New Records of Staffia Aenigmatica (Mammalia, Allotheria, Haramiyida) from the Upper Jurassic of Tendaguru in Southeastern Tanzania, East Africa

Mitt. Mus. Nat.kd. Berl.. Geowiss. Reihc 4 (2001) 239-255 05. I 1, 200 I

New records of Staffia aenigmatica (Mammalia, Allotheria, Haramiyida) from the Upper Jurassic of Tendaguru in southeastern Tanzania, East Africa

Wolf-Dieter Heinrich'

With 6 figurcs

Abstract

Tivo non-inultituberculate allothcrian cheek teeth are described from the Upper Jurassic of Tendaguru in southeastern Tanza- nia. East Africa. Both specimens were collected from dinosaur-bearing matrix of bone bed Wj of the Middle Saurian Bed at Tendaguru Site dy by thc German Tendaguru Expedition (1909-1913). Bone Bed Wj represents liinnic to brackish deposits of Kimnieridgian-Tithonian age. The cheek tceth, considered as lowcr posterior molar and upper molar, represent a single taxon ol the Haramiyida and are referred to S'tmffia nenigmnfica, known only from the Upper Jurassic of Tendaguru. This assignment reinforces evidcnce for the palaeogeographic dispersal of haramiyids to Gondwana and the temporal persistence of these non-multitubcrculate allotherians into the Late Jurassic. Characters that distinguish Staffin oenigmrtico from other haramiyids include the medial position of main cusp a1 at the front of the tooth crown and the presence of a largc, anterolingual main notch between cusps a1 and a2 in lower cheek teeth, as well as the development of a strong anterolabial cingular ridge in thc only known upper cheek tooth. Raffia shows the closest resemblance to Thomnsin from the Late Triassic to Early Jurassic of Europc, although these genera are disdinctly different. Retention of the basic tooth crown pattern of haramiyids and traces of wear in the Tendaguru teeth suggest that the masticatory movements in Stuffin were essentially rcstricted to 3 longitudinal direction, as in Thomasici. It is suggestcd that owing to its central position at the front of the tooth crown the Iowcr main cusp al could have occluded in the central basin of the opposing upper molar during masticaiory movements.

Key words: Mammalia. Allotheria, Haramiyida, Sfaffiu, Upper Jurassic, Tendaguru, Tanzania, East Africa, Gondwano

Zusammenfassung

ALISdcm Oberjura von Tendaguru in Tansania, Ostafrika, werden zwei Backenzahne eines Haramiyiden beschrieben. Beide Ziihne stammen aus knochenfuhrendcn Gesteinsproben, die von der Deutschen Tendaguru Expedition (1909- 19 13) in der Fundstclle dy gesammelt wurdcn. Fundschicht der Haramiyidcn-Zihne ist eine knochenfuhrende Lage (Wj) dcr Mittleren Saurierschicht, die im Zeitraum Kimmeridge-Tithon in einem kustennahen Ablagerungsraum entstand. Bcidc Backenzghne. ein hinterer unterer Molar und ein obercr Molar, werden ZLI Staffin acnigmrrticn gestellt, die bisher nur aus dein Oberjura von Tendaguru bekannt ist. Beide Nachweise bestatigen erneut. daR Haramiaiden einst in Gondwana verbreitet waren und dort noch in dei- sp#ten Jura-Zeit vorkamen. Merkmale, die Stn,ffiia rzenigmatica von anderen Haramiyiden unterscheiden. sind die 7entrale Position des al-Hockers im Vorderabschnitt der Zahnkrone und die tiere, breite anterolinguale Furchc zwischcn deni al- und a2-Hockcr der unteren Backenzahne sowie die starkc labiale Cingulumleiste am einzigen bisher bckannten oberen Molarcn. Zwischen Staffii/ aus dem Oberjura Ostafrikas und Thornasin aus der oberen Trias und dem unteren Jura Europas bcstehen Gemeinsamkeiten. aber auch wesentliche Unterschiede. Die Beibehaltung des Backcnzahn-Grundmusters der Hara- miyiden und Abkauungsspuren an den Zahnen aus Tendaguru zeigen, daR die Kaubewegung bei Sta,ffin im wesentlichcn in longitudinaler Richtung erfolgtc, wie bei Thornasin. Fur Staffia wird vermutet. daB der al-Haupthocker auf Grund seiner zentralen Lage im vorderen Abschnitt der Zahnkrone in das zentrale Becken des entsprechenden oberen Backcnzahnes pal3k und dort bei der Zerkleinerung von Nahrungspartikeln mitwirkte.

