The Tendaguru Formation, or Tendaguru Beds are a highly fossiliferous formation and Lagerstätte located in the Lindi Region of southeastern Tanzania. The formation represents the oldest sedimentary unit of the Mandawa Basin, overlying Neoproterozoic basement, separated by a long hiatus and unconformity. It reaches a total sedimentary thickness of more than . The formation ranges in age from the late Middle Jurassic to the Early Cretaceous, Oxfordian to Hauterivian stages, with the base of the formation possibly extending into the Callovian.
The Tendaguru Formation is subdivided into six members; these are from oldest to youngest the Lower Dinosaur Member, the Nerinella Member, the Middle Dinosaur Member, the Indotrigonia africana Member, the Upper Dinosaur Member, and the Rutitrigonia bornhardti-schwarzi member. The succession comprises a sequence of , , , conglomerates with minor oolite , deposited in an overall shallow marine to coastal plain environment, characterized by tidal, fluvial and lacustrine influence with a tsunami deposit occurring in the Indotrigonia africana member. The climate of the Late Jurassic and Early Cretaceous was semi-arid with seasonal rainfall and the eustatic sea level was rising in the Late Jurassic from low levels in the Middle Jurassic. Paleogeography reconstructions show the Tendaguru area was located in the subtropical southern hemisphere during the Late Jurassic.
The Tendaguru Formation is considered the richest Late Jurassic stratum in Africa. The formation has provided a wealth of fossils of different groups; early , several genus of , crocodyliformes, , fish, and flora. More than of material were shipped to Germany during excavations in the early 20th century. The faunal assemblage of the Tendaguru is similar to the Morrison Formation of the central-western United States, with an additional marine interbed fauna not present in the Morrison.
The dinosaur fauna found in the formation is similar to that of other highly fossiliferous stratigraphic units of the Late Jurassic; among others the Kimmeridge Clay and of England, the Sables de Glos, Argiles d'Octeville, Marnes de Bléville of France, the Alcobaça, Guimarota and Lourinhã Formations of Portugal, the Villar del Arzobispo Formation of Spain, the Shishugou, Kalazha and Shangshaximiao Formations in China, the Toqui Formation of Chile and Cañadón Calcáreo Formation and the Morrison Formation, with the presence of dinosaurs with similar counterparts, e.g., Brachiosaurus and Stegosaurus in the Morrison, and Giraffatitan and Kentrosaurus in the Tendaguru.Mateus, 2006, pp.223–232
Based on extended geological and paleontological observations, the "Tendaguruschichten" (Tendaguru Beds) were defined by Werner Janensch as expedition leader and Edwin Hennig in 1914 referring to a sequence of Late Jurassic to Early Cretaceous strata, exposed in the Tendaguru area, which is named after the Tendaguru Hill.Bussert et al., 2009, p.142
+ Stratigraphy of the Tendaguru FormationBussert et al., 2009, p.152 | |||||
Makonde | Early Albian Aptian | Fine to medium grained sandstones, intercalated conglomerates, siltstones and claystones | ~ | ||
Unconformity | |||||
Tendaguru | Hauterivian Valanginian | Rutitrigonia bornhardti-schwarzi | Fine to medium grained sandstones with basal conglomerate | ||
Unconformity | |||||
Tithonian | Upper Dinosaur | Ripple cross bedded fine grained sandstones and siltstones with intercalated claystone and micritic carbonates | ~ | ||
Indotrigonia africana | Calcite cemented sandstones, conglomerate beds, thin clay and silt layers with sandy limestones | ||||
Late Kimmeridgian | |||||
Middle Dinosaur | Ripple cross bedded fine grained calcareous sandstones and siltstones and massive to crudely bedded silt and claystones | ||||
Kimmeridgian Oxfordian | Nerinella | Trough cross bedded sandstone to massive sandstone | |||
Mid Oxfordian ?Callovian | Lower Dinosaur | Cross bedded fine grained sandstones and siltstones, with Interbedded clay-rich siltstones | > | ||
Hiatus | |||||
Triassic | |||||
Paleozoic | |||||
Basement | Neoproterozoic | Gneiss | |||
