The Taphonomy of Dinosaurs from the Upper Jurassic of Tendaguru (Tanzania) Based on Field Sketches of the German Tendaguru Expedition (1909-1913)1

Home , Dicraeosaurus, Eberhard Fraas, Edwin Hennig, Janenschia, Werner Janensch

Mitt . Mus. Nat.kd. Berl., Geowiss. Reihe 2 (1999) 25-61 19 .10.1999

Wolf-Dieter Heinriche

With 23 figures and 2 tables

Abstract

Tendaguru is one of the most important dinosaur localities in Africa . The Tendaguru Beds have produced a diverse Late Jurassic (Kimmeridgian to Tithonian) dinosaur assemblage, including sauropods (Brachiosaurus, Barosaurus, Dicraeosaurus, Janenschia), theropods (e.g., Elaphrosaurus, Ceratosaurus, Allosaurus), and ornithischians (Kentrosaurus, Dryosaurus) . Con- trary to the well studied skeletal anatomy of the Tendaguru dinosaurs, the available taphonomic information is rather limited, and a generally accepted taphonomic model has not yet been established . Assessment of unpublished excavation sketches by the German Tendaguru expedition (1909-1913) document bone assemblages of sauropod and ornithischian dinosaurs from the Middle Saurian Bed, Upper Saurian Bed, and the Transitional Sands above the Trigonia smeei Bed, and shed some light on the taphonomy of the Tendaguru dinosaurs. Stages of disarticulation range from incomplete skeletons to solitary bones, and strongly argue for carcass decay and post-mortem transport prior to burial . The sauropod bone accumulations are dominated by adult individuals, and juveniles are rare or missing . The occurrence of bones in different superimposed dinosaur- bearing horizons indicates that skeletal remains were accumulated over a long time span during the Late Jurassic, and the majority of the bone accumulations are probably attritional. These accumulations are likely to have resulted from long-term bone imput due to normal mortality events caused by starvation, seasonal drought, disease, old age and weakness. The depositional environment of the Middle and Upper Saurian Bed was mainly limnic to brackish in origin, while the palaeoenvironment of the Transitional Sands was marginal marine.

Key words: Dinosauria, taphonomy, Late Jurassic, Tendaguru, East Africa .

Zusammenfassung

Tendaguru zählt zu den bedeutendsten Dinosaurier-Lagerstätten Afrikas. Aus den Tendaguru-Schichten sind zahlreiche Skelettreste von Sauropoden (Brachiosaurus, Barosaurus, Dicraeosaurus, Janenschia), Theropoden (z.B . Elaphrosaurus, Cera- tosaurus, Allosaurus) und Ornithischiern (Kentrosaurus, Dryosaurus) geborgen worden . Sie stammen aus der späten Jura-Zeit (Kimmeridge - Tithon) . Während der Skelettbau der Tendagurusaurier gut untersucht ist, wirft die Taphonomie des Saurier- vorkommens von Tendaguru noch immer Fragen auf. Unklar ist bislang, wie die enormen Anreicherungen von Dinosaurier- knochen in den Tendaguru-Schichten zustandekamen . Unveröffentlichte Grabungsskizzen der Deutschen Tendaguru Ex- pedition (1909-1913) erweitern unsere Kenntnisse über die Taphonomie der Tendagurusaurier. In den ausgewerteten Grabungsskizzen sind Knochenansammlungen von Sauropoden und Ornithischiern aus dem Mittleren und Oberen Saurier- mergel sowie aus den Übergangsschichten über der Trigonia smeei-Schicht dokumentiert . Die Lage und der Erhaltungs- zustand der Funde lassen auf erheblichen Zerfall der Kadaver und post-mortalen Transport von Skelettelementen vor der Einbettung schließen . Das Vorkommen von Saurierknochen in mehreren übereinanderliegenden Profilabschnitten der Tenda- guru-Schichten zeigt, daß Skelettreste während der späten Jura-Zeit über einen längeren Zeitraum hinweg akkumuliert wur- den. Die Ansammlungen von Skelettresten gehen wahrscheinlich auf normale" Sterbe-Ereignisse zurück, wie z. B. Ver- hungern, Verdursten, Kankheit, Altersschwäche und jahreszeitliche Dürre. Als Ablagerungsraum der Mittleren und Oberen Saurierschicht kommt ein küstennaher limnischer, zeitweise wohl auch brackischer Küstenstreifen in Betracht. Die knochen- führenden Übergangsschichten unter- und oberhalb der Saurierschichten sind randlich marine Ablagerungen.

Schliisselwörter Dinosaurier, Taphonomy, Oberjura, Tendaguru, Ostafrika .

1. Introduction were discovered by B. Sattler in 1907 (Hennig 1914a, Wild 1991), and E. Fraas was the first to Tendaguru in southeastern Tanzania, East Africa, excavate dinosaurs in the vicinity of Tendaguru has yielded one of the most important Late Jur- Hill in September 1907 (Wild 1991). Shortly after- assic dinosaur assemblages. The first dinosaurs wards, Fraas (1908) described and figured some