Schliisselworter: Mammalia. Allotheria, Haramiyida, Sraffin, Oberjura, Tendaguru, Tansania, Ostafrika. Gondwana

Introduction that is based upon associated lower and upper jaws with teeth (Jenkins et al. 1997). The origin Haramiyids are an enigmatic group of non-multi- and phylogenetic relationships of haramiyids tuberculate allotherians. They are only known have been the subject of much discussion (e.g.. from isolated teeth, except for Hnrnrniyavia Clemens & Kielan-Jaworowska 1979, Hahn & clernrnensi from the Late Triassic of Greenland Hahn 1983, Hahn et al. 1989, Kielan-Jaworowska

' Museum fur Naturkunde, Institut fur Palaontologie, Tnvalidenstr. 43, D-10115 Berlin. Germany. E-mail: [email protected] Reccived June 2001, accepted July 2001

( WILEY-VCH Verlag Bcrlin GmbH. 13086 Berlln. 2001 I435-I943101/04I 1-0239 '$ 17 50- 50,O 240 Heinrich. W.-D.. New records of Jurassic mammals

1992. Butler 6: McIntyre 1994. Butler 2000. Kie- nian Tendaguru Expedition (1909-1913) from Ian-Jaworowska & Hurum 2001). The earliest re- the Middle Saurian Bed at Tendaguru has re- presentatives of the order Hatamiyida have been sulted in the discovery of two further haramiyid reported from the Upper Norian (Upper Middle cheek teeth. Both record? arc particularly impor- Keuper) P1mosmrrii.s Beds of Halberstadt in tant in that they add substantially to the knowl- Central Germany ( Tlio/urisiu hnliiii: Hahn 1973. edge of Gondwana haramiyids and also contri- Burler & MacIntyre 1994) and from the '? Nor- bute significantly to the understanding of the ian-Rhaetic Tiit Bjerg Beds of the Fleniniing Tendaguru local land vertebrate fauna, thus play- Fjord Formatim in East Grcenland (Harv/iii!,rr- ing a key role in rcsearch on Jurassic faunas of via clem?ierisi Jenkins et al. 1997). Most hara- the African mainland. miyids have ken recovered from Late Triassic The results presented here are part of a conti- to carly Jurasiic localities in Central and Wes- nuing Tendaguru project of the Institut fur Pa- tern Europe. Ilescribed genera include represen- laontologie der Humboldt-Universitiit 7u Berlin tatives of the families Therotcinidae ( Tlierorei- supported by the Deutsche Forschungsge- 11 I is : Sig)gnea it - Russel 1 I 9 8 3. Si g o gnea LI - R u s se 11 meinschaft (He 275711 -3). et 211. 19x6. BLtler 2000). and Haramiyidae ( Tho- mrsiri. includitig HuuuIiijw: e.g.. Plieninger 1837. Branca 19 15. Simpson 1928. Parrington 1947. Geological and palaentological setting Peycr 10%. Hahn 1973. Clcmens 1980. Hahn & H a I1 ti I 9 8 3. S I gogiiea ti - R usse I I I 9 89. Si g og n ea LI - Detailed accounts of the geological structure and Russell & Hal- ti 1994. Butler & MacInt>re 1994. the stratigraphy of the Mesozoic deposits in the Delsate 1995. 2000. Godefroit et al. 199s. Butler Tendaguru area are given by Hennig (1914, 2000). More rxently. Kermack et al. (1998) de- 1937). Janensch (1914a), Aitken (1961), and scribed Elr I i rl? vodot I osfh1.t 1tv I sis froin t he Brit - Kent et al. (1971). A brief summary of these ish Middle Jtii-assic (Bathonian) and pi-oposed data is presented by Russell et al. (1980) and the lieu allot iherian suborder Eleutherodontidu Heinrich (1999b). Therefore, only a few com- (order i/icrrttre setlis). ivhich \\:is assiped to the ments on the stratigraphy of the Tcndaguru Beds order Haranii!.ida by Butler (2000). In addition that are relevant to the description of the. non- to the genera mentioned abo\ e. possible hara- multituberculate allotlierian remains described o been recorded from the Lonu below. are noted here. Jurassic of North America (Jenkins et al. 1983. The stratigraphical terminology follows