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Selachians and Actinopterygians from the Upper Jurassic of Tendaguru, Tanzania

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Selachians and Actinopterygians from the Upper Jurassic of Tendaguru, Tanzania Mitt. Mus. Nat.kd. Berl., Geowiss. Reihe 5 (2002) 207-230 10.11.2002 Selachians and actinopterygians from the Upper Jurassic of Tendaguru, Tanzania Gloria Arratial, Jiirgen =wet2 & Wolf-Dieter Heinrich’ With 9 figures and 2 tables Abstract The first Late Jurassic selachian and actinopterygian fishes of Tendaguru in Tanzania were collected by the German-Tenda- guru expedition in 1909-1913. They are represented mainly by occasional teeth of a neoselachian (Sphenodus) and several specimens of a neopterygian (Lepidotes). New material collected by the German-Tanzanian Tendaguru expedition in 2000 includes additional selachians recovered from clay stones at site Dwa 5a and isolated actinopterygian remains. At least three hybodonts (Hybodus sp., Hybodontidae indet., Lonchidion sp.) and a new neoselachian batoid are presented here. This assem- blage is endemic to Tendaguru. A new batoid genus and species, Enguibutis schultzei, is described, which is the oldest record of a ray from Gondwana. The actinopterygians are represented by scarce and disarticulated scales of Lepidotes, ‘pholido- phoriform’-like scales, and teleostean vertebrae. This material includes new biogeographic records for Africa. In addition, a synopsis of Jurassic fishes from Africa is presented. Key words: Fishes, Enguibutis schultzei n. gen. and n. sp., Late Jurassic, Tendaguru, Tanzania, East Africa. Zusammenfassung Die Fischfauna aus dem Oberjura von Tendaguru ist bisher nur unzureichend bekannt. Das erste Material, das von der Deut- schen Tendaguru-Expedition (1909-1913) gesammelt wurde, besteht fast nur aus einigen wenigen isolierten Zahne des Haies Sphenodus und mehreren Exemplaren des Actinopterygiers Lepidotes. Neue Funde, die wahrend der Deutsch-Tansanischen- Tendaguru-Expedition im Jahre 2000 geborgen wurden, erlauben es, mehrere Taxa zu beschreiben, die aus dem Oberjura Afrikas noch nicht bekannt waren. Anhand von Zahnen konnten mindestens drei zu den urspriinglichen Hybodontiern zah- lende Taxa (Hybodus sp., Hybodontidae indet., Lonchidion sp.) nachgewiesen sowie ein fur Tendaguru neuer Rochen (Engu- ibutis schultzei n. gen. and n. sp.) festgestellt werden. Actinopterygier sind durch wenige und disartikulierte Schuppen von Lepidotes und einer ‘pholidophoriden’dhnlichenForm sowie durch Wirbel von Teleosteern vertreten. Das ,neue Material gibt wichtige Hinweise auf die palaobiogeographische Verbreitung der spatjurassischen Fische. Es wird ein Uberblick uber die bisher bekannten jurassischen Fischfaunen Afnkas gegeben. Schliisselworter: Fische, Enguibutis schultzei n. gen. und n. sp., Oberjura, Tendaguru, Tanzania, Ostafrika. Introduction mens assigned to Lepidotus minor are reason- ably complete individuals from a monospecific The first fish remains from the Tendaguru Beds mass-accumulation (Hennig 1914a: 296). Nearly were reported by Hennig (1914a). Prior to this 90 years elapsed until the subsequent study of study, fishes were unknown from the Mesozoic fish remains from the Tendaguru Beds. Arratia of East Africa. Hennig recognized selachians & Schultze (1999) restudied the available fish (Orthacodus, now interpreted as Sphenodus), a material and referred the Lepidotus specimens pycnodontid, and semionotids described as Lepi- from Tendaguru to a new species, Lepidotes ten- dotus minor (figured as Lepidotus aff. minor), daguruensis. Recent field work of the German- Lepidotus sp., and remains of fishes identified as Tanzanian Tendaguru expedition (Heinrich et al. Genus indet. (Table 1). Most of the material re- 2001) resulted in the discovery of further fish re- ported by Hennig consists of moderately well mains. This new material, collected mainly in ex- preserved teeth and scales. However, some speci- posures of the Tingutinguti and Dwanika stream ’ Museum fur Naturkunde der Humboldt Universitat, Institut fiir Palaontologie, Invalidenstr. 43, D-10 115 Berlin, Ger- many. * Department of Earth Sciences, University of Bristol, Wills Memorial Building, Queens Road, Bristol BS8 1RJ, UK. Received April, accepted June 2002 208 Arratia, G. et al., Selachians and actinopterygians from Tendaguru GEOLOGICAL SECTION LEGEND trough crossbedding rn low-angle crossbedding 140 horizontal bedclinp ripple c rosSbedding flaw or lenticular bedding clay-rich olstone with potfhedral texture massive, clay-rich Siltstone and claystone carbonate MAP LEGEND notexposed nNiuvium aTngonta smeei &d sample location OO.,~ exlraclasts Lipogiro gravels ---- intraclasts BMiddle Saurian Bed A excavation sde e.@e concretions Tqonta schwarzr Menn~aEed rJ rnerlstream Y D I biotuibaions Bed CI= clsystone. sI = sitdone, L/I Sa~~~Bed Lower Saurian Bed - - - roadntack IS: fne-graned sandstone. 