The paper is dedicated to Professor Dr. W. Janensch (1878-1969), leader of the German Tendaguru expedition (1909-1913), in appreciation of his outstanding contributions to the understanding of African dinosaurs . z Museum fiir Naturkunde, Institut fiir Pal5ontologie, Invalidenstr . 43, D-10115 Berlin, Germany. Received January 1999, accepted May 1999 26 Heinrich, W.-D., Taphonomy of Dinosaurs from Tendaguru of these bones as "Gigantosaurus" africanus downstream and buried close to the mouth of (= Barosaurus africanus) and "Gigantosaurus" rivers that flowed into the sea. Reck (1925), robustus (= Janenschia robusta). Extensive col- however, doubted catastrophic mass mortality lections of dinosaurs were made by the German and interpreted the depositional environment of Tendaguru expedition between 1909 and 1913 the Saurian Beds as saline marshes. To date (Janensch & Hennig 1909; Hennig 1912a, b; Ja- there is no taphonomic model that convincingly nensch 1914a, c), followed by field seasons of explains the enormous accumulations of dino- the British Tendaguru expedition between 1924 saur bones in the Tendaguru Beds. and 1931 (e.g., Migeod 1927, 1930, 1931 ; Parkin- The archive of the Institute of Palaeontology son 1930). The German and British expeditions of the Museum of Natural History of the Hum- yielded spectacular collections of dinosaurs and boldt University, Berlin, retains unpublished since that time Tendaguru has been known as field sketches made by the German Tendaguru one of the world's most important Upper Juras- expedition (1909-1913). These field sketches are sic dinosaur localities. extremely important because primary field re- Whereas the British dinosaur collection records of the expeditions are very scarce. The un- mained undescribed (Russell et al. 1980), the published field records reveal details of the pre- German dinosaur material from Tendaguru has servation characteristics of Tendaguru dinosaurs been well studied. Sauropods (e.g., Brachio- and enable some reassessment of the tapho- saurus, Dicraeosaurus), theropods (e.g., Elaphro- nomic interpretations suggested by previous saurus, Ceratosaurus, Allosaurus), and ornitho- workers. The main focus of this paper is to docu- pods (Kentrosaurus, Dryosaurus) were identified ment and assess these unpublished Tendaguru and described in a series of monographs. Most of field records. these works concentrated on the anatomical description of the Tendaguru dinosaurs (e.g., Hen- nig 1925; Janensch 1914b; 1929a, b; 1935, 1950a, Setting 1955, 1961a, b). By contrast, the available taphonomical accounts are rather limited (e.g., Hennig 2.1 Geological and palaeontological background 1912a, b; Janensch 1914a, c, 1929a, b) and detailed excavation plans have not been published The Tendaguru Beds that have yielded the dino- so far. saur assemblages crop out in southeastern Tan- The depositional environment and the origin zania (Hennig 1914b, 1937 ; Janensch 1914a, c; of the bone accumulations recovered from the Aitken 1956, 1961 ; Kent et al. 1971 ; Kapilima Tendaguru Beds are still under debate. Several 1984; Zils et al. 1995a, b; Fig. 1). The sedimen- taphonomic models have been suggested, among tary sequence was named after Tendaguru Hill them the interpretation of Janensch (1914a, c), (= steep hill; Dr. J Kabudi, Daressaalam, oral according to which the Tendaguru Saurian Beds communication), which is located about 60 km are marginal-marine in origin. The attritional northwest of the seaport of Lindi (Fraas 1908, and catastrophic bone accumulations and re- Janensch 1914a, c) . The following paragraph markable pre-burial taphonomic biases have briefly summarises the original description by Ja- been taken into consideration in this model. Ja- nensch (1914c) . nensch (1914a, c) argued that dinosaurs had The Tendaguru Beds represent shoreline de- been trapped and killed in the mud or 'mire- posits that were laid down near an oscillating holes' of lagoons that were temporarily exposed strandline during Late Jurassic and Early Cretac- by ebbing tides (see also Hennig 1912a, b). This eous times (Janensch 1914a, c; Russell et al. was followed by carcass flotation and decay in 1980) . There are three sedimentary cylces in the shallow water, prior to burial. Bone beds mainly Tendaguru Beds (Hennig 1914b, Janensch consisting of disarticulated skeletal elements of 1914c). The series of sediments begins with sandy Dryosaurus or Kentrosaurus were thought to re- marls of the Lower Saurian Bed (Untere erste sult from sudden, catastrophic, mass-mortality Saurierzone, Janensch 1914c; Unterer Saurier- events (e.g., Hennig 1912a, 1925; Janensch 1914a, mergel, Janensch 1961a) which measure more c) . Other workers argue that an estuarine en- than 20 m in thickness (Fig. 1). These strata pass vironment existed in what is now the Tendaguru gradually into the marine sandstones of the Ner- area (e.g., Abel 1927, Kitchin 1929, Parkinson inea Bed (Untere Sandsteinzone, Nerineenzone, 1930, Colbert 1984), and that dinosaur carcasses Janensch 1914c; Untere Zwischenschichten, Ja- were washed in by rivers. They were transported nensch 1961a) which are overlain by greenish- Mitt. Mus. Nat.kd. Berl., Geowiss. Reihe 2 (1999) 27 grey and reddish sandy marls of the Middle the Trigonia schwarzi Bed, a view that is strongly Saurian Bed (Mittlere zweite Saurierzone, Ja- supported by palaeontological (Dietrich 1933, nensch 1914c; Mittlerer Sauriermergel, Janensch Spath 1927-1933, Aitken 1956, 1961) and geolo- 1961a). Janensch (1914c) reports a thickness of gical evidence (Aitken 1956, 1961) . The inverte- about 25 m for the Nerinea Bed and some 15 m brate fossil record indicates an Upper Valangi- for the Middle Saurian Bed. The succeeding nian (Oberes Valendis) to Aptian age (Dietrich fine to coarse grained marine sandstones of 1933) or Hauterivian to Aptian age (Spath the Trigonia smeei Bed (Mittlere Sandsteinzone 1927-1933) for the Trigonia schwarzi Bed (Ait- mit Trigonia smeei, Janensch 1914c; Obere ken 1956, 1961) . The Jurassic-Cretaceous bound- Zwischenschichten, Janensch 1961a), which mea- ary is placed above the Upper Saurian Bed, sure about 20 m in thickness, separate the Mid- which is thus regarded as entirely Late Jurassic dle Saurian Bed from the Upper Saurian Bed in age (e.g., Dietrich 1933, Aitken 1956), and be- (Oberste dritte Saurierzone, Janensch 1914c; low the Trigonia schwarzi Bed, contrary to the Oberer Sauriermergel, Janensch 1961a). The opinon of previous workers who positioned this latter consists of greenish-grey and reddish sandy boundary either at the base (e.g., Hennig 1914b, marls which reach some 40 m in thickness. The Janensch 1914c), or within the Upper Saurian Upper Saurian Bed is overlain by sandy shallow Bed (e.g., Hennig 1937). The stratigraphic work marine sediments of the Trigonia schwarzi Bed in the Tendaguru region is concisely summarized (Obere Sandsteinzone mit Trigonia schwarzi, by Aitken (1961) . Janensch 1914c), the thickness of which was esti- The present report is especially concerned mated at approximately 5 m. with the Middle and Upper Saurian Beds and Sandy or oolitic sediments at the boundaries the Transitional Sands above the Trigonia smeei between the Saurian Beds and the marine sand- Bed. These units of the Tendaguru Beds are now stone units (Zwischenschichten, Nerinea Bed, generally considered to be Kimmeridgian to Trigonia smeei Bed) have been described as Tithonian in age (Aitken 1956, 1961; Russell et Transitional Beds (Obergangschichten), since al. 1980; see also Schrank 1999, Schudack et al. marine invertebrates (e.g, belemnites) are asso- 1999). Moreover, charophytes suggest a Kimmer- ciated with skeletal remains of dinosaurs (Hen- idgian age for the Middle Saurian Bed (Schu- nig 1914b, Janensch 1914c). Transitional Beds dack 1999). containing marine invertebrates and dinosaurs The majority of dinosaur remains were recov- have also been reported from the Nerinea Bed ered from the Middle and Upper Saurian Bed, (Janensch 1961a) . Sandy deposits below and occuring mainly as incomplete skeletons and soli- above the Trigonia smeei Bed have been distin- tary bones (Janensch 1914a, c; 1929a, 1961a). By guished as the Lower and Upper Transitional contrast, the Lower Saurian Bed has only Sands by Russell et al. (1980). yielded a small number of isolated sauropod The age of the Tendaguru Beds has been con- bones and theropod teeth (Janensch 1929a, troversial since the first dinosaurs were discov- 1961a). ered in the vicinity of Tendaguru Hill (e.g., Ja- The Middle and the Upper Saurian Bed show nensch 1914c; Dietrich 1925, 1933; Kitchin 1929 ; clear evidence of redeposition (Janensch 1961a: Hennig 1937; Aitken 1956, 1961) . The age of the 179) . This is consistent with the lithological char- Tendaguru Beds ranges from Late Jurassic to acters of the matrix from the Middle Saurian Early Cretaceous (e.g., Hennig 1914b; Janensch Bed from Site Jg (WJ) which was searched for 1914c; Schuchert 1918, 1934; Dietrich 1933 ; Hen- mammals (Heinrich 1998, 1999). The matrix oc- nig 1937 ; Aitken 1956, 1961; Kapilima 1984), casionally contains assemblages of oncoids, mud contrary to previous views that placed the se- clasts and lithoclasts consisting of reworked ca- quence in the Cretaceous (e.g., Fraas 1908, liche nodules (Dr. M. Aberhan, Prof. Keupp, Dr. Kitchin 1929), and there is general agreement Schudack; oral communication). The latter were that the Trigonia schwarzi Bed, above the Upper derived from palaeosoils that had developed in Saurian Bed, correlates well with the Early Cre- the Tendaguru palaeoenvironment, under a taceous (e.g., Janensch 1914c, Lange 1914, warm and seasonally dry (arid or semiarid) cli- Zwierzycki 1914, Dietrich 1933, Aitken 1961) . mate. In this connection it is interesting to note Hennig (1914b, 1937) believed that the Ten- that the known invertebrate fossil record from daguru Beds represent a nearly continuous se- the Middle and Upper Saurian Beds, which in- quence without major breaks. By contrast, Par- cludes records of freshwater gastropods (e.g., kinson (1930) suggested a disconformity below Dietrich 1914, Hennig 1914c), evidence of As- 28 Heinrich, W.-D ., Taphonomy of Dinosaurs from Tendaguru mussia (= Estheria) (Janensch 1933), and limnic Labrosaurus (?) stechowi and Ceratosaurus (?) to brackish ostracods (Schudak et al. 1999) is roechlingi (Janensch 1914b, 1929a, 1961, Hennig consistent with the sedimentological data. 1925) . Pterosaurs (Reck 1931) and a mammal Dinosaur bone assemblages, including concen- (Dietrich 1927, Heinrich 1991, 1999) were also trations of water worn limb bones of sauropods reported. were also frequently found in the Transitional The Transitional Sands below the Upper Sau- Sands above and below the Trigonia smeei Bed, rian Bed and above the Trigonia smeei Bed and some of them show evidence of reworking yielded among others Brachiosaurus brancai, (Janensch 1961a). Dinosaur remains have also Barosaurus africanus, Janenschia robusta, Di- been reported from the Transitional Sands above craeosaurus sattleri, and Kentrurosaurus aethiopi- the Nerinea Bed (Janensch 1961a). The co-occur- cus (Janensch 1961a). Pterosaurs were also de- rence of dinosaur bones and marine to brackish scribed (Reck 1931, Galton 1980) . invertebrates in these beds suggests a shallow marine to brackish depositional environment for 2.2 Sites the Transitional Sands (Janensch 1914c, 1961a). The marine depositional environment of the Dinosaurs were first discovered at a few sites in Nerinea Bed, Trigonia smeei Bed, and Trigonia the vicinity of Tendaguru Hill (Fraas 1908), and schwarzi Bed is indicated by the presence of were subsequently collected from approximately marine invertebrates, including ammonites (e.g., one hundred sites during the field seasons of the Zwierziecky 1914, Dietrich 1933, Zeiss 1975), German Tendaguru expedition between 1909 gastropods (e.g., Dietrich 1914, 1933; Hennig and 1913. Most of these sites are located close to 1914c), lamellibranchs (e.g., Lange 1914, Dietrich Tendaguru Hill (see Janensch 1914a: 45; 1925a: 1933, Aitken 1961), and brachiopods (e.g., Lange 18), except for a small number of localities in 1914), but these all require further study. More- the Mchuya and Makangaga area that are not over, with regard to the stratigraphic terminol- considered here. The former is about 40 km, the ogy, Aitken (1961) doubted that Trigonia smeei latter approximately 80 km north of the Tenda- described from the Tendaguru Beds (e.g., Lange guru region (Janensch 1914a, Fig. 1). 1914, Dietrich 1933) was identical to Trigonia The German Tendaguru expedition was smeei from the type locality of Shapoor in Cutch run by W Janensch (1909-1911), assisted by (India), contrary to Cox (1952) who accepted E. Hennig (1909-1911), H. von Staff (1911), and their synonymy. H. Reck (1912-1913) who was supported by his The field scetches of the German Tendaguru wife Mrs. Ina Reck nee von Grumbkow. The expedition cover sites in the Middle Saurian Tendaguru sites were worked intensively, and at Bed, the Upper Saurian Bed, and the Transi- important localities, the collecting period ex- tional Sands between the Trigonia smeei Bed tended through three field seasons from 1909 to and the Upper Saurian Bed that yielded a di- 1911, for example at Site S (Fig. 1) which yielded verse vertebrate fauna. As far as the land verte- the type specimen of Brachiosaurus brancai (Ja- brate assemblage of the Middle Saurian Bed is nensch 1914a, c). In 1911, about 500 Africans concerned, the dinosaurs are the most common were involved in the excavations . Together with element and include among others Brachio- their families about 900 people temporarily inha- saurus brancai, Barosaurus africanus, Dicraeo- bitated the region of Tendaguru Hill (Jaeger saurus hansemanni, Kentrurosaurus aethiopicus, 1971) . Dryosaurus lettow-vorbecki, Elaphrosaurus bam- The excavations met with outstanding success. bergi, Labrosaurus (?) stechowi, Ceratosaurus (?) African field crews supervised by the leaders of roechlingi, and Allosaurus (?) tendagurensis (Ja- the German Tendaguru expedition and native nensch 1914b, 1925b, 1929a, 1955, 1961a; Hennig foremen uncovered the bones, which were 1925). Pterosaurs (Janensch 1914b, Reck 1931, mapped, labelled, catalogued, packed, carried to Unwin & Heinrich 1999) and lizards (Broschins- the seaport of Lindi, and shipped to Germany ki 1999) and mammals (Heinrich 1998, 1999) (Janensch 1914a). Each site was indicated with also occur. lower case or capital letters, Roman numerals or The land vertebrate fauna of the Upper Sau- other symbols (e.g., Nr.) combined with Arabic rian Bed is also dominated by dinosaurs, namely numerals (Janensch 1914a) . Janensch (1914a, by Brachiosaurus brancai, Barosaurus africanus, 1925a) published maps indicating the topo- Janenschia robusta, Dicraeosaurus sattleri, Ken- graphic and stratigraphic position of the princi- trurosaurus aethiopicus, Elaphrosaurus bambergi, pal sites of the Tendaguru area, most of which Mitt . Mus. Nat.kd. Berl., Geowiss. Reihe 2 (1999) 29 are from the Upper and Middle Saurian Beds. er not more than 15 sites. Skeletal elements are Only a few sites are known from the Tansitional roughly sketched in the quarry plans, partly in Sands at the base and top of the Saurian Beds. pencil, partly with ink. The bones are mostly la- A detailed account of the diggings at Tendaguru belled with arabic numerals combined occasion- and the recovery of dinosaur bones is given by ally with the symbols of the sites (e.g., Cl, C2). Janensch (1914a). Preliminary anatomical identifications of the bones are given in the field sketches and the . Hand-written notes 2.3 Excavation records Tendaguru field catalogue by W. Janensch indicate that some of these iden- Documentation of the German Tendaguru expe- tifications were later reassessed. With a few ex- dition is largely incomplete. Records include a ceptions, the length and width of the bones are detailed field catalogue, containing lists of the not indicated in the field sketches. dinosaur bones recovered with hand-written Scale drawings of bone accumulations made notes by Werner Janensch, Edwin Hennig, and with grids were probably lost during World War Hans Reck. Sketchbooks of the German Tenda- II, and arrows indicating compass directions are guru expedition with pencil and ink drawings by often lacking. Discrepancies between the speci- Werner Janensch, Edwin Hennig and Hans men count given in the Tendaguru field catalo- Reck, documenting single bones and skeletons, gue and bones documented in the excavation and watercolours of individual skeletal elements, sketches indicate that the quarry plans often re- possibly by Ina Reck, are also available . A few present only "snapshots" of the diggings, and do topographic quarry plans are also present in the not completely reflect all details of the fossil ver- sketchbook. The collection of records also con- tebrate record encountered during the excava- tains Werner Janensch's photo album containing tions. Moreover, the field sketches lack detailed black and white photographs documenting the data with respect to the contemporary fauna and field seasons from 1909 to 1911. Most important sedimentology of the dinosaur-bearing beds ex- are photographs of uncovered sauropod skele- posed by the digging activities. tons including Dicraeosaurus hansemanni (Site Despite these gaps, there is no doubt that the m) and Brachiosaurus brancai (Sites D, S, no) . unpublished field records of the German Tenda- Three oil paintings completed by Ina Reck at guru expedition can contribute to a better under- Tendaguru Hill in 1912 were recently rediscov- standing of the taphonomy of the Tendaguru di- ered in the Institute of Palaeontology of the Mu- nosaurs. The field sketches record approximately seum of Natural History of the Humboldt-Uni- the original position of skeletal elements and versity, Berlin . These paintings give some idea of their position relative to one another. A broad the size of the quarries and field work at Tenda- spectrum of disarticulation stages can be recog- guru, possibly of Sites Jg (WJ) and St. This doc- nized, and reveal interesting details of the tapho- umentation, detailed above, forms the basis for nomic history of the Tendaguru dinosaurs that this study. have not been documented so far. It is note- Nevertheless, taphonomical analysis of the worthy that much of the skeletal material docu- Tendaguru dinosaurs are greatly hampered by mented in the field sketches is housed today in the loss of field records. There is, for instance, the collections of the Palaeontological Institute only a partial field book of the German Tenda- of the Museum of Natural History, Humboldt guru expedition and this covers the time span University, Berlin. Therefore, some of the pre- before the expedition's arrival at Tendaguru Hill served field sketches could be "updated" and, on April 16, 1909. Missing parts of the field for example, augmented with scales and ontoge- book, quarry maps and excavation plans appear netic data obtained from palaeohistological ex- to have been destroyed by fire during World aminations of sauropod limb bones (Sander War II, possibly as a result of an air raid on No- 1999, in press) . vember 12, 1943 that completely destroyed a collection room containing dinosaurs from Tenda- guru. The type material of Dryosaurus lettow- 3. The field sketches vorbecki has been missing since that time. As mentioned previously, almost 100 dino- The majority of field sketches document dinosaur-bearing sites were discovered by the Ger- saur bone assemblages from the Upper Saurian man Tendaguru expedition between 1909 and Bed. Only a few quarry maps cover bone accu- 1913 . The available field sketches, however, cov- mulations from the Middle Saurian Bed and the

30 Heinrich, W-D., Taphonomy of Dinosaurs from Tendaguru

Geographic location and geological structure of Tendaguru, Tanzania Trigonia schwarzi Bed

Upper

Saurian

Bed

_TS TS Trigonia

smeei

Bed

TS TS Middle

Saurian

Bed TS TS

Nerinea

Bed

2 3 0 500 1000 1500 m TS TS Lower E -~_ SB E w TSMB 0 Saurian -MSB ~NB + ,- + + + + ~LSB + r + + + + + + + + + + + + + + _+ + Bed + + + + + + + + + + + + + + + + -G + r + + + + + + + + + + + + + +

Fig. 1 . Map of the Tendaguru area with geological sections and the location of the sites considered in the present account. Data from Hennig (1914b) and Janensch (1914c, 1925a). Note that the marine Nerinea Bed, the two Trigonia Beds, and the Transitional Sands were not mapped in the original site plan (Janensch 1925a: 18) and the Transitional Sands were not distinguished as separate stratigraphical units by Janensch (1914c) . 1, Lower Saurian Bed; 2, Middle Saurian Bed; 3, Upper Saurian

Mitt. Mus. Nat.kd. Berl ., Geowiss. Reihe 2 (1999) 31

Table 1 Stratigraphic distribution of selected dinosaur-bearing sites in the Tendaguru area .

Stratigraphy Sites

Upper Saurian Bed C,D,N,0,P,k,ab,cc,X Transitional Sands between Trigonia smeei Bed and Upper Saurian Bed H, IX Trigonia smeei Bed Transitional Sands between Trigonia smeei Bed and Middle Saurian Bed Middle Saurian Bed S, Y,Jg(=WJ),m Transitional Sands between Middle Saurian Bed and Nerinea Bed Nerinea Bed

Transitional Sands overlaying the Trigonia smeei The assignment of bone accumulations to dis- Bed. No records are available from the Lower articulation stages such as incomplete skeletons Saurian Bed, the Transitional Sands above and or partial skeletons is based on a simplified de- below the Nerinea Bed, and the Transitional scriptive scheme that provides unambiguously Sands below the Trigonia smeei Bed. The topo- defined taphonomic categories (Table. 2). Ac- graphic and stratigraphic position of the sites de- cording to this classification, partial skeletons tailed in the present taphonomic account is indi- only consist of one discrete skeletal unit (e.g., cated in Figure 1 and Table 1 . tail or hind limb), in contrast to skeletons that As in other vertebrates, the Tendaguru dino- always comprise more than one of those units saurs began their taphonomic history as car- (e.g., tail and hind limb; fore limb, shoulder gir- casses with complete skeletons, followed by a dle and neck) . broad spectrum of disarticulation stages and scat- The skeletal elements are annotated with sym- tering of individual skeletal elements. Analysis of bols used by the German Tendaguru field crew. the field sketches and data given by Janensch With the exception of Fig. 11, triangles indicate (1929a) reveal that four main preservational ca- specimens documented either by arabic numerals tegories occur in the Tendaguru Beds (Table 2) : or by hand-written notes in the Tendaguru field (1) skeletons, (2) partial skeletons, (3) bone ac- catalogue and descriptions in the literature. The cumulations dominated by disarticulated indivi- precise position and orientation of these bones is dual elements, and (4) solitary bones. These uncertain. The assessment of quantitative data main preservational types can be further subdi- and the azimuth orientation of bones must be vided by using additional taphonomical criteria treated with caution because of the incomplete- such as the degree of disarticulation and loss of ness of the excavation records. Reference speci- bones (see Table 2). mens that were used to calculate the scale for

Table 2 Disarticulation stages of a dinosaur skeleton (simplified). For explanation see text.