Ja- Butler 2000) aiid the Middle Jiiriissic (Bat1ioni:iii) nensch (191 4a) and Russell et al. (1980). Accord- of England ( FIeenian 1976. Butler 2000). ing to Jancnsch (1914a), the Tendaguru Beds are Until recently. haramiyids were unknown out- part of the Lindi Formation, a stratigraphic term side Laurasia. However. microvertebrate sam- introduced by Dacque & Krenkel (1909). Strati- pling in dinosaur-bearing matrix from the Tcnda- graphic names such as Tendaguru Beds, Tvigonia guru Beds in .outheastern Tanzania led. in 1999. sriieei Bed and Middle Saurian Bed (Fig. 1) do to the discovet y of Stci,f'fi"i rreriigri~citicrr.extending not meet adequately the international strati- the hitherto known range of haramiyids to the graphic guidelines of the International Union of Late Jurassic and the known palaeogeographic Geological Sciences. More appropriate names distribution of thcse non-multituberculate al- such as Tendaguru Formation or Middle Saurian lotherians fro n the Northern Hemisphere to Member used sporadically in the literature are Gondwana (Hlinrich 1999a). ignored here. since the introduction of new stra- Sriiffici neriignrnricri is, as yet. a poorly known tigraphic terms should be based on new litho- taxon. The holotype and only known specinien is and biostratigraphic investigations in the Tenda- a moderately well preserved cheek tooth which guru region and these are still in progress was tentativel:.r identified as a lower posterior (Heinrich et al. 2001). premolar- and provisionally placed in the family The Tendaguru Beds, covcring large areas in Haramiyidae (Heinrich 1999a). Unfortunately. southeastern Tanzania, are best exposed in the the fragmental y nature of the type specimen did surroundings of Tendaguru Hill and most well not allow prtxisc conclusions concerning the known for the abundance of dinosaur bones tooth ct-oiv ti ni 1m-ph ol c)gy. (e.g.. Janensch 1914b, 1929, 1961, Parkinson Since thc fitst report of a Late Jurassic hara- 1930. Russell et al. 1980, Heinrich 1999b). The miyid from Crondwana. continued micro\.crte- sedimentary sequence reaches about 140 m in hate samplin: in matrix collected by the Ger- thickness and consists of three dinosaur-bearing Mitt. Mus. Nat.kd. Berl.. Geowiss. Reihc 4 (2001) 24 1 deposits, the Lower, Middle, and Upper Saurian rediscovered hand-written letter of H. Reck, who Beds, and three marine sandstone units, the Neri- directed the excavations at Tendaguru Site dy in nea Bed, Trigonia smeei Bed and the Trigonia 1912 and 1913, was very helpful, as it contains a schwarzi Bed (Fig. 1). short comment and a simplified sketch of the Ever since dinosaurs were first discovered in profile showing the precise stratigraphic rclation- the vicinity of Tendaguru Hill in 1906, the age of ships of the two previously-mentioned bone beds the Tendaguru Beds has been the subject of con- (Fig. 3). The comment reads as follows: “Dcr troversial discussion. Although the sequence was Graben ist gewifi der gleiche. Aber das Jg Mate- once considered as Cretaceous in age (e.g., Fraas rial Hennig’s stellt cine scharf begrenzte Knoche- 1908, Kitchin 1929), the Tendaguru Beds are nansanimlung dar, welche irn Liegenden meiner now dated as Late Jurassic, except for the Trigo- Wj Herde ruht. Teilweise licgen die Herden so- nia schwarzi Bed which is equated with the gar direkt ubereinander, jedoch ist eine fossil- Early Cretaceous (e. g. Dietrich 1933, Aitken leere oder doch sehr arme Bank eingeschaltet” 1961, Russell et al. 1980, Heinrich et al. 2001). (The quarry is certainly the same one. But Hen- The age of the haramiyid-bearing deposits will nig’s Jg material consists