16= mediumgrained SandStonf gS = coarse-grained sandstone. 0 1 krn co = conglomerate Mitt. Mus. Nat.kd. Berl., Geowiss. Reihe 5 (2002) 209 sections, differs from the hitherto known fish The biostratigraphic position of fish-bearing fauna of the Tendaguru Beds in its greater taxo- deposits in the Tendaguru Beds can be outlined nomic diversity. Several taxa previously un- as follows: The age of the Lower Saurian Bed is known from the Late Jurassic of East Africa are tentatively considered to be Oxfordian, since the recognized. of the Nerineu Bed, from which fish remains have yet to be recovered, is Oxfordian ? to Late Kimmeridgian in age (Aberhan et al. 2002). Based Geological and paleontological setting on charophyta, the middle part of the Trigoniu smeei Bed is referrable to the early Tithonian Tendaguru Hill is located in southeastern Tanza- (Heinrich et al. 2001, Aberhan et al. 2002). A nia, about 60 km northwest of the seaport of bone bed (Jg) in the Middle Saurian Bed at the Lindi. Details of the geological structure of the Tendaguru site dy was considered to be either Tendaguru region and the stratigraphy of the fa- Kimmeridgian (Schudack 1999) or Kimmeridgian mous Tendaguru Beds, which have produced a to Tithonian (Schrank 1999) in age. Ammonites diverse dinosaur assemblage, are given by Hen- collected from the Nerineu Bed and the Trigoniu nig (1914b), Janensch (1914, 1925), Aitken smeei Bed indicate a Late Kimmeridgian to (1961), and Russell et al. (1980), with important Tithonian age of the Middle and Upper Saurian additions and refinements by Bussert (2001), Bed (Aitken 1961, Russell etal. 1980). Ammo- Heinrich et al. (2001), and Aberhan et al. (2002, nites (Aitken 1961) and palynomorphs (Heinrich this volume). et al. 2001) refer the Trigoniu schwurzi Bed to The stratigraphic sequence exposed at Tenda- the Early Cretaceous (possibly Valangian to guru Hill consists of three dinosaur-bearing stra- Hauterivian: Aberhan et al. 2002). ta, the Lower, Middle, and Upper Saurian Bed, separated from one another by three marine sandstone units, the Nerineu Bed, the Trigoniu Synopsis of Jurassic fishes from Africa smeei Bed, and the Trigoniu schwurzi Beds (Figs 1 and 2). The age of the Tendaguru Beds is While the Cretaceous fish fauna of Africa has considered to be Late Jurassic to Early Cretac- attracted much scientific attention, there are eous. The age of the fish-bearing strata will be comparably few studies on Triassic and Jurassic discussed below. fishes (for references see Murray 2000). Isolated The fish remains described here come from teeth and scales referred to Colobodus cf. muxi- the Lower and the Middle Saurian Beds, the Tri- mus, Colobodus sp., Hybodus cf. minor, and Eu- goniu smeei Bed, and the Upper Saurian Bed gnuthus (= Furo) from the Lugh Series of north- (Figs 1 and 2, Tab. 2). They were collected from ern Somalia by d’Erasmo (1931) are considered sites in the Dwanika stream section and the Tin- to be Late Triassic or Early Jurassic in age gutinguti stream section (Fig. 1, Tab. 2), and com- (MacFadyen 1933). Goodwin et al. (1999) re- plemented by material from previous excavation ported scales from the Late Triassic-Early Juras- localities of the German-Tendaguru expedition sic Adigrat Sandstones of Ethiopia west of Addis (IV, XX, dy) located in the surroundings of Ten- Ababa. Gaudant et al. (1972) reported the pre- daguru Hill. Most of the material has been sence of a small teleost from the Toarcian of Tu- found in the fine-grained sandstones in the chan- nisia, which was identified as Leptolepis. In gen- nel facies, followed by specimens recovered from eral, there are few Early Jurassic fish remains fine-grained sediments in the tidal flat facies reported from Africa. (Tab. 2). The data on the petrography and de- Saint-Seine (1955) and Saint-Seine & Casier positional environment of the fish-bearing strata (1962) described two fish faunas from the Juras- are summarized in Table 2. All the material de- sic Stanleyville Beds of the interior of what was scribed here was obtained from acetic acid resi- then the Congo. The deposits were interpreted dues of rock samples processed for recovery of as Kimmeridgian in age by Saint-Seine (1955) microvertebrates. but are now considered to be Aalenian-Batho- 4 Fig. 1. Map of the Tendaguru area in southeastern Tanzania with a synthetic section of the Tendaguru Beds and the location of the sites and the position of the fish-bearing horizons. Source: Hennig (1914a), Janensch (1914b, 1925), Heinrich et al. (2001), and data from the German Tanzanian Tendaguru expedition 2000 (after R. Bussert). Arratia, G. et al., Selachians and actinopterygians from Tendaguru nian (Colin 1994). A highly diverse fauna includ- ing coelacanths but no selachians was described from supposed fresh water deposits (Saint-Seine 1955). The marine fish fauna of the Stanleyville
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