Disarticulation stages Designation Skeletal elements

A Complete skeleton articulated, no loss of bones B Complete skeleton partly articulated, partly disarticulated; no loss of bones C Complete skeleton disarticulated, no loss of bones D Incomplete skeleton articulated, loss of bones E Incomplete skeleton partly articulated, partly disarticulated ; loss of bones F Incomplete skeleton disarticulated, loss of bones G Complete partial skeleton articulated, no loss of bones H Complete partial skeleton partly articulated, partly disarticulated ; no loss of bones I Complete partial skeleton disarticulated, no loss of bones K Incomplete partial skeleton articulated, loss of bones L Incomplete partial skeleton partly articulated, partly disarticulated ; loss of bones M Incomplete partial skeleton disarticulated, loss of bones N Bone field partly associated, partly disassociated O Bone field disassociated P Solitary Bone isolated

A Continued legende Fig. 1 Bed; TSB, Trigonia schwarzi Bed; USB, Upper Saurian Bed; TSMB, Trigonia smeei Bed ; MSB, Middle Saurian Bed ; NB, Nerinea Bed ; LSB, Lower Saurian Bed ; G, gneiss; TS, Transitional Sands. A - Sites in the Middle Saurian Bed, " - Sites in the Upper Saurian Bed, 0 - Sites in the Transitional Sands between the Trigonia smeei Bed and the base of the Upper Bed 32 Heinrich, W-D., Taphonomy of Dinosaurs from Tendaguru several quarry plans are quoted in the captions Further abbreviations: GTE - German Ten- of the textfigures. daguru expedition, MNHB - Museum für Na- The definition of taphonomic variables used turkunde der Humboldt-Universität zu Berlin, in the description is as follows (Behrensmeyer IPHB - Institut für Paldontologie am Museum 1991, Lyman 1996): MNI - minimum number of Mr Naturkunde der Humboldt-Universität zu identified individuals; NISP - number of speci- Berlin. mens documented in the field sketches; size of accumulation - area 2 over which skeletal (m ) 3.1 Sites in the Upper Saurian Bed remains are recovered; density - number of specimens documented in the field sketches 3.1.1 Tendaguru Site C per m2. The following anatomical abbreviations are Location : Site C is situated approximately used in the quarry maps: as - astragalus, ca - 0.7 km south of Tendaguru Hill. cc - cervical rib, cd - caudal vertebra, c - rib, Stratigraphy : Upper Saurian Bed dorsal rib, co - coracoid, cr. - skull, cv - cervical vertebra, do - dorsal vertebra, fe - femur, Ta x o n: Barosaurus africanus fi - fibula, he - haemapophysis, hu - humerus, Figure : 2 il - ilium, is - ischium, me - metacarpal, mt - Reference : Janensch (1914a, 1929a, 1961a), metatarsal, pb - pubis, pm - phalange (manus), GTE field catalogue: 3 pp - phalange (pes), ra - radius, sa - sacrum, Assemblage data sc - scapula, st - sternal plate, tb - tibia, ul - NISP: 37 ulna, v - vertebra, ~) - specimen of known ontogenetic age. The right side is indicated by d MNI: 1 and the left by s. Capital or lower case letters Number of taxa : 1 with arabic numerals are catalogue numbers of Designation : Single-individual, single-taxon the German Tendaguru expedition. assemblage.

Tendaguru, Site C C20 Upper Saurian Bed C21 cv C19 C18 Barosaurus africanus

C22 ~C23 tb fi

.wIONN~~i

'W10111111 .0, 11111-111

~IIOII IIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIII~h

vertebral column and ribs

shoulder girdle pelvic girdle

IIIII~IIIIIIIIIIIIII hind limb indet . approx . 1 m

Fig . 2. Incomplete skeleton of Barosaurus africanus from Tendaguru Site C. A few bones are still articulated, but most are disarticulated . The length of the string of cervical vertebrae C18-21 (2.7 m) was used to calculate the scale for the quarry plan. Redrawn after a field sketch from W. Janensch

Mitt. Mus. Nat.kd . Berl ., Geowiss. Reihe 2 (1999) 33

Quarry data (Janensch, GTE field catalogue: 3) . The asso- Size of accumulation : approxmately24 .00m2 ciated tibia (C22) and fibula (C23) were found just below a string of articulated cervical verteb- Density : approximately 1 .54 rae (C18-21) which measure 2,7 m in length. A Modification data string of articulated caudal vertebrae (C4 and 5) Disarticulation stage : E is also present. These articulated specimens sug- Measurements : The length of the caudal ver- gest that some skeletal elements were still con- tebrae was measured as 180 mm (C 2), 200 mm, nected by soft tissue (ligaments or muscles) prior (C3), and 200 mm (C5) ; that of the femur (C15) to burial. Several caudal vertebrae were disarti- at 1320 mm (Janensch, GTE sketch book). The culated, but apparently still close to their proper length of the left tibia (C13) was measured at anatomical order (Janensch, GTE field catalo- 890 mm (Janensch 1961a). gue: 3), indicating minimal transport from the position in which the carcass finally came to rest. Ontogenetic data : Unknown Extensive damage occured to the ilium and one Comments : The digging at Tendaguru Site C of the femora prior or subsequent to burial. commenced in April 1909 (Janensch 1914a: 43) . Most of the long bones and the string of cervical Cervical and caudal vertebrae, an ilium, a femur, vertebrae show a similar (? west to east) orienta- tibia, fibula, and an astragalus assigned to an in- tion. dividual of Barosaurus africanus were recovered (Janensch 1929a: 5). Note the dominance of elements of the posterior body region. Most of the 3.1.2. Tendaguru Site D skeletal elements were disarticulated, but still closely associated (e.g., tibia and astragalus) . The Lo c a t i o n: Site D is located about 1.0 km imperfect femur (C15), the tibia (C13), and the north-north-east of Tendaguru Hill. astragalus (C14) belong to one single hind limb Stratigraphy : Upper Saurian Bed

Fig . 3. Incomplete skeleton of Brachiosaurus brancai uncovered from the Upper Saurian Bed at Tendaguru Site D. Photo- graph: W. Janensch

34 Heinrich, W.-D., Taphonomy of Dinosaurs from Tendaguru

Tendaguru, Site D Upper Saurian Bed Brachiosaurus brancai D33 cv D34 1111111111101 ra D32 fe 111111111IIIIII-iniivnnyn IIIIIIIIIIIIIIIIIIIillllllll y1111 _D35 mc II 1IIIP~~ tb

sa mc D43 D1-31 sc

D41

D38 tb D36 D42 fe sc

vertebral column and ribs

shoulder girdle fore limb 1 m 11111111111111111111 hind limb approx .

Fig. 4. Incomplete skeleton of Brachiosaurus brancai from Tendaguru Site D. The majority of the skeletal elements are disarticulated, but closely associated . Most caudal vertebrae were found in natural order. Some limb bones rest on top of one another. The measured length of the humerus D 39 (1600 mm) was used to calculate the scale for the quarry map. The orientation of the map is not known. Redrawn after a field sketch from W. Janensch

Taxon : Brachiosaurus brancai catalogue: 4) . The dig yielded a skeleton consist- a poorly Figures : 3, 4 ing of a series of 29 caudal vertebrae, preserved sacrum, pelvic bones (ilium, pubis), Fig. 16, 1929a; Reference : Janensch (1914a, dorsal ribs, two anterior limbs without feet, two 4-5 1950a), GTE field catalogue: scapulae, femora, tibiae, and fibulae (Janensch Assemblage data 1914a, 1929a: 7 ; 1950a; GTE field catalogue: NISP: 50 4-5). Only some of these are documented in the MNI: 2 field sketch. The dimensions of the limb bones indicate a medium-sized individual of Brachio- of taxa : 1 Number saurus brancai (Janensch 1961a: 230, tab. 18) . Designation : Multi-individual but single-taxon The series of caudal vertebrae resting next to the assemblage badly preserved sacrum was found almost in nat- Quarry data ural articulation (Janensch 1950a). The association of the remaining disarticulated elements is Size of accumulation : approximately evident. The scapula (D42) was apparently 22.00 m2 found lying upon the head of humerus D39, and Density : approximately 2.27 several other limb bones were found one on top Modification data of the other (e.g., tibia D38 and femur D36; ra- and femur D32). Plate-like elements Disarticulation stage : F dius D34 such as the shoulder blades seem to be oriented limb bones Measurements : The length of the in a nearly horizontal plane, and the scapula : 230, were measured as follows (Janensch 1961a D43 and ulna D41 are broken. Most of the elon- tab. 18): humerus (1339): 1600 mm, femur (1336) : gated limb bones are oriented nearly parallel, 1550 cm, tibia (D?) : 900 mm, and fibula (D?) : and the two scapulae are positioned transverse 940 mm. to the prevailing compass direction of the heavy Ontogenetic data : Unknown limb bones. Comments : The excavations at Site D com- Most bones documented appear to derive menced on June 21, 1909 (Janensch, GTE field from a single individual preserved as an incom- Mitt. Mus. Nat.kd . Berl ., Geowiss. Reihe 2 (1999) 35 plete skeleton. Originally, the leg bones D34 and Quarry data D35 were tentatively identified as fibula (?) and Size of accumulation : Unknown radius (?) respectively (Janensch, GTE sketch Density : Unknown book). Later, Janensch (GTE field catalogue: 4, hand-written note) identified these bones as fibu- Modification data la and tibia. If properly identified, the existence Disarticulation stage : F of three tibiae indicates the presence of at least Measurements : No measurements are avail- two individuals of Brachiosaurus brancai. Unfor- able. tunately, a reassessment of the limb bones from Ontogenetic data : Unknown Site D is not possible, since the location of the collection is not known. Comments : The excavation at Site N commenced in September 1909 (Janensch, GTE field catalogue : 9). The dig produced a disarticulated 3.1.3 Tendaguru Site N incomplete skeleton, with a scapula, complete and fragmentary humerus, radius, and ribs. At a Loc a t i o n : Site N is located about 0.9 km east little distance to these bones there were two of Tendaguru Hill. ischia, a dorsal vertebra, a caudal vertebra, ribs, Stratigraphy : Upper Saurian Bed and a large fragmentary bone (N9) tentatively Tax on : Brachiosaurus brancai identified as a fragmentary scapula (Janensch, GTE field catalogue: 19). The bone accumula- Figure : 5 tion at Site N is dominated by elements of the Reference : Janensch (1914a: 44), GTE field anterior appendicular skeleton. In total, 21 skele- catalogue: 19 tal elements assigned to Brachiosaurus brancai Assemblage data were collected from site N (Janensch, GTE field NISP: 21 catalogue: 19). The bones are attributable to one individual of Brachiosaurus brancai. Except for a MNI: 1 few skeletal elements, most of the long bones Number of taxa : 1 are oriented nearly parallel, possibly reflecting Designation : Single-individual, single-taxon wave action or current action. Some of them assemblage such as the humerus and ischia are broken.

Tendaguru, Site N Upper Saurian Bed Brachiosaurus brancai N9 SO

N12 cd

N10 cv?

vertebral column and ribs shoulder girdle pelvic girdle N7 is fore limb CNP N8 indet. do

Fig. 5 . Incomplete skeleton of Brachiosaurus brancai from Tendaguru Site N. All skeletal elements are completely disarticulated . Scale and orientation of the map are not known. Redrawn after a field sketch from W Janensch

36 Heinrich, W.-D.. Tiplionomy of Dinosaurs from TLndaguru

3.1.4 Tendaguru Site 0 M c a S u r e in e n t s : The length of tllc right limb bones were measured as follows (Jancnsch 1961 Location : Site O is located about (I.S km a) : humerus (03) : 610 mm, ulna : 410 min, femur north of Tendaguru Hill . (02) : 9ti0 mn1, tibia (04) : 580 mm, fibula (OS): Stratigi- aphy : tJppcr Saurian Bed 620 mm. Ta x o i1 : Dlci-acostim-its Saidei-i Ontogenetic data : Janensch (1929x: 10) believed that the skeletal elements ori`inatcd from Figure : 6 (colour figure on p. 36) a subadult individual of Dicr-aeosniu-its snulcri, Reference : Janensch (1914x, 1929x, 1961a), howcvei, hislological examinations of' a right fe- GTE field catalogue : 20 mur (02) and a right 11umeru5 (03) Su0`11est that Assemblage data both limb bones belong to in adult individual of NISP: 17 this species (Sander 1999, in press). MNI : I Comments : Digging by the German Tcncla- guru expedition's crew at Site O commenced at Number of taxa : 1 the beginning of October 1909 (Janensch, GTE Designation : Single-individual, single-taxon field catalogue: 47). Janensch (1914x: 44, 1929x: assemblage 10) reported a scapula, pelvic bones, cents of Quarry data caudal vertebrae, and an imprecisely determined Sire of accumulation : approximately number of fore and hind limb elements, all as- 7.00 i11 2 signed to Dicrncosaurtrs siadcri. All bones are disarticulated, but closely associated . There are Density : approximately 2 .43 two groups of long hones oriented with their Modification data long axes in two perpendicular directions. The Disarticulation stage : F boric accumulation at Site O is dominated by

Tendaguru, Site O Upper Saurian Bed Dicraeosaurus sattleri

02 0+ fe.d

08 sc .d

015 mt 013 011 ca . ca 014 ca vertebral column shoulder girdle pelvic girdle fore limb hind limb approx . 1 m indet.