of a sharply-defined be further discussed below. bone accumulation, which lies beneath my W.1 The haramiyid material described herein “herd”. In part, the “herds” even lie directly comes from Tendaguru Site dy (Jg, Wj) that is atop one another, however, a non-fossiliferous or situated approximately 2.5 km north-north-west at least very (fossil) poor layer is intercalated be- of Tendaguru Hill (Fig. 1). It was recovered from tween them). Consequently, the bone bed WJ. as matrix of the Middle Saurian Bed which overlies he defined it, is above bone bed Jg, and it the Nerinea Bed and underlies the Trigonia should be noted that the symbols Jg and Wj rc- smeri Bed in the Tendaguru region (Hennig fer merely to two bone-bearing strata in a single 1914, Janensch 1914a). The locality was origin- site (dy) and do not relate to two different local- ally named as Site Jg (Janensch 1914b: 51, map ities in the Tendaguru area. H. Reck’s letter, on p. 45) or Jg (Wj) (Janensch 1925: map on p. written on February 10, 1914, is of importance as XVIII) and later re-named as dy (Janensch 1955: it makes possible, for the first time, to distinguish 107, foot note l), a symbol that derives from unequivocally two successive assemblages of ter- Dysalotosnurus (now Dryosaiirus), because the restrial vertebrates within the Middle Saurian Middle Saurian Bed at this site was found to Bed. contain the ornithischian dinosaur Dryosaurus Both allotherian cheek teeth described below Zerrowvorbecki, unrecorded elsewhere in the Ten- were obtained from blocks of bone bed W,j, col- daguru area (Janensch 1955). Site dy was one of lected during the excavations of the German the richest and most extensively hand-quarried Tendaguru expedition between 1911 and 1912 at vertebrate localities in the Tendaguru region Site dy and housed now in the Museum fur Nat- (Fig. 2). urkunde Berlin (Institut fur Palaontologie). The The Middle Saurian Bed at Tendaguru Site dy specimens were hand picked under a microscope contains two bone-bearing strata, characterized from the residue of matrix which was dissolved by disarticulated dinosaur bones of Dryosaurus in acetic acid. Iettowvorbecki that almost completely dominate Specimens recovered from the matrix of bone the deposits (Janensch 1914a, b, 1955; Heinrich bed Wj also include skeletal remains of or- 1999b). Apart from Dryosaurus lettowvorbecki nithischian dinosaurs (Kentrosaurus aethiopiirs, only a few isolated stegosaur and sauropod re- Dryosaurus lettowvorbecki), unidentified thero- mains have been found (Branca 1914, Janensch pod and sauropod dinosaurs, a dwarf crocodile 1914a, b). The depositional environment of the similar to Bernissartia (Heinrich, unpublished), Middle Saurian Bed at Tendaguru Site dy was pterosaurs (Tendaguripteriis recki: Unwin & mainly limnic to brackish in origin (Heinrich Heinrich 1999), and the holotype of the hara- 1999 b) . miyid Staffia aenignznricn (Heinrich 1999a). The Unfortunately, Janensch (1914a, 1929, 1955) mammalian records of Tendagiirodon janeiisclii did not provide profile sketches or unequivocal and Tendagurutherium dietrichi described pre- names for the two above mentioned bone-bear- viously from Tendaguru Site dy (Heinrich 1998) ing strata of Tendaguru Site dy. This presents a are from bone bed Jg and associated with a para- problem, since the material of Site dy which was macellodid lizard (Broschinski 1999), an unidenti- designated as Jg or Wj cannot be related to fied dwarf crocodile (? Bernissarria), and Dryo- either of the two bone beds with certainty. A saurus lettowvorbecki (Heinrich, unpublished). 242 Heinrich, W.-D., New records of Jurassic mammals

