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A New Polypterid Fish: Polypterus Faraou Sp. Nov. (Cladistia

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A New Polypterid Fish: Polypterus Faraou Sp. Nov. (Cladistia Blackwell Science, LtdOxford, UKZOJZoological Journal of the Linnean Society0024-4082The Lin- nean Society of London, 2006? 2006 146? 227237 Original Article A NEW POLYPTERID FROM THE LATE MIOCENE OF CHADO. OTERO ET AL. Zoological Journal of the Linnean Society, 2006, 146, 227–237. With 10 figures A new polypterid fish: Polypterus faraou sp. nov. (Cladistia, Polypteridae) from the Late Miocene, Toros-Menalla, Chad OLGA OTERO1*, ANDOSSA LIKIUS2, PATRICK VIGNAUD1 and MICHEL BRUNET1 Downloaded from https://academic.oup.com/zoolinnean/article/146/2/227/2631032 by guest on 26 October 2020 1Laboratoire de Géobiologie, Biochronologie et Paléontologie humaine, CNRS UMR 6046, Faculté des Sciences fondamentales et appliquées, Université Poitiers, 40 av. du Recteur Pineau, F-86 022 Poitiers cedex, France 2Département de Paléontologie, Université de N’Djaména, N’Djaména, Tchad Received June 2004; accepted for publication July 2005 Polypterus faraou sp. nov. (Cladistia, Polypteridae) from the Late Miocene of Toros-Menalla (western Djourab, Chad) is described on the basis of a subcomplete articulated skeleton preserved in three dimensions. This is the first time such a complete fossil polypterid skeleton has been described. It is the only verifiable fossil record for the genus Polypterus. P. faraou closely resembles P. bichir and P. endlicheri, extant fish found in the Chad–Chari system. Intrarelationships among the polypterids are not yet resolved: however, P. faraou shows a primitive shape of the body and a primitive shape of the opening of the lateral line on the scales, similar to that of three living species (P. bichir, P. endlicheri and P. ansorgii). © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society, 2006, 146, 227–237. ADDITIONAL KEYWORDS: anatomy – Anthracotheriid Unit – Polypteriformes – western Djourab. INTRODUCTION two extant genera, i.e. Polypterus, with ten species and 15 subspecies (Poll, 1941a, b, 1942; Hanssens, Teugels Palaeontological expeditions have been conducted in & Thys van der Audenaerde, 1995; Britz, 2004; there Chad since the early 1930s. In 1994, the MPFT are 11 species if P. polli is considered as a valid spe- (Mission Paléoanthropologique Franco-Tchadienne) cies), and the monotypic genus Erpetoichthys. Nine initiated a series of field missions to the Mio-Pliocene fossil genera have been described. Two have been sites of the eastern Djurab Desert in northern Chad described on the basis of isolated skull bones and (Brunet et al., 1995, 1998; Brunet & MPFT, 2000) and, scales (Meunier & Gayet, 1996; Gayet & Meunier, since 1997, to the Miocene sites of Toros-Menalla, 1991), and six others have been described on the basis western Djurab. The Anthracoteriid Unit in Toros- of the articular head of pinnules (Gayet & Meunier, Menalla has yielded a rich vertebrate fauna (Vignaud 1996; Werner & Gayet, 1997). Finally, Serenoichthys et al., 2002), the oldest known species of hominid, has been described on the basis of scale and finlet Sahelanthropus tchadensis (Brunet et al., 2002), as counts and distribution (Dutheil, 1999). Serenoichthys well as the first nearly complete and articulated skel- is the only other fossil articulated skeleton known so eton of a fossil polypterid. This polypterid belongs to a far, but the head is missing. These fossil genera are new species that is described here. The estimated age Cretaceous in age and African, except for Dagetella of the Anthracotheriid Unit is c. 7 Myr (Vignaud et al., and Latinopollia, which are of Late Cretaceous and 2002). Early Palaeocene age, respectively, and South Ameri- Polypterids (Cladistia) are basal actinopterygian can. Post-Palaeocene polypterid history is exclusively fishes (Rosen et al., 1981). They are represented by African. Isolated scales and skull bones attributed to Polypterus sp. testify to the presence of the genus in *Corresponding author. E-mail: [email protected] African freshwaters since the Eocene. Moreover, two © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society, 2006, 146, 227–237 227 228 O. OTERO ET AL. types of African Cretaceous pinnules were tentatively bital space (as in P. bichir katangae and possibly in attributed to two fossil Polypterus species, P. dageti P. teugelsi, whereas it is more than 60% in P. ansorgii, (Gayet & Meunier, 1996) and P. sudanensis (Werner & and less than 45% in the other species) and orbit Gayet, 1997). The intrarelationships among poly- smaller than suboperculum width (as in P. bichir and pterids have yet to be clarified; indeed, the monophyly P. endlicheri only); prognathus lower jaw (as in and the intrarelationships among extant species have P. bichir and P. endlicheri only); at least 5–7 prespi- not been established. racular bones (the number of prespiracular bones is This paper provides the first