Fig. 6. Incomplete skeleton of Dicrucosuurus eullleri from lcndaguru Sitc 0. The spccimcn is completely disarliculated . Note tlic dominance of large liml, hones. The measured lem0th of the humerus 03 (610 inm) was used to calculate tlic scale for the quarry mali . The orientation ol tlic oral) is not known . Redrawn after a field sketch from W. Jancnsch

Milt . Mus. NatId. Berl- Geowiss. Reihe 2 (1999) )7

heavy linlh hones (11un1erus, ulna, femora, tibiae, NLinlhcr of taxa : 1 fibula) . This may indicate that most smaller D e s i g n a t i o n : Multi-i11dividttal but sin,le-tax- hones had been almost completely removed on assenlblage from the carcass by ctu - rents prior to burial. Ouarrv data Size of accumulation : appoxilllatCly 3.1.5 Tendaguru Site P 24 .(1(1 n1 - Location : Site P is located at Nterego, ap- D c n s i t v : approximately 2.74 proxinuttely 1 .2 km north-north-east of Tenda- Modification data ouru Hill . Disarticulation stage : H (find I), 1< (find i o S t r a t r a p h y : Upper Saurian Bed III, IV) and L (find II) Ta x o n : Jruw»schia mbusia Measurements : The length of the left limb Figure: 7 (colour figure on p. 37) hones were measured as follows: 1lttmerus (Pty) : Reference : Janensch (1914x, 1922, 1929x, 890 111111 . radius (Pl l ): 620 mn1, ulna (P12) : 1961x), Bonaparte et al. (in press), GTE field 670 mn1, tibia (P5): 850 111, ilstragaltts (P6) : catalogue: 21-22 210 mm. Assemblage data Ontogenetic data : Unknown NISP: 65 Comments : Dioging at Site P commenced in MN1 : 2 October 19(19 (Janensch, GTE field catalogue:

Tendaguru, Site P Upper Saurian Bed Janenschia robusta

vertebral column pelvic girdle fore limb hind limb approx . 1 m indet .

Fig. 7. Partial skeletons of hmen .schia rohnsra from Tendaguru Site P The hone asscmhlawc consists nmink, of two articukiled left fore limbs and two articulated left hind limbs. Pelvic elements rest neat to the sacrum, and some vcrtchrac are scattered over the llltat- I-N flour. The measured length of the 1111n1ct1IS Pi; (890 nun) wns used to calculate the scale for the yuKtrrN mat) . Tlic oricnlation of the mat) is not known. Redrassn after a field sketch lioin W. Janensch

38 Heinrich, W-D., Taphonomy of Dinosaurs from Tendaguru

21-22) Four partial skeletons of Janenschia 3.1.6 Tendaguru Site k robusta were uncovered, along with additional scattered postcranial bones (GTE field catalo- Location : Site k is situated about 0.7 km gue; Janensch 1922, 1929a; Bonaparte et al., in south of Tendaguru Hill. press). According to Janensch's unpublished ex- Stratigraphy : Upper Saurian Bed cavation sketches and hand-written notes (GTE Tax on : Barosaurus africanus field catalogue: 21), partial skeleton I consists of Fi g u r e : 8 (colour figure on p. 39) a reasonably complete left hind limb, with femur, tibia, fibula, and the foot, including astragalus, Reference : Janensch (1914a, 1929a, 1961a), metacarpals, and phalanges. The foot bones were GTE field catalogue still close to their natural anatomical position, Assemblage data while femur, tibia and fibula were slightly disarti- NISP: 50 culated. Close to this bone accumulation lay par- MNI: 1 tial skeleton IV, represented by an incomplete left fore limb, with radius, ulna, carpal, metacar- Number of taxa : 1 pals, and phalanges. Nearly all these bones were Designation : Single-individual, single-taxon found in their proper anatomical order. Partial assemblage skeleton II consists of an incomplete hind limb, Quarry data including tibia, fibula, and slightly disarticulated Size of accumulation : approximately and scattered foot bones (astragalus, metatarsals, 19.00 m2 phalanges). Partial skeleton III includes the radius, ulna, and the manus of an incomplete left Density : approximately 2.62 fore limb. In addition to the four partial skele- Modification data tons, dinosaur bones found at site P include, Disarticulation stage : E among others, a left humerus, a sacrum, and sev- Measurements : The length of skeletal ele- eral disarticulated vertebrae scattered over the ments is as follows (Janensch 1961a) : left scapula quarry floor. Janensch (unpublished sketchbook) (k34) : 1340 mm, right humerus (k37): 970 mm, also noted ribs in his excavation map but no lo- left ulna (k38) : 740 mm, left ischium (k44): cation data are given. The association of these 880 mm, left tibia (k41): 860 mm. bones with the four partial skeletons remains far from clear. The long axes of the two anterior Ontogenetic data : Unknown lower legs (partial skeletons III, IV), the femur Comments : Skeleton k was found close to the of partial skeleton I, and the single humerus remains of Barosaurus africanus from Sites A were similarly oriented. The two posterior lower and B (Janensch 1914a, 1929a). The collecting legs (finds I, II) are deposited with their long site was discovered in 1910 (Janensch 1914a). In axes perpendicular to the preferred orientation specimen k, all the main body regions (e.g., of partial skeletons III and IV skull, axial skeleton, appendicular skeleton) are The low degree of disarticulation, especially represented. Janensch (1929a: 6) reported a of the feet, suggests that soft tissue must have braincase, maxilla, cervical and dorsal vertebrae, remained attached to the limb bones when they anterior caudal vertebrae, ribs, scapula, coracoid, were deposited. The taphonomic evidence also humerus, ulna, ischium, femur, tibia, and fibula, suggests that the partial skeletons were buried all assigned to Barosaurus africanus. There are close to the point where the original corpses two clusters of skeletal elements, one of which came to rest. In addition, several limb bones comprises mostly bones of the anterior body re- show signs of strong weathering due to pro- gion (e.g., braincase, cervical vertebrae, scapula, cesses subsequent to burial (Janensch 1914a: 44; coracoid, humerus), whereas the other contains 1961a). bones from the posterior body region (e.g., Taphonomic and anatomical analyses show ischium, femur, tibia, fibula, caudal vertebrae) . that the four partial skeletons belong to a single Only a few skeletal elements are articulated: two species, and represent two individuals of Ja- strings of dorsal vertebrae (k12 and k13, k14 and nenschia robusta (Janensch 1922, 1929a: 7; GTE k15) and two articulated caudal vertebrae (k 20, field catalogue: 21-22). Tendaguru Site P is the k21) have been found (Janensch GTE field cata- only known site, so far, that has provided spe- logue: 48). The remaining skeletal elements were cific information on the fore and hind limbs of disarticulated but still closely associated. Most of Janenschia robusta. the cervical vertebrae and caudal vertebrae were

Mitt . Mus. Nm .kcl . Beil.. Geowiss. Rcilic 2 ( 1999) ;9

apparently found in their natural anatomical or- 3.1.7 Tendaguru Site ah der (Janensch, GTE field catalogue : 48), and haemapophyses were associated with the centra Location : Site ab is located approximately of the caudal vertebrae. 1 .2 km nortllnorthcast of Tendagtlru Hill. The excavation plan (Fig. S) shows that the Strati`l'ra1'11y : tJpper Saurian Bed limb hones, ischia, scapula, and most of dorsal Ta x a : Brrrosata -us a/i-icamis, Dicracosaunts ,sat- rips are embedded with their long axes trending tlcri, Brachiosaurus hralwai '?, Sauropoda indet . approximately northwest to southeast. The heads Figure : 9 (colour figure on p. 40) of all the dorsal ribs are directed southwards. The fibula is water-worn (Janensch, GTE field Reference : Janensch (1961a), GTE field cata- catalogue: 49) . logue : 40-41 The arrangement of the skeletal elements ap- Assemhlaac data pears to have been modified by water currents. NISY: 37 However, the presence of articulated and disarti- MNI : 5 culated vertebrae found in almost natural anatomical order, the orientation of the dorsal ribs, NLtt1lhCr of taxa : (4) and the presence of a bralncase indicate that the Designation : Multi-individual, n1ttlti-taxon carcass was embedded and decayed close to the asscmhlaoc place where it came to rest. Quarry clata

Tendaguru, Site k Upper Saurian Bed Barosaurus africanus

skull

vertebral column and ribs shoulder girdle pelvic girdle fore limb hind limb

CJ k2

k1 cr

approx . 1 m

Pig. S. Incomplete skeleton of Rewo.wun -u.s rr/iieluno from Tend,iIIuru Sitc k, in an advanced staoe of disarticuLitirm, with hrain- cnsc and cervical vertebrae resting next to the anterior appendicular MW trunk element, . The eoudal vertebrae arc close lc, the pelvic and hind limb dements. Most skeletal elements are dISPINICUIatCCI, but still closely associalrd . The measured length of the scalnila k?4 (1 3,4(1 mm) was used to calculate the scale lür tltc quarry map. Redrawn after a field sketcli from W. .lanensch

40 Heinrich . W.-D., haphonomy of Dinosaurs from Iendaouru

Size of accumulation : approximately entl_v randomly, across the excavation floor. Sev- 10.00111 , eral bones are imperfect and show distinct signs and mixing of skeletal D e n s i t y : approximately 3.61 of breakage. Scattering elements from different taxa and individuals su1- Modification data gests that there Was mach transport of hones Disarticulation stage : O prior to burial. Some of the heavy limb bones M e a s u r e m c n t s : Dicraeosam-rrs saideri, lengt11 arc oriented northwest to southeast, others are of three left humcri: 620 mm (ahl), 620 mm embedded with long axes trending approxi- (ah2), SRO1nin (abl0); Barczsarrrus all-icunus, mately west to cast. Tlie presence of three left length of a left tibia (ab4) : 650 mnl. hUtllerI (abl, ab2, abIO) reveals that the diplodo- is represented by at 0ntoQenetic data : Histological data ob- cid Dicracosourus .scrttlcri individuals. Two tibiae (ab4, ah5) be- tained from two left humcri (ab2, abIO) indicate least three . apparently, to one individual of Barosaurtrs that these limb hones are froth two adult indivi- long -icanus (Jancnsch, GTE field catalogue: 4()), duals of Dicraeosaunrs ,scrttlcri (Sander 1999, in all femora arc tentatively assigned to press) . and two large Brachiosarrrus hrarwai (Jancnsch, GTE field cat- arc no excavation data giv- C o m m e n t s : There alogue : 4()). A huge uncatalogued limb bone that 1929x, 1961 a), except for en by ,Jancnsch (1914x, was not collected (Jancnsch, field sketch, hand- . The dig commenced at the GTE field catalogue Written note) Was tentatively Identified as a October, 1909. Remains of at the beginning of humerus of Brachiosaurits branwai. least five individual sauropods were recovered from the Upper Saurian Bed at Site ab. All bones are disarticulated and scattered, appar-

Tendaguru, Site ab Upper Saurian Bed

Barosaurus africanus Brachiosaurus brancai?

L Dicraeosaurus sattleri Sauropoda indet.

approx . 1 m

Fig. d. Bone field from TCI1Chlgurlr Site ab . The accumulation consists nr

Mitt . Mus. Nat.kd. Berl ., Gcowiss. Rcihc 2 (1999)

3.1 .8 Tendaguru Site cc Comments : Tapllonomic data for Site cc are poor. Remains found here include cervical ver- Location : Site cc is situated approximately tebrae, a dorsal vertebra, several dorsal ribs, a 2.9 km eastnortheast from Zenc!aguru Hill . scapula. a 11tt111CruS, and hone fragments (,la- Stratigraphy : Upper Saurian Bed nensch, GTE field cataluouc: 142-1 ) . All elements are clisarticttlatecl . but still associated. A pre- Ta x oil : Brachiusaurus brancai ferred orientation of the hones is not discern- Figure : 1() (colour fiL0ure Oil p. 41) able. R e fe r e n c e : GTE-field catalogue: ! 42 The documented skeletal elements are mainly Assemblage clata from the anterior body region (neck, anterior ap- penclicular skeleton) of a carcass that underwent NISP: 15 maceration and decay prior to hurial. MNI : 1 Number of taxa : I 3.1.9 Tendaguru Site X Designation : Single-individual, single-taxon assemblage Location : Site X is located about 1 .t) kn1 Quarry data south-south-east from 'Tendagttru Hill. Size of accumulation : approximately S t r a t i ,o r a p h y : LJpper Saurian Bed 24.(1(1 n1~ Ta x o n : Kciarosam"its acthiopicus, Sauropocla in- Density : approximately 1 .11 clet. Fi 0 urc : Il Modification data Reference : Hennig (1925), GTE field catalo- Measurements : Length of a right hutnerus gue : 113-114 (cc2 ) : 1080 111111 . Assemblage clata DlsartICttlation stage : F NISP: 115 Ontogenetic data : According to Sander 11999, in press), hurnerus cc2 belongs to an adult MNI : (7) individual of Brachinsaur-it.s brancai. Number of taxi : (2)

Tendaguru, Site cc Upper Saurian Bed ce2 L Brachiosaurus brancai 11u .d

vertebral column and ribs shoulder girdle fore limb indet.

cc14 do

approx. 1 m

Fig. 10 . Incomplete skeleton of Brachin.saurus hrcuwai from Site cc in an advanced state of disarticulatiun. All hones are diSM'tiCUlatCd. The measured length of the Itumertü CC" (108(1 nun) was used to c~dculalc the sctde Ior the quarry map. The orientation of the map is not known. Redrawn after a field sketch from W. Jancnsch

42 Heinrich, W-D., Taphonomy of Dinosaurs from Tendaguru

De s i g na t i o n : Multi-individual, multi-taxon 113). The quarry map (Fig. 11) is based on a assemblage field plan preserved in the GTE sketch book list of skeletal elements assigned Quarry data and a detailed to Kentrosaurus (Hennig 1925: 228-229). Bones Size of accumulation : approximately 30.00 m2 of Kentrosaurus aethiopicus are the most com- Density : approximately 3.83 mon skeletal elements of dinosaurs recovered Saurian Bed at Site X, constitut- Modification data from the Upper ing more than 95% of the total dinosaur finds. Measurements : See Hennig (1925) The documented bone cluster is dominated by Disarticulation stage : N (O?) isolated hand and foot bones which formed a pavement on the quarry floor. Scattering and mix- Ontogenetic data : Unknown ing of the skeletal elements is evident. Judging Comments : Digging at Site X commenced in from the data by Hennig (1925: 228-229), the August, 1909 (Janensch, GTE field catalogue: skeletal remains belong to at least 6 individuals