At time of deposition the area was undergoing a semi-arid climate with coastal influences that maintained somewhat higher moisture levels than seen inland.Noto & Grossmann, 2010, p.7 The upper parts of the formation, the Middle Dinosaur and Rutitrigonia bornhardti-schwarzi Members in particular, showed prevailing semiarid conditions with pronounced dry seasons, based on palynologic analysis.Schrank, 1999, p.181 The Tendaguru fauna was stable through the Late Jurassic.Noto & Grossmann, 2010, p.9
During the Late Jurassic and Early Cretaceous, the Gondwana paleocontinent was breaking up and the separation of the and Gondwana supercontinents resulted from the connection of the Tethys Ocean with the proto-Atlantic and the Pacific Ocean. In addition, the South Atlantic developed towards the end of the Late Jurassic with the separation of South America and Africa. Africa became increasingly isolated from most other continents by marine barriers from the Kimmeridgian into the Early Cretaceous, but retained a continental connection with South America. Global sea levels dropped significantly in the Early Jurassic and remained low through the Middle Jurassic but rose considerably towards the Late Jurassic, deepening the marine oceanic trench between continents.Arratia et al., 2002, p.227
In the Tingutinguti stream section, the Indotrigonia africana Member exhibits several up to thick, poorly sorted, conglomeratic sandstone beds. They contain , reworked and/or accumulations of thick-shelled bivalves (mainly Indotrigonia and Seebachia), and exhibit megaripple surfaces. These conglomeratic sandstone layers are interpreted as storm deposits. In the Dwanika and Bolachikombe stream sections, and in a small tributary of the Bolachikombe creek, a discrete, up to thick conglomerate in the lower portion of the Indotrigonia africana Member displays evidence of a tsunami deposit. Overall, lithofacies and the
diverse macroinvertebrate and microfossil assemblages of the Indotrigonia africana Member suggest a shallow marine environment. Based on the diverse mesoflora and the abundance of Classopollis, a nearby vegetated hinterland is postulated that was dominated by xerophytic .
Following the discovery in 1906, teams from the Museum für Naturkunde, Berlin (1907–1913), and later the British Museum (Natural History), London (1924–1931) launched a series of collecting expeditions that remain unequalled in scope and ambition. Led by the vision and influence of geologist Wilhelm von Branca, the German expeditions were particularly successful. This was largely due as the project was regarded as a matter of national ambition of the German Empire and enjoyed the benevolence of wealthy patrons. Eventually, nearly 250 tons of fossils were shipped to Berlin, representing an entirely new dinosaur fauna that remains the best understood assemblage from all of former Gondwana.Cifelli, 2003, p.608
From there, the material was transported to Fraas' institution, the Royal Natural History Collection in Stuttgart, Germany. Fraas described two species in the badly known genus "Gigantosaurus"; G. robustus and G. africanus (today Janenschia robusta and Tornieria africana, respectively.
The small-scale trough and ripple cross-bedded fine-grained sandstone at the base of the Upper Dinosaur Member is interpreted as tidal flat deposits. Unfossiliferous sandstone in the upper part
was most likely deposited in small fluvial channels in a coastal plain environment, whereas argillaceous deposits were laid down in still water bodies such as small lakes and ponds. Rare occurrences of the ostracod Cypridea and charophytes signal the influence of freshwater, whereas the sporadic occurrence of marine invertebrates suggests a depositional environment close to the sea.
Fining upward sequences of the basal part of the Rutitrigonia bornhardti-schwarzi Member are interpreted as tidal channel fills, the overlying fine-grained sandstone, silt- and claystone as tidal flat deposits. From the immediate surroundings of Tendaguru Hill, invertebrates and vertebrates are poorly known and limit the palaeoenvironmental interpretation of this member. The composition of the land-derived sporomorph assemblage suggests a terrestrial vegetation which was dominated by cheirolepidiacean conifers in association with .