"II 'FTrigonia schwarzi

Saurian - c/) -

Trigonia - smeei - Bed

- Nerinea

Bed t- t- -

E lower : Saurian j Bed 0

Fig. I. Map of the Tendaguru area in southwestern Tanraiiia nith a generalized section of the Tendaguru Beds and the topographic location ol the mammal-hearing Site dy considered in present account. TS. Transitional Sands; MK, remains of coarse to pebblepined mluviatil deposits on top of Tendaguru Hill assigned to the Makonde Beds by Janensch (1914a), true thickness is not shown. Uote also that age of the Makondc Beds is uncertain (see Ken1 et al. 1971). Contour heights in fcct. Source: Hcnnig (1914). Janensch (10141. 1925). Heinrich (1999h): sheet 'Nanjirinje' (no. 281/1) of the topographic maps (wries: Y 742: cdilion I-TSD). Tanzinia. Mitt. Mus. Nat.kd. Bcrl.. Gcowiss. Rcihe 4 (2001) 14.;

Fig. 2. Photograph of Tendaguru Site dy, the type locality of the haramiyid Stufjia rienignzaticn. Type specimens of fi~ltft7gllrO- dot7 j(rt7wschi3 TL.nrlayirrii~heriiit~zrlietrichi and Dryosn~irlrslertowvorbrcki have also been recovered here. in exposures of the Upper Jui-assic Middle Saul-Ian Bed. The position of the two super-imposed bone beds Jg and Wj is likely to have been close to the base of the exposure. Photograph by Hans G. Reck in 1912. Source: Institut fur Palaontologic der Huniboldt-1Jniversi- tat zu Berlin

Miospore and dinoflagellate assemblages re- the U-shaped ridge is mesial and the row of covered from matrix, housed in the Museum fur three cusps (row A) is labial (Jenkins et al. Naturkunde Berlin (Institut fur Palaontologie), 1997). The cusp terminology used in this paper suggest a Kimmeridgian to Tithonian age for follows Jenkins et al. (1997) and Butler (2000) bone bed Wj (Schrank 1999, 2000), whereas as- unless otherwise noted. The cusps are labelled sociated charophytes (Schudack 1999) and ostra- with letters (row symbols) combined with arabic cods (Schudack & Schudack 2001) indicate a numerals and numbered from mesial to distal in Kimmeridgian age for this bone-bearing deposit. lower cheek teeth, and from distal to mesial in Further biostratigraphic studies are needed to upper cheek teeth (Fig. 4), as suggested by Jen- determine more precisely the age of the two kins et al. (1997) and Butler (2000). Capital let- bone beds at Tendaguru Site dy. ters are used for cusps of upper (maxillary) cheek teeth, lower case letters for those of lower (dentary) cheek teeth. The terms applied here to Cusp terminology cusps of non-multituberculate cheek teeth are primarily descriptive and do not imply homolo- Tooth crowns of haramiyid chcek teeth typically gies with cusps of cheek teeth in multitubercu- have two rows of mcsiodistally aligned cusps, a late allotherians which are similarily named in central basin, and an U-shaped ridge connecting the literature. Thc cusp teniinology used is illu- both cusp rows (Fig. 4). In lower chcek teeth, strated in Figure 4. the U-shaped ridge is distal and the row with the Butler (2000) has recently demonstrated, with highest mesial cusp (row A) is lingual (Hahn reasonable certainty, the cusp homologies of the 1973, Jenkins et al. 1997). Contrary to previous molariforms in the haramiyids. In the lower mo- intcrpretations, however, in upper cheek teeth lars of Hararniyavia, cusp b2 is placed posterior 233 Heinrich. W.-D.. New records of Jurassic mammals