Tendaguru, Site X Upper Saurian Bed N Site plan and disposition of bones in the central part of the excavation

sac

Kentrosaurus aethiopicus + + radiale" intermedium ulnare metacarpal phalange (manus) A metatarsal + ++ " phalange (pes) ?+ X140 + findet . (manus or pes) do ' cervical vertebra + caudal vertebra vertebra ++ fe femur hu humerus 0 sac sacrum tb tibia " D? .+ Fig . 11 . Bone field in the cen- Sauropoda indet . tral part of Tendaguru Site X. The accumulation is dominated 0 caudal vertebra _L 0 by manus and pes elements of do dorsal vertebra 40 Kentrosaurus aethiopicus . Scat- tering is evident . Because of sac 0 space limitation the catalogue numbers could only be shown ?A A for some bones. Redrawn after approx . 1 m *A a field sketch from W Ja- 0 nensch or E. Hennig

Nlilt. Mus. Nat .kd. Berl ., (icowiss . Reilic 2 (I`)`)`I)

of Kcml-oscnlnls acthinlficus . Moreover, isolated 3.2 Sites in the Transitional Sands above the Tri- sauropod vertebrae are also present. gonia snneei !lied The dominance of scattered and mixed hand and foot hones confirms the impression that 3.2 .1 Tendaguru Site H there was substantial transport and sorting of bones prior to burial . However. the origin of the Location : Site H is located 1 .3 km northeast nnass accumulation of kentrosaurian bones is dif- of Tendagurul Hill ficult to explain owing to the lack of precise S t r a t i g r a p h y : Transitional Sands between the data. It is possible that carcasses of mired and 7i-i,golliu stncei Bed and illc base of the Upper drowned kentrosaurs floated or were washed Saurian Bed away, whereas the hands and feet rennainccl in Ta x a : Ba1'os(llll"11.~ 11111CY11111 S, the mud, which was subsequently reworked by Figure : 12 (colour figure on p. 4 3)) currents or waves to form a pavement of bones. This interpretation would agree well with the ta- Reference~Janensch: (1929,1), GTE field cata- phonollnic model proposed by Janensch (1914c). logue: 1 1 However, it is also possible that the hand and Assemblage data foot bones found at Site X were winnowed away N I SP: 24 frcmn the decaying carcasses of Kcim-osaurlls MNI : 1 acihiopicus and thus resulted from current sorting. Whatever happened, the place of hurial is Number of taxa : 1 apparently not the place of death . Designation : Multi-individual but Sill" 1C-taXOll asselllblage Ouarry data Size of accunnulation : approximately 14 .00 In -

Tendaguru, Site H Transitional Sands between Trigonia smeei- Bed and Upper Saurian Bed

hind limb

indet . approx . 1 m

Fig. 12 . IncompIcle skeleton of Karo.sauru .s africanus Irom Tendamuru Sitc H . The acetunulatiun is dominated hV limb homes Mid pelvic elements. Some limb hones rest on lop of one another . The orientation of the map is not known . Redrawn after a field sketch from W. Janensch 44 Heinrich, W.-D., Taphonomy of Dinosaurs from Tendaguru

Density : approximately 1.71 such as the right femur (H1) and the right tibia much of the mate- Modification data (H2), are still associated, but rial recovered is imperfect (e.g. scapula H13, Disarticulation stage : F pubes H10). The dominance of heavy long Measurements : Right radius (H6) : 700 mm; bones and the remarkable scarcity of small ske- left ulna (H7): 750 mm; right femur (H1) : letal elements suggests that there was much 1200 mm; left femur (H4): 1290 m; right tibia post-mortem transportation of bones from the (H2) : 880 mm; left fibula carcasses prior to burial. Because of the size dif- (H3): 920 mm; right fibula (H5): 960 mm. ferences between the two femora found (H1: 1200 mm; H4: 1290 mm), it is concluded that the Ontogenetic data : Unknown. bone accumulation from Site H contains skeletal Comments : The dig at Site H commenced in remains of at least two individuals of Brachio- August, 1909 (GTE field catalogue: 11), and saurus brancai. produced disarticulated and scattered bones of In addition to these skeletal remains, Site H Barosaurus africanus, including ribs, scapula, has yielded non-sauropod bones not documented ulna, radius, ischium, pubis, femur, tibia, and fi- in the field sketch, among them a coelurosaurian bula (Janensch 1929a: 6). Some of these bones, tibia and a metatarsal of Kentrosaurus aethiopi-

Tendaguru, Site IX Site plan and disposition of bones Transitional Sands between in the excavation Trigonia smeei- Bed Kentrosaurus aethiopicus and Upper Saurian Bed Barosaurus africanus 1111111111111111 Brachiosaurus brancai Janenschia robusta indet.

IX01

Fig. 13 . Bone field from Tendaguru Site TX . The accumulation consists mainly of disarticulated limb bones from sauropods scattered over the quarry floor. Redrawn after a field sketch from W. Janensch

Mitt . Mus. Nat.kd . Berl., Geowiss. Reihe 2 (1999) 45

cus (GTE field catalogue: 11). Their precise pro- Assemblage data venance is uncertain. Other discoveries from Site NISP: Unknown H include marine invertebrates (e.g., belemnites) MNI: Unknown indicating a marginal marine palaeoenvironment for the dinosaur-bearing Transitional Sands Number of taxa : (4) above the Trigonia smeei Bed, as formerly sug- De si g n a t i o n: Multi-individual, multi-taxon gested by Janensch (1914c) . assemblage Quarry data 3.2.2 Tendaguru Site IX Size of accumulation : The precise size of the excavation floor is not known. The exposed Location : Site IX is situated about 1.4 km quarry floors a-f measure, in total, roughly northeast of Tendaguru Hill. 13.0 m by 2.5 m (s. Fig. 13) S t r ati g r aph y : Transitional Sands between the Density : Unknown Trigonia smeei Bed and the base of the Upper Modification data Saurian Bed Disarticulation stage : O (N?) Ta x a : Barosaurus africanus, Brachiosaurus brancai, Janenschia robusta, Kentrosaurus aethio- Measurements : Brachiosaurus brancai, picus length of a right femur (IXal), 1180 mm; length of left femur (IX1) : 880 mm; Barosaurus africa- Figure : 13 nus, length of left humerus (IX94) : 640 mm; Re fe r e n c e : Janensch (1914c), GTE field cata- length of right humeri: 660 mm (IXkl1), 730 mm logue: 108-112. (IXvl), 380 mm (IXx9).

As~"~"v e

Fig. 14. Incomplete skeleton of Dryosaurus lettow-vorbecki from Tendaguru Site Jg (WJ) . Original field sketch from 1. Reck or H. Reck

46 Heinrich, W-D., Taphonomy of Dinosaurs from Tendaguru

O n t o g e n e t ic data : According to Sander langes, ribs) are under-represented or completely (1999, in press), humerus IX94 is from a juvenile missing (e.g., skull elements, lower jaws) . individual of Barosaurus africanus, while femur Some bones are well preserved, others in poor IX1 is from an adult individual of Brachiosaurus condition. The epiphyses of several limb bones are brancai. considerably worn due to rolling and abrasion by Comments : Excavations at Site IX com- the embedding sands (Janensch 1914a, c), and menced on August 17, 1909. About 150 indivi- there is no doubt that the skeletal remains were dual bones of dinosaurs were collected from Site current modified. Moreover, the scattering and the IX which was located between the Trigonia state of preservation of the skeletal elements sug- smeei-Bed and the Upper Saurian Bed (GTE gest a long period of time during which the bones field catalogue : 108), although only some of were transported by waves and/or currents from these were mapped (Fig. 13) . Partial skeletons the carcasses to the place of permanent burial. were not found, and the majority of bones were Only a few catalogued limb bones have been disarticulated and scattered over the excavation identified taxonomically. Most of them appar- area, though some of them may have still been ently belong to sauropod dinosaurs. A minimum associated. Janensch (1914c: 44) reported that of four individuals of Barosaurus africanus are most of the bones were embedded horizontally indicated by four right humeri . Skeletal remains and concentrated in a distinct bone-bearing of Brachiosaurus brancai, Janenschia robusta, layer. He also noted the dominance of bones and Kentrurosaurus aethiopicus are apparently from the appendicular skeleton (Janensch 1914c: less common than those of Barosaurus africanus, 44) . A closer study of the 130 identified bones but precise specimen counts are not available . from Site IX reveals that 85 (= 65,4%) are leg Belemnites have frequently been found at Site IX elements, including humeri, radii, ulnae, femora, (GTE field catalogue : 108) . tibiae, and fibulae (GTE field catalogue : 108-112) . Other elements (e.g., vertebrae, pha-

Fig. 15. Tendaguru Site S, the type locality of Brachiosaurus brancai . Photograph: W. Janensch

Mitt . Mus. Nat.kd. Berl ., Gcowiss. Reihe 2 (1999) 47

3.3 Sites in the Middle Saurian Bed DisartiCttlatIOn Stage : '? (complete or incomplete skeleton) 3.3.1 Tendaguru Site Jg (= WJ) Ontooenetic data : Unknown L o c a t i o n : Tendaguru Site J O (= WJ) is located o approximately 2.3 km north-north-west of Ten- Comments : Excavations at Site J (WJ ) com- daguru Hill . menced in 191() and continued through the years 1911 and 1912 (Branca 1914 ; Janensch 1914c; S t r a t i o r h y : a p Middle Saurian Bed GTE field cataloouc : 73). Quarry Jo (WJ) is one Tax on : Diyosenn-us lcttotv-vorhecki of the richest dinosaur-bearing sites in the Sur- Figure : 14 roundings of Tendaguru Hill . It produced thou- R c f e r c n c c : GTE sketchbook sands of hones of the small ornithopod dinosaur Assemblage data Dl-yusnurus blow-ivrrheeki (Janensch 195(lc, 1955, 196Ib). Jttdgino from matrix kept in the NISP: Unknown Berlin Tendaluru collection, isolated vertebrae MNI : 1 and limp hones of Drvosaiwus lcftow-volhecki Number of taxa : 1 are the most common elements found in the Designation: Single-individual, single-taxon Middle Saurian Bed at Site Jg (WJ). Skull frag- assemblage ments and teeth are fairly rare. Site Jg (WJ) has Quarry data not only produced isolated bones but also partial and fairly complete skeletons (Flo. 14). Strings of Size of accumulation : Unknown Vertebrae with closely associated chevrons also Density : Unknown occur. The dryosaurs from Site Jg (WJ) are Modification data thought to have died close to the places where McaSttrcmcnts : Not available they were buried (Janensch 1914a, c) .

Tendaguru, Site S Middle Saurian Bed Brachiosaurus brancai

N vertebral column and ribs

shoulder girdle

pelvic girdle

fore limb

hind limb S43 cv indet.

S35-39 mc

cd S14 approx . Ilm

Fig. I(,. Incomplete skeletons of Brachiosurrs (run-11i from Tendaouru Site S. Anatomical identifications of skeleton SII arc underlined . Elements of skeleton SI could not he indentificd with certainty. The mc~isured length of the hun1CIuS S9 (213 .0011) was used 10 cnICUIAC the scale for the quarry mal, . Redrawn niter a field sketch from W. Jancnsch

I Icinrich . W.-D. . Taphonomy ol I)inoseurs from Icndaguru

I A drawing of in in-situ skeleton of Drvo- S t r a t , , r a p h _Y : Middle Saurian Bed srn1r.u.c l(gtonv-vorbccki, preserved in the GTE Ta x o n : Brachiosaurus brcuwai sketchbook shows, a fairly complete skeleton re- 15,Fi g e I 'C s : 16 (colour figure on p. 47) and 18 covered in 1912 (Fig. 14) . An articulated skull Janensch (1914a, c; 1929a, 1950a, h : seems to he present, and most of the vertebral Reference: field catalogue: 25-2t; co1L1n111 appears to be preserved in natural ar- 1961a), GTE ticulation, as do the dorsal ribs. The caudal ver- Assemblage data tebral column is partly preserved . Frotn the fore NISI': Not precisely known the proximal section of a right limbs, nothing but MNI : 2 hunicrLIS 01' ulna was documented. The left fe- taxi : 1 lnur is spread slightly away from the trunk, and Number of the proximal sections of both femora are appar- Designation : Multi-individual, Sing lC - 1aXOI1 ently closely associated with the pelvic region . assenlbla-e The lower Icgs were either not found or not Quarry data Unfortunately, cloctunented in the field sketch . Size of accurn elation : the location of the documented skeleton of Density : '? Drvosaürits lclfoiv-vorbccki is not known . Modification data : E (skeleton S11), L 3.3.2 Tendaguru Site S Dlsarticulatlon stage (skeleton S 1) Location : Site S is located approximately Measurements : Skeleton S11, length of se- 1 .0 km SOLIthsouthwest of Tendaguru Hill lected elements of the appc:nclICLIlar skeleton

Tendaguru, Site Y Middle Saurian Bed Brachiosaurus brancai N

Y1 cr is skull vertebral column and ribs shoulder girdle fore limb Y27 Ca hind limb

Y25 as .d approx . 1 m

Fig. 17 . Brochinsaurus brancai from Tendaguru Site Y. The incomplete skeleton consists of disarticulatcd elements from the calculate the scale for Uic quarn anterior 11,11. 1 of the holy. The measured length of the scapula Y8 (154 .11 cm) was used to map. Redrawn after a field sketch from W. Janensch