Excavation history
German Tendaguru Expedition
Public discussion about provenance and restitution
In popular culture
Paleontological significance
Fossil content
Mammaliaformes
Allostaffia A. aenigmatica Quarry Ig Middle Dinosaur Three isolated molars Originally described as Staffia, later renamed Allostaffia as Staffia was preoccupied bya foraminifera.Heinrich, 2004
Assigned to Haramiyida (though possibly a gondwanathere instead).Chimento et al., 2016 Brancatherulum B. tendagurense Unspecified Upper Dinosaur Dentary without teeth Either a stem- or .Averianov & Martin, 2015, p.327 Tendagurodon T. janenschi Quarry Ig Middle Dinosaur Single tooth One of the earliest AmphilestidaeHeinrich, 1998, p.269 Tendagurutherium T. dietrichi Quarry Ig Middle Dinosaur Partial dentary with damaged last molar Either a Peramuridae or an
Squamates
Paramacellodidae? Indeterminate An osteoderm Unconfirmed
Pterosaurs
Tendaguripterus T. recki Quarry Ig Middle Dinosaur A partial mandible with teeth Barrett et al., 2008 ?Indeterminate archaeopterodactyloid Mkoawa Mtwara Humerus Costa & Kellner, 2009, p.814 Indeterminate azhdarchid Mkoawa Mtwara Indeterminate dsungaripteroid Upper Dinosaur Humerus Pterodactylus P. maximus Mkoawa Mtwara Later determined to be an indeterminate pterodactyloid P. brancai Mkoawa Mtwara TibiotarsiGalton, 1980 Later determined to be an indeterminate dsungaripteroid P. arningi Mkoawa Mtwara Later determined to be an indeterminate pterosaur Rhamphorhynchus R. tendagurensis Mkoawa Mtwara Later determined to be an indeterminate "rhamphorhynchoid"
Ornithischians
Dysalotosaurus D. lettowvorbecki Quarry Ig Middle Dinosaur "Large number of mostly disassociated cranial and postcranial elements" A Dryosauridae"Table 19.1," in Weishampel et al., 2004, p.414 Kentrosaurus K. aethiopicus Quarry Q, Ig, St, S, Ny, Li, XX, r, y, d, Ng, X, H, IX, Om, bb, Ha, XIV, II, IV, V, VIII, G, e, g, Ki Lower, Middle & Upper Dinosaur "Two composite mounted skeletons, four braincases, seven sacra, more than seventy femora, approximately 25 isolated elements, juvenile to adult" A Stegosauria"Table 16.1," in Weishampel et al., 2004, p.344
Sauropods
Australodocus A. bohetii Quarry G Upper Dinosaur Two neck vertebrae; more undescribed remains destroyed during World War II A SomphospondylanRemes, 2007 Dicraeosaurus D. hansemanni Quarry Q, m, St, dd, Sa Lower, Middle & Upper Dinosaur "Skeleton lacking skull and forelimbs, two partial skeletons, isolated vertebrae, and limb elements" A Dicraeosauridae"Table 13.1," in Weishampel et al., 2004, p.264 D. sattleri Quarry La, s, O, ab, E, M, o, Ob, bb, XIV, G, GD Middle & Upper Dinosaur "Two partial skeletons without skulls, isolated postcranial remains" Giraffatitan G. brancai Quarry Q, J, Ig, Y, St, dd, S, TL, XX, Ma, JR, Ng, Bo, To, p, t, Lw, D, N, ab, cc, X, IX, Z, T, Aa, l, E, XIV, II, G, e, Ki, No, R, F, XII, GD, XV, Sa, U, i Lower, Middle & Upper Dinosaur Brachiosaurid. The new genus Giraffatitan was erected to hold the former Brachiosaurus species, B. brancai after scientists concluded that it was distinct enough from the Brachiosaurus type species, B. altithorax, to warrant such a reclassification.Taylor, M.P., 2009, pp.787-806 Janenschia J. robusta Quarry dd, P, IX, B, G, Oa, NB Middle & Upper Dinosaur Known from hindlimb and forelimb material, left pubis and two right Ischium