelys group,?). Glastal, ca. 19 myr, Testudininei indet. n.g. a (large form) sp. Southern part, N Orange River, (from North to South and West to East), Early-Middle Miocene, Rooilepel: - wardi level, Middle Miocene, ca. 10-12 myr, Testudininei n.g. a sp .; - laini level, Middle Mocene, ca. 8 myr, Testudininei n.g. a sp. Karingarab, wardi level, Middle Miocene, ca. 10-12 myr, Testudininei n.g. a (large form) sp. North of Gypsum Plate Pan, ca. wardi level, Middle Miocene, 10-14 myr, Testudininei n.g. a (large form) sp. Arrisdrift, Early Miocene, ca. 17 myr, Eymnochelyinae indet., Testudininei n.g. a (large form), n. sp. [‘Geochelone namaquensis (Stromer 1926’): Meylan & Auffenberg 1986]; Testudininei n.g., n. sp. ( ‘Chersina sp.’: Meylan & Auffenberg 1986); (Hendey 1978). Auchas, Early Miocene, ca. 18 myr, Erymnochelyinae indet., Testudininei n.g. a (large form), n. sp. (also OMS). Historic site: Brandberg, north of Namibia, 1600-1750 AD, Stigmochelys pardalis: Cooper & Branch (1999). NIGER ‘Contin