Mitt. Mus. Nat.kd. Berl., Geowiss. Reihe 2 (1999) 49

(Janensch 1961a): left sternal plate : 1100 mm; Brachiosaurus brancai, is a partial skeleton, com- right coracoid: ca. 840 mm; right humerus: prising the skull, six cervical vertebrae and (?) 2130 mm; right radius: 1240 mm; right Ulna: dorsals (Janensch 1929a: 8) . Specimen SII which 1300 mm; me I dext. : 595 mm; me II dext: is larger than SI, is an incomplete skeleton of 635 mm; me III dext: 595 mm; me IV dext. : Brachiosaurus brancai (Janensch 1914a, c; 1929a; 570 mm; me V dext.: 490 mm; right femur: ca. 1950a, b; 1961a), including skull bones, a nearly 1960 mm, right fibula: 1190 mm. complete presacral vertebral column (11 cervical and 11 dorsal vertebrae), cervical and dorsal O n t o g e n e tic data : Judging from solitary ribs, most of the anterior appendicular skeleton limb bones that are at least one tenth larger (left scapula, coracoids, sternals, right anterior than those of skeleton SII, it could be concluded limb, left humerus, ulna, and radius), parts of the that the SII specimen was not yet fully grown pelvic girdle (e.g. both pubes bones), and incom- (Janensch 1938) . pletely preserved hind limbs (e.g., imperfect right Comments : The dig commenced on October femur, right tibia and fibula, and foot bones). 11, 1909 and Tendaguru Site S was worked by The sacrum and the caudal series were already the German Tendaguru until 1911 (Janensch missing when the specimen was discovered, most 1914a) . Site S is the type locality for Brachio- likely due to erosion (Janensch 1914c). saurus brancai and remains of two individuals Janensch (1914a, c) reported that the humerus were found. Skeleton SI, the type specimen of and a tibia of the SII individual were found in

Fig. 18. Brachiosaurus brancai from Tendaguru Site S. The articulated series of vertebrae from skeleton S11 includes anterior dorsals and cervical vertebrae. Original field sketch from W. Janensch . The skeletal elements in the figure are tentatively identified as follows: S73 - left scapula; S76 - dorsal vertebra 2; S77 - dorsal vertebra 1; S78 - cervical vertebra 13; S82-S87 (? S80, ? S88) - cervical vertebra 12; S89-S91 - cervical ribs; S92-S93 - dorsal ribs; S96-S97 (? 94) - cervical vertebra 11

50 Heinrich, W.-D ., Taphonomy of Dinosaurs from Tendaguru

an upright position, suggesting that the animal S t r a t i g r a p hy : Middle Saurian Bed mired in soft mud. The proximal section of this Taxon : Brachiosaurus brancai humerus shows distinct traces of abrasion, indicating that the skeleton was affected by moving Fi g u r e : 17 (colour figure on p. 48) water prior to burial (Janensch 1914c) . The Re fe r e n c e : Janensch (1929a), GTE field cata- bones of the right fore limb were found in natur- logue: 37 al order, but not articulated, and several bones (e.g., coracoids, right humerus, radius ulna) were Assemblage data embedded in a horizontal position (Fig. 16). The NISP: 23 distal part of the left anterior limb was absent MNI: 1 upon discovery, possibly due to post-depositional erosion (Janensch 1914a, c) . Number of taxa : 1 (2) Sections of the reasonably complete presacral Designation : Single-individual, single-taxon vertebral column of specimen SII are roughly assemblage documented in two separate field sketches, partly in pencil, partly in ink. One of these two Quarry data field sketches is reproduced in the present ac- Size of accumulation : approximately count (Fig. 18) . According to Janensch (1950a: 21 .00 m2 33), the anterior section of the vertebral column, including presacral vertebrae 3 to 15, were found Density : approximately 1.1 in natural order. The three anteriormost presa- Modification data cral vertebrae (3-5) are well preserved and do Measurements : length of a left scapula: not show signs of abrasion, while the remaining 1540 cm; length of a right astragalus: 160 mm presacral vertebrae (6-15) are articulated and (Janensch 1961a). display close, bone-to-bone contact, except for presacral vertebra 8 which was slightly dislo- Disarticulation stage : F cated. The dorsal parts of presacral vertebrae 9 Ontogenetic data : The remains of Brachio- to 15 are missing, possibly due to attrition by the saurus brancai recovered from Site Y are from a fine sands in which the remains were embedded medium-sized individual (Janensch 1950a: 33). (Janensch 1950a). The string of presacral vertebrae (3 to 15) was recovered from a massive limy Comments : The excavations at Site Y com- sandstone bed, unike presacral vertebrae 17 to menced in 1910 (GTE field catalogue: 37) . The 24 which were found together with other bones bone accumulation from Site Y represents an in- from skeleton SII in marls below the limy sand- complete skeleton, dominated by bones from the stone bed (Janensch 1950a: 33) . The majority of anterior body half. From the posterior skeleton these posterior presacral vertebrae were disarti- only a fibula was found. Site Y has yielded a culated, except for vertebrae 23 and 24 which monospecific bone accumulation assigned to Bra- were found in bone to bone contact (Janensch chiosaurus brancai. All bones documented are 1950a: 33). disarticulated. Janensch (1929a : 8) noted a brain- The sediments exposed at the place of burial case, eight cervicals, ribs, scapula, coracoid, hu- show that low-energy dinosaur-bearing marls merus, and a dorsal vertebra. The skeleton is ap- passed into higher-energy sandy deposits. The parently current-modified, with a scapula resting shape of most bones recovered from Site S has next to a humerus and dorsal ribs. The cervical not been substantially changed by taphonomic or vertebrae are approximately in anatomical order diagenetic processes. The taphonomic evidence (Janensch 1929a: 8), and the braincase lies close obtained from skeleton SII suggests carcass ma- to them. The majority of the long bones were ceration and decay of a mired individual of Bra- embedded with their long axes oriented west to chiosaurus brancai prior to burial (Janensch east (e.g, right scapula, left humerus, dorsal ribs). 1914c). Only a few elongated bones (e.g., left scapula, left fibula) are approximately oriented north to south. Some bones were embedded on top of 3.3.3 Tendaguru Site Y one another (e.g., right scapula, left humerus). The taxonomic allocation of a small elongate Location : Site Y is located about 3.1 km caudal vertebra that was found south of the left north of Tendaguru Hill. scapula is far from clear.

Mitt . Mus. Nat.kd. Berl., Geowiss. Reihe 2 (1999) 51

3.3.4 Tendaguru Site m mur (m5) : 1220 mm; left tibia (m1): 760 mm; left fibula (m3) : 800 mm, left astragalus: 165 mm Lo c a t i o n : This site was situated close to Kin- Disarticulation stage : E dope, about 3.2 km north of Tendaguru Hill . Stratigraphy : Middle Saurian Bed Ontogenetic data : Unknown Tax on : Dicraeosaurus hansemanni Comments : One of the most important dis- Figure : 19 coveries of the 1910 field season was the type specimen of Dicraeosaurus hansemanni, recov- Reference : Janensch (1914a; 1929a, b, 1936, ered from the Middle Saurian Bed at Site m 1961a), GTE field catalogue near Kindope (Janensch 1914a). Unfortunately, a Assemblage data quarry plan is not available. The more important NISP: Unknown taphonomic data published by Janensch (1929a, MNI: 1 b) are briefly summarized below. The skeleton is incomplete, but remarkably Number o f t a x a : One-individual assemblage well preserved. Much was found in natural ar- Designation : Single individual assemblage, ticulation. The skull, proatlas, and axis have not single-taxon assemblage been not recorded. A series of vertebrae ranging Quarry data from cervical 2 to caudal 19 lay in natural order. Size of accumulation : Unknown Only the anterior end of the tail was recovered; the posterior tail section beyond caudal 19 was Density : Unknown lost prior to discovery. The preserved part of the Modification data tail is slightly recurved (Janensch 1929b: 41). Measurements : The length of the preserved Ilium, pubis, parts of the ischium, and the femur pelvic elements and limb bones is as follows of the right body side were discovered in situ. (Janensch 1961a) : right ilium: 840 mm; right The left femur was found close to the sacrum. pubis: 795 mm; right ischium: 750 mm; right fe- The sternal plates, coracoids, scapulae, and ante-

Fig. 19. Type specimen of Dicraeosaurus hansemanni from Tendaguru Site m. Photograph: W. Janensch 52 Heinrich, W-D., Taphonomy of Dinosaurs from Tendaguru rior limbs have not been recorded. Except for disassociation. They belong to a sequence of dis- the femora, the hind limbs are represented by articulation stages recognized in the Tendaguru the articulated left lower leg (tibia, fibula, and Beds (Tab. 2), including (1) incomplete skeletons astragalus), found just above the posterior sec- with bones partially articulated (e.g., Dicraeo- tion of the neck. Feet are not recorded. The saurus hansemanni, Site m; Brachiosaurus bran- well-preserved right dorsal ribs are still articu- cai, Site S), (2) complete or incomplete partial lated with the dorsals. The left dorsal ribs are skeletons (e.g. Janenschia robusta, Site P), (3) incompletely preserved and scattered, but close assemblages of predominantly disarticulated to the skeleton. Some vertebrae suffered consid- skeletal elements concentrated in bone fields erable damage: nearly all the left diapophyses (e.g. Barosaurus africanus, Site C; Kentrosaurus are missing in the section ranging from dorsal 13 aethiopicus, Site X), and (4) solitary bones. Vir- to caudal 1, and of the posterior dorsals, the left tually complete dinosaur skeletons have not parapophyses, the left neurapophyses and the been found in the Tendaguru Beds so far (Hen- centra are water worn. Three caudals were nig 1912b). Partial skeletons show a similar found at some distance from the skeleton. Their broad spectrum of disarticulation stages as in- association with the preserved caudal series of complete skeletons (Table 2), but both may re- skeleton m is uncertain. flect different taphonomic histories. A partial According to Janensch (1929b: 41), the car- skeleton, for example, may not only result from cass floated quite a long distance in quiet water gradual "in-situ" carcass decay at the site of before it came to rest on its right side. After- death or final deposition but also from early wards, the anterior section of the neck was post-mortem truncation and pre-burial transport flexed ventrally, rotated and became separated of carcass parts from the site of death or from from the posterior cervicals . Subsequently, this the original location of carcass deposition. section came to rest in an oblique position adja- Mono- and multispecific assemblages of dino- cent to the carcass prior to burial. Afterwards, saurian skeletal elements have been recognized. exposed parts of the skeleton, including the left Monospecific assemblages, for instance, are dorsal and sacral diapophyses, the left rib cage, documented for Barosaurus africanus (e.g., Site the left pelvic girdle, and perhaps the left hind C), Brachiosaurus brancai (e.g., Site S), Dicraeo- limb were abraded, truncated and dispersed saurus hansemanni (Site m), and Dryosaurus let- close to the skeleton. The skull, most of the ap- tow-vorbecki (Site Jg = WJ). These assemblages pendicular skeleton (e.g., sternal plates, anterior are usually represented by incomplete or partial limbs, right lower leg, both feet, and ? left hind skeletons, and they are mainly known from the limb) and the posterior section of the tail were Middle Saurian Bed (e.g., Sites m and S) and the probably lost during post-mortem transport from Upper Saurian Bed (Sites C, D, N) . Multispecific the place of death to the site of burial. assemblages have been documented, for instance, from Site ab (Barosaurus africanus, Bra- chiosaurus brancai, Dicraeosaurus sattleri), and 4. Discussion: taphonomic patterns in Tendaguru Sites IX and X (Barosaurus africanus, Brachio- dinosaurs saurus brancai, Janenschia robusta, Kentrosaurus aethiopicus) . Multispecific assemblages are typi- 4.1 Occurrence of bones cal of the Transitional Sands above the Trigonia smeei Bed (Site IX), but have also been docu- Most field sketches preserved in the achives of mented in the Upper Saurian Bed (e.g., Sites ab, the IPHB document bone accumulations for the X). sauropod dinosaurs Brachiosaurus, Barosaurus, Calculations of the MIN'S reveals that single- Dicraeosaurus, and Janenschia . Quarry plans individual assemblages (e.g., Barosaurus africa- documenting bone assemblages for ornithischians nus, Sites C; Brachiosaurus brancai, Sites D, M; (Dryosaurus, Kentrosaurus) are rare. Nothing Dicraeosaurus sattleri, Site O) and multi-indivi- can be said about the original position of dual assemblages (e.g., Brachiosaurus brancai, skeletal remains of theropods (e.g., Elaphro- Site S; Janenschia robusta, Site P) both occur. saurus) due to the lack of field sketches for The Middle and Upper Saurian Beds have these taxa. yielded both single-individual assemblages (e.g., The documented skeletal remains recovered Dicraeosaurus hansemanni, Middle Saurian Bed, from quarries in the vicinity of Tendaguru Hill Site m; Barosaurus africanus, Upper Saurian represent a broad spectum of disarticulation and Bed, Site k) and multi-individual assemblages Mitt . Mus. Nat.kd. Berl ., Geowiss. Reihe 2 (1999) 53