non-Neosauropoda Eusauropoda Tendaguria T. tanzaniensis Nambango site Upper Dinosaur "Two associated cranial dorsal vertebrae" A Turiasauria"Table 13.1," in Weishampel et al., 2004, p.261 Tornieria T. africana Quarry St, k, MD, A, e, Sa Middle & Upper Dinosaur "More than three partial skeletons, a few skull elements, and many isolated postcranial elements" Diplodocidae"Table 13.1," in Weishampel et al., 2004, p.265 Wamweracaudia W. keranjei A sequence of caudal vertebrae MamenchisauridaeMannion et al., 2019 Brachiosaurus B. brancai Mkoawa Mtwara "Five partial skeletons, more than three skulls, and isolated limb elements" B. brancai was distinct enough from the non-Tendaguru Brachiosaurus type species B. altithorax that it was moved to its own genus, Giraffatitan."Table 13.1," in Weishampel et al., 2004, p.267 B. fraasi Remains attributed to B. fraasi were later referred to B. brancai, and thus now Giraffatitan Diplodocinae indet. Indeterminate Kijenjere Upper Dinosaur Partial skull Belonging to a form that is closely related to DiplodocusRemes, 2009, p.26 Diplodocidae indet. Indeterminate Kijenjere Upper Dinosaur Caudal vertebrae and metatarsal Originally referred to as "Barosaurus"Remes, 2009, p.28 Diplodocidae indet. Indeterminate Trench XIV Upper Dinosaur Articulated pedes Possibly representing two different taxaRemes, 2009, p.30 Flagellicaudata indet. Indeterminate Upper Dinosaur Braincase Referred to Flagellicaudata indet. based on the derived characters shared with this groupRemes, 2009, p.34 "The Archbishop" Brachiosaur, distinct from Giraffatitan
Theropods
?Abelisauridae indet. Indeterminate Quarry TL Upper Dinosaur and Middle Dinosaur A left tibia, a right tibia, and a femur Possibly an indeterminate abelisaurid. Megalosauroidea indet. Indeterminate Quarry MW Upper Dinosaur left tibia and left astragalus A large indeterminate megalosauroid. Elaphrosaurus E. bambergi Quarry Ig, dd, ?RD Middle Dinosaur, ?Upper Dinosaur "Postcranial skeleton""Table 3.1," in Weishampel et al., 2004, p.48 An Elaphrosaurinae noasaurid Ostafrikasaurus O. crassiserratus Quarry Om Upper Dinosaur "Tooth" A tooth of controverisial affinitiies, had either been suggested to have been a spinosaurid or a ceratosaurid.Buffetaut, 2012, p.2 Veterupristisaurus V. milneri Quarry St Middle Dinosaur "Vertebrae" The earliest known carcharodontosaurid. Allosaurus ?A. tendagurensis Quarry TL Middle Dinosaur A tibia"Table 4.1," in Weishampel et al., 2004, p.75 Remains now considered "Tetanurae indet." Possibly a megalosauroid or carcharodontosaurid. Originally referred to Allosauridae' Ceratosaurus C. roechlingi Quarry St, MW Middle & Upper Dinosaur Caudal vertebra Tentatively referred to Ceratosauridae. Originally referred to Allosauridae. Labrosaurus L. stechowi Quarry St, MW Middle Dinosaur Teeth A Ceratosauridae, a species of Ceratosaurus. Originally referred to Allosauridae. ? Torvosaurus T. sp Quarry St, MW Upper Dinosaur Teeth Includes remains previously referred to "Megalosaurus" ingens - now known as " Torvosaurus sp".
Crocodyliformes
Bernissartia B. sp Upper & Middle Dinosaur Bussert et al., 2009, p.164
Amphibians
Fish
Invertebrates
Gastropods
Bivalves
Bussert et al., 2009, p.165 Bussert et al., 2009, p.162
Coral
Ostracods
Flora
See also
Notes and references
Notes
Bibliography
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