description of a near possibly equal to or above seven only in P. weeksii, in complete fossil polypterid skeleton including cranium, P. bichir bichir and in P. endlicheri congicus); deep and it is the only clearly recognizable fossil record for body with approximately 23 transversal scales, 58–60 the genus Polypterus. The new species from the Late scale rows, and 14 predorsal scales (such body shape Downloaded from https://academic.oup.com/zoolinnean/article/146/2/227/2631032 by guest on 26 October 2020 Miocene of Toros-Menalla (western Djurab, Chad) is exists elsewhere only in P. ansorgii, P. bichir and assigned to the genus Polypterus based on its morpho- P. endlicheri); 15 finlets with successive pinnules of logical characteristics, which fall within the anatomi- the finlets overlapping the base of the following cal range of Recent species of the genus, from which it one (such dorsal fin mophology elsewhere only in differs by a unique combination of characters. We also P. ansorgii, P. bichir and P. endlicheri congicus); discuss the primitive or derived state of certain osteo- grooved lateral line scales with notched posterior mar- logical features observed in this species. gins (as in P. ansorgii, P. bichir and P. endlicheri only). The holotype is 590 mm in standard length. Such dimensions are also reached in P. bichir and SYSTEMATIC PALAEONTOLOGY P. endlicheri only. ACTINOPTERYGII COPE, 1887 Occurrence. Anthracotheriid Unit of Toros-Menalla, (SENSU ROSEN ET AL., 1981) Late Miocene of Chad. CLADISTIA COPE, 1871 POLYPTERIDAE GÜNTHER, 1870 MATERIAL AND METHODS POLYPTERUS ST HILAIRE, 1802 Type species. Polypterus bichir St Hilaire, 1802. Specimen TM090-001-039 was found lying on its back. It is preserved in three dimensions. The dorsal and lateral body surfaces have globally kept their living POLYPTERUS FARAOU SP. NOV. shape, whereas the ventral surface has collapsed (Fig. 1A–C). Most of the scales of the body are present Derivation of name. In Chadian Arabian, faraou and articulated with each other. All the bones of the means flattened. The name is given in reference to the right half of the skull are in natural position, whereas dorso-ventrally depressed head of the fish. some bones of the left side are slightly displaced. This Type specimen. TM090-001-039, housed in the Centre three-dimension preservation allows us to observe the National d’Appui à la Recherche (CNAR, N’Djamena, natural position of the bones and the shape of the cra- Chad). nium; it also allows us to measure the dimensions of the specimen (Fig. 2) and to compare some dimensions Diagnosis. A Polypterus species characterized by: and meristic data between the specimen and Recent depressed head (as in P. weeksii and P. endlicheri species. Jollie’s (1984) nomenclature of the bones is only), head length approximately 20% of body length followed. (less than 20% in other Polypterus species except P. ansorgii, P. bichir and P. endlicheri) and head width at the level of the preoperculum reaches 70% of its ANATOMICAL DESCRIPTION length (more than in any other Polypterus species); HEAD AND BODY SHAPE (FIGS 1–3) opening of the lateral line on the nasal 3 close to its posterior border (as in P. bichir and P. endlicheri, The standard length of the specimen is 59 cm (Fig. 2, whereas it is central in other species); large and a). We observe a natural dorso-ventral depression of rounded snout, shorter than the interorbital distance the head. The head length (from the front of the mouth (as in most Polypterus species except P. bichir and to the posterior level of the opercular bone) is equiva- P. palmas the snout of which is longer than the lent to about 20% of the standard length (Fig. 2, m). interorbital distance); superolateral position of the The width of the head, measured at the level of the orbit (as in P. ansorgii, P. weeksii, P. endlicheri and preoperculum, is around 70% of its length (Fig. 2, n). P. bichir, whereas the orbit is lateral in P. delhezi, The distance between the lateral border of the spira- P. ornatipinnis, P. senegalus, P. retropinnis and cular series at the level of the preoperculum is 45% of P. palmas); orbit length around the half the interor- the total length of the head (Fig. 2, o). The lower jaw is © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society, 2006, 146, 227–237 A NEW POLYPTERID FROM THE LATE MIOCENE OF CHAD 229 Downloaded from https://academic.oup.com/zoolinnean/article/146/2/227/2631032 by guest on 26 October 2020 Figure 1. Photograph of specimen TM90-01-39, holotype of Polypterus faraou sp. nov., in A, dorsal, B, left lateral, and C, ventral views. Figure 2. Measurements in specimen TM90-01-39, holotype of Polypterus faraou sp. nov., and ratios used in the text. prognathus. The general shape of the snout is wide (nasals 3) meet in the midline. The suboperculum is and rounded, and it is shorter than the interorbital deeper than the orbit length. The gular plates are space is wide (Fig. 2, compare c and e).
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