(Brachiosaurus brancai, Middle Saurian Bed, lettow-vorbecki are extraordinarily abundant at Site S; Janenschia robusta, Upper Saurian Bed, Site Jg (WJ) . Here, the matrix is dominated by Site P). The documented bone assemblages from vertebrae and limb bones, some partly articu- the Transitional Sands above the Trigonia smeei lated, and some partly disarticulated. Isolated Bed are multi-individual in nature (e.g. Baro- bones are also very common. By contrast, skull saurus africanus, Site H; Barosaurus africanus, bones and teeth are very rare, as are small hand Brachiosaurus brancai, Janenschia robusta, Ken- and foot bones. It should be mentioned that trosaurus aethiopicus, Site IX) . Single-individual bones from Dryosaurus lettow-vorbecki (Site Jg) assemblages are more frequently documented and sauropod caudal vertebrae (Site dd) en- than multi-individual assemblages. crusted with calcite have repeatedly been found. The documented multi-individual assemblages These calcite rinds, measuring 2 mm to 5 mm in consisting of incomplete or partial skeletons of thickness, are infillings of post burial shrinking sauropods are always monospecific. Examples in- cracks. Janensch (1925b: 7) also reported calcar- clude the skeletons SI and SII of Brachiosaurus eous rinds that covered bones of Elaphrosaurus brancai from the Middle Saurian Bed Site S, and bambergi recovered from Tendaguru Site dd. the partial skeletons I-IV of Janenschia robusta Skull material from sauropods is extremely from Upper Saurian Bed Site P In addition, Ja- rare, although the Middle Saurian Bed at Site t nensch (1929a: 9) reported two individuals of Di- has yielded a disarticulated complete skull of craeosaurus hansemanni from the Middle Saur- Brachiosaurus brancai with 56 teeth in its jaws, ian Bed at Site dd. and Tendaguru site dd, for instance, produced The Middle Saurian Bed has yielded the best braincases of Dicraeosaurus hansemanni and preserved skeletons from Tendaguru, some of Barosaurus africanus (Janensch 1935). which, notably those of Brachiosaurus brancai Janensch (1914c) reported water worn bones (Site S) and Dicraeosaurus hansemanni (Site m), of Brachiosaurus brancai (skeleton SII) from are on display in the Museum of Natural History Site S as well from Dicraeosaurus hansemanni of Humboldt University, Berlin. An unpublished from Site m. The material from the Middle Sau- field sketch of Site Jg (WJ) indicates that reason- rian Bed is evidently current modified. Most ske- ably complete skeletons of ornithischians (Dryo- letal elements were apparently carried away saurus lettow-vorbecki) were also found. More- from carcasses, skeletons or partial skeletons, over, the Middle Saurian Bed has produced possibly by currents or wave action. enormous concentrations of disarticulated skele- Bone accumulations from the Transitional tal remains of smaller ornithischians such as Sands above the Trigonia smeei Bed are domi- Kentrosaurus aethiopicus from Sites St, He and nated by disarticulated sauropod dinosaur limb X (Janensch 1914c, Hennig 1915, 1916, 1925) bones. Much of the material is in poor condition, and Dryosaurus lettow-vorbecki from Site Jg confirming Janensch's (1914a, c) interpretation (Janensch 1914c, 1955, 1961b; Galton 1981) . that there was much destruction, transportation, Reported recoveries of theropod dinosaurs in- and sorting of bones. Abraded epiphyses are clude several disarticulated and scattered skele- thought to be evidence of water action (Janensch tal elements from Tendaguru Site dd that are ap- 1914a, c; 1961a). Most of the bones rest horizon- parently derived from a single skeleton of tally next to one another, possibly due to wave Elaphrosaurus bambergi (Janensch 1925b: 7; Gal- action or wave-induced currents (Janensch ton 1982). Included in this collection, but undo- 1914a, c) . There is no doubt, that this material cumented in field sketches, are 17 presacral ver- underwent sorting through a broad spectrum of tebrae, 18 caudal vertebrae, a chevron bone, two post-mortem processes prior to burial. imperfect ribs, both scapulae and coracoidea, Russell et al. (1980: 172) reported the domi- right humerus, two metacarpals, sacrum, left nance of Barosaurus africanus in the Transi- pubis, two ilia and ischia, left femur, tibia, fibula, tionals Sands above and below the Trigonia astragalus, and two phalanges (Janensch 1925b: smeei Bed and noted "a significantly higher 7, 1929c: 280). In addition, several localities of number of elements from the left side" of the the Middle Saurian Bed, such as the collecting barosaur body. From Site Ki, Transitional Sands sites St and H (Janensch 1920, 1925b), produced below the Trigonia smeei Bed, Hennig (1925: isolated limb bones and teeth. 247) mentioned a complete but somewhat dis- Samples from the Middle Saurian Bed housed articulated left lower leg of Kentrosaurus aethio- in the Berlin Tendaguru collection confirm the picus, including the tibia, fibula, and bones of impression that skeletal remains of Dryosaurus the pes. 54 Heinrich, W.-D ., Taphonomy of Dinosaurs from Tendaguru

Most quarry maps from the Upper Saurian 4.2 Bone representation Bed primarily document disarticulated skeletal materials from sauropods (e.g., Brachiosaurus The great majority of sauropod bone acculuma- brancai, Sites D, N, cc; Dicraeosaurus sattleri, tions either documented in the field sketches or Site O) . Incomplete skeletons (e.g., Barosaurus listed by Janensch (1929a, GTE field catalogue) africanus, Site k) and partial skeletons (e.g., are incomplete skeletons or partial skeletons. Janenschia robusta, Site P) also occur, with The field sketches examined in this account and both partly articulated, and partly disarticulated data given by Janensch (1929a) show that some bones. As mentioned above, most of the bone skeletal elements are more frequently recorded accumulations appear to represent single-indivi- than others. Skull remains for instance, are extre- dual assemblages, although, multi-individual as- mely rare, whereas appendicular skeletal ele- semblages have also been recognized. Janensch ments usually dominate the bone assemblages (1922, 1961a) reported an articulated manus examined. This raises the question, to what ex- skeleton of Janenschia robusta from Site Nr. 5 tent certain skeletal elements better survived the and Brachiosaurus brancai from Site R, as well taphonomical filters than others. This question is as two articulated pes skeletons of Barosaurus difficult to answer, however, because no first- africanus from Sites XIII and 28, suggesting hand data are available . According to the re- that sauropod dinosaurs became mired in soft cords of sites, not all bones found have been mud (Janensch 1914c: 249). Fore- and hind- documented in field sketches. In addition, poorly limb skeletons (e.g. Janenschia robusta from preserved bones were not collected at some sites Site P) and strings of vertebrae (e.g. Baro- (Janensch, GTE field catalogue). In order to saurus africanus from Site C) and accumula- minimize the affects caused by collector prefer- tions of disarticulated bones (e.g. Barosaurus ences, individual bone representation diagrams africanus, Dicraeosaurus sattleri, and ? Brachio- were replaced in the present account by a body- saurus brancai from Site ab) show that the car- part representation diagram. This diagram casses underwent considerable pre-burial decay, (Fig. 20) reflects only "summarily" the presence as formerly suggested by Janensch (1914c, of body parts recognized in skeletons, but does 1961a) and Hennig (1925). not consider the abundance of individual skeletal The Upper Saurian Bed produced mass accu- elements within the distinguished body parts. mulations of skeletal elements of the or- The analysis of incomplete skeletons and par- nithischian Kentrosaurus aethiopicus (Hennig tial skeletons of Tendaguru sauropods from 1925). A bone field uncovered at Site X, for in- about 30 sites, partly documented in the field stance, consists mainly of isolated and jumbled sketches, partly listed by Janensch (1929a) led to elements from the manus and pes (Janensch the following conclusions (Fig. 20) : (1) As ex- 1914c, Hennig 1925) reflecting a secondary con- pected, the paired elements of the appendicular centration of dinosaurian material (Fig. 10). Hen- skeleton dominate the fossil record of Tendaguru nig (1915 : 244; 1925: 247) reported an incom- sauropods. (2) In Tendaguru diplodocids (Baro- plete skeleton of Kentrosaurus aethiopicus from saurus africanus, Dicraeosaurus hansemanni and Site bb which is close to the top of the Upper Dicraeosaurus sattleri) that have longer hind Saurian Bed. The skeleton includes a left scapula limbs than fore limbs, the rear limbs and pelvic and coracoid, imperfect left pubis, fragmentary elements are dominant. (3) In Tendaguru bra- ischium, both femora, a caudal vertebra, and chiosaurids (Brachiosaurus) that have longer fragments of ribs and vertebrae (Hennig 1925: fore limbs than hind limbs, the anterior appendi- 247). Field sketches are not available, the stage cular skeleton with its huge fore limbs and of disarticulation is unknown. shoulder girdle dominate the fossil record. Isolated bones and teeth indicate the presence Body-part representation is strongly influ- of theropod dinosaurs in the Upper Saurian Bed, enced by the degree of carcass decay that was among them Ceratosaurus, Allosaurus and reached before burial. Therefore, the tapho- Elaphrosaurus (Janensch 1925b, 1929c). Bones nomic pattern found is not only an "overprint- are very rare, and teeth are more frequently pre- ing" of distinctive skeletal traits of the sauropod served. According to Janensch (1925b: 61), a left skeletons. It also suggests a considerable amount quadratum, left fibula, and three imperfect cau- of pre-burial sorting of dinosaur bones in the dal vertebrae recovered from Site Mw are as- Tendaguru Beds. The presence of tiny and very signable to one individual of Ceratosaurus (?) delicate pterosaurian and mammalian remains in roechlingi. the Middle and Upper Saurian Bed at Tenda-

Mitt . Mus. Nat.kd . Berl., Geowiss. Reihe 2 (1999) 55

02 Barosaurus, n-11 =Dicraeosaurus,n- 8

Fig .20. Frequency distribution of main body regions in sauropod skeletons from Tendaguru . For explanation see text . 1 - skull and neck, 2 - shoulder girdle and fore limbs, 3 - sacral and dorsal vertebrae, dorsal 5 ribs, 4 - pelvic elements and body region posterior limbs, 5 - tail

guru (Reck 1931, Galton 1980, Unwin & Hein- trated close to these two limb bones, indicating rich 1999, Dietrich 1927, Heinrich 1998, 1999) in- that dispersal of bones was minimal. The place dicates, however, that this taphonomic pattern of death must therefore have been the place of cannot be due to post-burial destruction of smal- final deposition. ler sauropod skeletal elements.

4.4 Individual size and age class distributions 4.3 Burial position The following account of the size distribution of The burial position of carcasses of dinosaurs is Tendaguru sauropods (Brachiosaurus brancai, difficult to assess because of the relatively small Barosaurus africanus) is based upon the humeri number of reasonably complete skeletons recov- and femora, and that of the ornithischians ered from the Tendaguru Beds. Moreover, the (Dryosaurus lettow-vorbecki) upon the femora, skeletons documented in the field sketches dis- because these bones are better represented in play different burial positions. the collection than other limb bones of these The specimen of the dryosaurid Dryosaurus taxa. Metric data were taken from Janensch lettow-vorbecki recovered from the Middle Sau- (1961a and unpublished) and from Sander rian Bed at Site Jg (WJ) was buried on its belly, (1999, in press), augmented by my own meas- with the left femur spread away from the trunk, urements. and a cervical vertebral column that was not The number of the measured specimens is straight or only slightly recurved (Fig. 13) . The rather small, making quantitative assessments skull and neck were apparently still articulated. difficult . Therefore, sauropod specimens from Russell et al. (1980: 172) mentioned that Ten- different dinosaur-bearing horizons were put to- daguru barosaurs "preferred to die laying on gether to form time-averaged populations, as their left side". The upright position of both left explained in the diagrams (Figs 21 and 22). humerus and left tibia of skeleton SII of the bra- Both humeri and femora of Brachiosaurus bran- chiosaurid Brachiosaurus brancai (Fig. 16) sug- cai were assigned to 30.0 cm intervals, those of gests that the animal became mired in soft mud Barosaurus africanus to 14.0 cm intervals. Life- and died laying on its belly. In-situ decomposi- age data of sauropod individuals are based on tion of soft tissues is indicated by the series of histological examinations by Sander (1999, in articulated cervical vertebrae as well as by paral- press). lel oriented dorsal ribs resting closely on each A data set published by Janensch (1961b) as other (Fig. 16) . Except for the cervical vertebrae, well as my own measurements can be used to the majority of skeletal elements are concen- establish a femur-size distribution-diagram for 56 Heinrich, W.-D., Taphonomy of Dinosaurs from Tendaguru

Dryosaurus lettow-vorbecki. The femora of Dryo- nately, histological data that could calibrate the saurus lettow-vorbecki are assigned to 5.0 cm in- size classes with ontogenetic data are not avail- tervalls. The diagram (Fig. 23) is based on a col- able. Paleohistological studies on femora of lection of femora recovered from the Middle Dryosaurus lettow-vorbecki from Tendaguru indi- Saurian Bed at Tendaguru Site Jg (WJ) Unfortu- cate rapid and uninterrupted growth (Chinsamy 1995). Fig. 21 shows the humerus- and femur-size distribution of Brachiosaurus brancai. The histo- grams reveals that the populations are dominated by humeri and femora ranging in length from 144 to 174 cm. Based on life history studies by Sander (1999, in press), humeri and femora of these dimensions are probably assignable to

30- n=23 25- 20- 15- 10- 5- A 0

11 0/0 - 40- n=17 35- 30- 25- 20- 15- 10- 5- B 0

0/0 40- n=15 35- 30- 25- 20- 15- 10- 5 - 0- 10 24 38 52 66 Fig. 21. Size-class distribution of humeri and femora of Bra- chiosaurus brancai from Tendaguru . Data grouped together as follows: A - subset of humeri from the Middle Saurian Fig. 22 . Size-class distribution of humeri and femora of Baro- Bed (n = 3), Upper Saurian Bed (n = 12), and Transitional saurus africanus from Tendaguru . Data grouped together as Sands overlaying the Trigonia smeei Bed (n = 2), B - subset follows : A - subset of humeri from the Upper Saurian Bed of humeri from the Middle Saurian Bed (n = 3) and Upper (n = 5), and Transitional Sands overlaying the Trigonia smeei Saurian Bed (n =12), C - subset of femora from the Middle Bed (n = 18), B - humeri from the Transitional Sands over- Saurian Bed (n = 11), Upper Saurian Bed (n = 3) and Transi- laying the Trigonia smeei Bed (n = 17), C - subset of femora tional Sands (n = 3), D - subset of femora from the Middle from the Upper Saurian Bed (n = 7), and Transitional Sands Saurian Bed (n = 11) . 1 - juvenile individuals ; 2 - adult in- overlaying the Trigonia smeei Bed (n = 8) . 1 - juvenile indi- dividuals ; 3 - old adult individuals. The age determination is viduals; 2 - adult individuals ; 3 - old adult individuals . The based upon life history studies by Sander (1999, in press) . age determination is based on life history studies by Sander Metric data from Janensch (1961a), Sander (1999, in press) (1999, in press) . Data from Janensch (1961a), Sander (1999, and own measurements in press) and my own measurements Mitt . Mus. Nat.kd . Berl., Geowiss. Reihe 2 (1999) 57 adult individuals. Adult individuals have also the sample is based on time-averaged bone as- been found in the adjacent humerus size classes semblages. (114-143 cm, 174-203 cm), suggesting that In this connection, it is worth mentioning that about 60% of the measured specimens of Bra- histological data obtained from three Humeri of chioaurus brancai may belong to adult indivi- Dicraeosaurus sattleri from the Upper Saurian duals. Old adult individuals are most likely re- Bed (Sites ab and O) also suggest that adult in- presented in the highest Humeri-size class dividuals predominate in the bone assemblages (204-233 cm). (Sander 1999, in press), as is the case for Bra- Humeri of juvenile individuals of Brachio- chiosaurus brancai and Barosaurus africanus. A saurus brancai are distinctly underrepresented, femur of Janenschia robusta collected from the and neonates have not been found so far (Figs. Upper Saurian Bed at Tendaguru Site 22 is attri- 21A, 21B) . The "first" appearance of old adult butable to an old adult individual (Sander 1999, individuals in femur-size class 174-203 cm and in press). the "last" appearance of adults in femur-size Dryosaurus lettow-vorbecki (Fig. 23) displays a class 204-233 cm (Figs. 21C, 21D) indicates that bimodal femur-size-distribution, with two peaks, the age groups overlap to significant degree, and one at the beginning and one close to end of the include individuals of different size. A similar in- plot, suggesting either that juvenile and adult in- terval of overlap is to be expected between juve- dividuals prevail or that sexual dimorphism oc- nile and adult specimens. curs. Sexual dimorphism cannot be excluded Consideration of the size frequency histo- with certainty, but it seems more likely that the grams for the humerus and femur of Barosaurus bimodality is the result of individual size varia- africanus reveals that adult individuals appar- tion. If this is correct, the pattern of size-distribu- ently prevail (Fig. 22), as in Brachiosaurus bran- tion may allow conclusions to be drawn regard- cai. Diagram 22A, which approaches a unimodal ing on the origin of the mass accumulations of size-class distribution is difficult to assess, since Dryosa.urus lettow-vorbecki, discussed below.

4.5 Attritional or mass mortality?

25- The origin of the extraordinary concentrations of dinosaur bones in the Tendaguru Beds has been the of subject of speculations ever since the first discovery of dinosaurs in the Tendaguru area. 20J The interpretation has to consider attributes of the recovered bone assemblages, paleoecological aspects, and pecularities of the depositional environment that influenced the involved taphonomical processes. According to Janensch (1914a, c; 1961a), the dinosaur-bearing deposits were possibly laid down in a lagoonal environment separated from the sea by reefs. The sedimentation was thought to have been affected by tides, episodic storms (hurricanes), wind waves, earthquakes, and changes in the regional climate (Janensch 1914c). The invertebrate fossil record suggests a brackish to limnic depositional environment for the Mid- dle and Upper Saurian Bed (Dietrich 1914; Hen- nig 1914b, c; Schudack 1999; Schudack et al. size classes/cm 1999), and records of Asmussia (= Estheria) from the Upper Saurian Bed may indicate peri- Fig. 23. Size-class distribution for the femur of Dryosaurus ods of drought (Janensch 1933, Russell et al. lettow-vorbecki from the Middle Saurian Bed at Tendaguru 1980) . Site Jg (WJ) . Metric data from Janensch (1961b: 164) and measurements obtained from recently recovered femora . To- As mentioned above, Janensch (1914c) sug- tal number of femora : 28 gested that the dinosaur bone assemblages from 58 Heinrich, W.-D ., Taphonomy of Dinosaurs from Tendaguru

Tendaguru resulted mainly from abrupt cata- In addition, articulated sauropod limb bones strophic killing events. Unfortunately, the precise buried in an upright position (Janensch 1914c) relationships of the quarried dinosaur bones to indicate that the body weight of the animals the enclosing sediments are far from clear, due could have been an important selective tapho- to the loss of many records. In addition, the pre- nomic filter for the dinosaurs from Tendaguru . cise stratigraphic position of the dinosaur-bearing So, for instance, the body weight of the juvenile sites within the Lower, Middle, and Upper Sau- Tendaguru sauropods may have been below the rian Bed is unknown . This is especially regretta- threshold for becoming stuck in the mud, unlike ble, since the depositional environment of the the adult animals that passed more frequently Tendaguru Beds underwent remarkable changes the "point of no return" due to their heavier during the Late Jurassic and Early Cretaceous. body weight. The dinosaur-bearing sites in the Transitional A similar taphonomic model has been sug- Sands that connect the Saurian Beds with the gested for the Late Triassic (Norian) prosauro- shallow marine sandstone beds are undoubtedly pods Plateosaurus (Sander 1992) and Sellosaurus marginal marine in origin because marine in- (Hungerbtihler 1998). Sander (1992: 256) argues vertebrates are associated with the dinosaurs. A that the absence of juvenile individuals of Plateo- humerus of Brachiosaurus brancai covered by saurus engelhardti in several European Knollen- oysters is reported, for instance, from Site Aa mergel sites is due to the "lower foot pressure" which is in the Transitional Sands above the of juvenile animals that were thus "below the Middle Saurian Bed (Janensch, GTE field catalo- threshold for becoming mired" in shallow de- gue : 140) . As previously mentioned, the state of pressions. preservation of bones recovered from the Transi- Hennig (1925) believed that spatially limited tional Sands suggests transport and redeposition mass accumulations of disarticulated bones of of bones prior to burial. Therefore, these enor- Kentrosaurus aethiopicus from the Middle mous concentrations of disarticulated dinosaurs Saurian Bed (Site St) and the Upper Saurian bones probably represent time-averaged bone Bed (e.g., Sites He and X), containing indivi- accumulations rather than a short-term mass duals of various size and age, resulted from a mortality event. sudden mass death event (see also Weigelt As mentioned above, most of the documented 1927) . Rich concentrations of disarticulated ske- skeletal accumulations from the Middle and Upper letal remains of Dryosaurus lettow-vorbecki re- Saurian Beds are single-individual and mono- covered from the Middle Saurian Bed at Site specific. Multi-individual assemblages are rare, Jg (WJ) were also thought to represent an in- and death assemblages of Brachiosaurus brancai, stance of sudden mass mortality (Janensch Dicraeosaurus sattleri, and Barosaurus africanus 1914c) . Most limb bones (e.g., femur, tibia, fibu- are apparently dominated by limb bones of adult la) were found parallel to one another, with individuals. This taphonomic pattern does not their long axis oriented northwest to southeast, necessarily support the catastrophic mass mortal- indicating that these bones were accumulated ity model, since killing agents that produced under the influence of wave action (breakers) mass mortality should have affected all indivi- or wave-induced currents (Janensch 1914c: 256). dual age classes of the sauropod populations. Janensch (1914c) supposed that herds of small However, this conclusion should be made ornithischians that had entered lagoons during with caution because actualistic studies on large periods of drought were killed by the returning extant land mammals reveal that patterns of flood and became embedded close to the place bone accumulations may result from patterning of death. If this scenario is correct, Dryosaurus in behaviour (Haynes 1993). Younger African lettow-vorbecki left rapidly formed mass assem- elephants, for instance, sometimes die "much blages of bones in the Tendaguru Beds that re- nearer the inner areas" of die-off sites "than older sulted from an external catastrophic killing individuals" due to aggressive interactions at ex- event. Accumulations of sauropod bones in the cavated wells during severe drought (Haynes Middle and Upper Saurian Beds are also 1993:131). As in extant African elephant popula- thought to have been caused by similar killing tions (Haynes 1990, 1993), seasonal drought events, since the huge sauropod dinosaurs were could have resulted in social fissioning and also considered to be gregarious land animals ephemeral aggregations of adult and old adult (Janensch 1914c). Tendaguru sauropods at shrinking water sources By contrast, Reck (1925) who quarried Ten- and feeding sites. daguru Site Jg (WJ) in 1912 doubted the cata- Mitt. Mus. Nat.kd. Berl., Geowiss. Reihe 2 (1999) 59

strophic mass mortality. He argued (Reck 1925: mals. Nevertheless, it is worth noting, that the 9-10) : "So sehr auch das herdenartige Auftreten interpretation as an attritional mortality profile einzelner Arten, und besonders die Herden- also agrees more closely with the data obtained zusammensetzung aus Tieren jeder Größe und from the Tendaguru sauropods (Figs. 21, 22) as jeden Alters für sie zu sprechen scheint, so well as with the distribution pattern of bones stehen doch auch ihr unüberwindliche Hinder- within the Tendaguru dinosaur beds. nisse entgegen, wenn man dem Sprachgebrauch The invertebrate fossil record and seasonality entsprechend unter Katastrophe nur ein seltenes, indicated by reworked caliche nodules might in- verheerendes Naturereignis versteht .. ... Aber dicate playa lake-like water bodies close to an zwei Hauptmomente führen . . .. . zur Ablehnung oscillating shoreline, and subject to either seaso- solcher Theorie: die großzügig gleichmäßigen, nal or long-term droughts. Cross-bedded sand- außer durch die Transgressionen nicht unter- stone beds within the Middle and Upper Saurian brochenen, feinklastischen Sedimentationsver- Beds, in turn, might be explained by seasonally hältisse und die regellose Verteilung der Funde rainy intervals that resulted in water flows, tem- in der Vertikalen innerhalb der Mergelmassen. porary lakes, and ponds. These ephemeral water Dadurch wird die Supposition jeder Katastrophe bodies might have been refuges and die-off sites unhaltbar, wenn man nicht jeweils einen ganzen for Tendaguru dinosaurs during seasonal Mergelhorizont als Niederschlag und Zeugnis drought . Most likely, the palaeoenvironment in einer Katastrophe ansprechen will. Dagegen the Tendaguru region was much more diverse aber spricht alles . . .. . Als generelles Entstehung- than generally thought so far. sprinzip der Lagerstätte kommt ... .. die Ka- In the absence of direct taphonomic studies tastrophe nicht in Betracht" . Reck (1925), on the and substantial parts of the doumentation of the basis of his studies of the Tendaguru region, original expedition, it is beyond the scope of the concluded that the dinosaur bones were ir- present account to establish a detailed tapho- regularly dispersed within the Tendaguru saurian nomic model for the Tendaguru dinosaur accu- beds, and that the origin of the bone acumu- mulation. However, the existence of a number of lation could not be explained by catastrophic superimposed dinosaur-bearing horizons (Sau- mass mortality. This distribution pattern suggests rian Beds, Transitional Sands) shows that dino- that the bone accumulations resulted from recur- saur bones were accumulated over a long time ring "normal" death events of dinosaur indivi- span during the Late Jurassic. The fossil record duals over a long period of time rather than also reveals that extraordinarily suitable condi- from a catastrophic, short-term mass mortality tions for the preservation of dinosaurian bones event. occured repeatedly. The year-to-year input of In addition to this argument, further support bones resulting from short-term intervals of mor- for an attritional origin is found in the femur-size tality due to drought might have been consider- distribution pattern of Dryosaurus lettow-vor- able and "simulates" sudden catastrophic mass becki which represents an age profile, dominated mortality events. If so, the Tendaguru fossil re- by juvenile and adult individuals (Fig. 23). This cord may reflect attritional mortality rather than pattern (Fig. 23) shows a striking similarity to the sudden catastrophic mass mortality. so-called U-shaped or attritional mortality type of profile that is due to a long-term input of bones resulting from "normal", ecologically re- Acknowledgements lated deaths in different-aged population mem- Unwin bers (Lyman 1996). This pattern differs remark- I am grateful to Prof. Dr. H.-E Schultze and Dr. D. (Museum für Naturkunde der Humboldt-Universität zu Ber- ably from the unimodal L-shaped, catastrophic lin, Institut für Paläontologie) for much-valued comments or mass mortality pattern in which younger age and improvement of the English, and to Dr. E. Cook (Uni- classes dominate and older age classes become versity of Bristol) for reviewing the manuscript . I extend my thanks, to Dr. E M. Sander (Universität Bonn) for discus- successively underrepresented (Lyman 1996). At sions and histological data of Tendaguru sauropods, to Dr. B. the present stage of investigation, I do not draw Curtice and Dr. L. Curtice (Phoenix, AZ, USA) for helping the conclusion, that the mortality profile in with the identification of cervical vertebrae of Brachiosaurus brancai, to Dr. M. Aberhan (Museum für Naturkunde der Dryosaurus lettow-vorbecki is attritional in ori- Humboldt-Universität zu Berlin, Institut für Paläontologie), gin, since possible influence of the sexual di- Prof. Dr. H. Keupp and Dr. M. Schudack (Freie Universität the litho- morphism is far from clear and the interpreta- Berlin, Institut für Paläontologie) for comments on logy of the Tendaguru Beds, to Mrs. E. Glass (Humboldt- tion of U-shaped mortality profiles is mainly Universität zu Berlin, Institut für Mineralogie) for X ray dif- based on experience with extant and fossil main- fraction analysis, to Mr. J.-P. Mendau for the redrawing of 60 Heinrich, W.-D., Taphonomy of Dinosaurs from Tendaguru the site maps, to Mrs . V. Heinrich for the diagrams, and to Mead, J. I, Nelson, L. W. (eds .). Megafauna and man: dis- Mrs . W. Harre (all Museum für Naturkunde der Humboldt- covery of America's Heartland : 22-31, The Mammoth Universität zu Berlin) for the photographs . Financial support Site of Hot Springs, South Dakota Incorporation, Hot from the Deutsche Forschungsgemeinschaft (He 2757/1-1) is Springs, South Dakota . gratefully acknowledged . - 1993. Mammoths, mastodonts & elephants . Biology, Be- havior, and fossil record. 413 pp. Cambridge University Press, Cambridge. Heinrich, W-D. 1991 . Über Brancatherulum tendagurense References DIETRICH, 1927 (Mammalia : Eupantotheria) aus dem Oberjura von Tendaguru, Tansania. Vorläufige Mittei- Abel, O. 1927. Lebensbilder der Vorzeit . 637 pp., Fischer, lung . - Mitteilungen aus dem Zoologischen Museum Jena. Berlin 67(1): 97-104 . Aitken, W G. 1956. 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