Fig. 3. Excerpt of in unpublished hand-\\ritteil letter of Hans G. Reck. leader ol the Gcrnman Tendaguru Expedition in 1012 and 1913. showing the superpositional relationships of the two hone-bearing beds Jg and Wj exposed at Tcndaguru Site dy between 1912 and 1913. The Icrter dates from February 10. 191-1. Sourcc: Institut for Palaontologie der Humboldt-IJniversitat zu Berlin.

to the main cu;p at (Jenkins et al. 1997: 717. fig. New fossil material of Stuffiu 4: Butler 2000: 321. fig. 1). This suggests that the cusp in Thonwsia. that was traditionally consid- Lower posterior molar (MB.Ma.50069) ered as bl (e.:.. Hahn 1973. Sigogneau-Russell 1989, Butler & McIntyre 1994). is probably The crown of the new lower cheek tooth is com- homologous w th cusp b2 of HriraiiiiJwvicr (But- plete. well preserved, and practically unworn; the ler 2000: 320. f.g. 1) aiid should be labelled as b2 root has been broken away. The occlusal outline rather than bl. Since the dental morphology of of the crown is approximately pear-shaped Staffin is simi1;ir to that of Thoriirisirr in several (Fig. 511A). The margins of the tooth crown, as aspects. it is also probable that cusp bl in thc seen from above, are rounded, with well devel- Tendaguru specimens (Fig. 4) might be homolo- oped indentations, especially in the labial and gous with cusp b2 in Hmniiziyavia. as recently posterior wall. corresponding to the external suggested by 13utler (2000). A distinct swelling openings of the notches. The tooth crown is on the labial dope of the main cusp a1 in the slightly longer than wide, with the greatest width type specimen of Sraffa aciiigmarica. that might approximately across cusps a2 and bl (Fig. 5/1A). be the result of the fusion of cusp a1 with bl. Specimen MB.Ma.50069 exhibits the basic har- would support this interpretation ( Heinrich amiyid tooth crown pattern. The two rows of 1999a). cusps bordering the central basin are curved However. Tlioina.sia from the Late Triassic to (Fig. S/lA). aiid each consists of three cusps de- Early Jurassic of Europe and Stoffiri from the creasing gradually in height from front to back Late Jurassic If East Africa are separated by (Fig. Y1B-1C). Cusp row a is slightly higher long temporal interval. Moreover. the available than row b, with principal cusp a1 distinctly lar- haramiyid mat’xial from Tendaguru is too poor ger and higher than the other cusps of the tooth to yield defini! e data regarding cusp homology. crown. In all cusps, the basin surface is steeper More complete material is needed before signifi- than the marginal surface. cant conclusior!s can be made. Thus the descrip- The cusp height gradient of row a is: tive cusp termi iology is retained here (Fig. 4). a1 > a2 > a3. The main cusp a1 is shifted medi- Mitt. Mus. Nat.kd. Bcrl., Geowiss. Reihe 4 (2001) 245

a1 a1

bl bl 62 b2 A2 b3 IA b3

B4 B3 82 BI L1-W

...... 3B ...... 28 BI B2 A2 A3 A4

......

...... - ...... -... A1 3c

y /.../J AA2 A4 3D

a2 B2twA2...... B1 A1 1E b3 2E 3E

Fig. 4. Dental terminology for cheek teeth of Stuffiu aenigrnuficu from the Upper Jurassic of Tendaguru. 1. right lower posterior molar tentatively identified as m3; 2. holotype specimen, tentatively identified as right lower posterior premolar (? anterior molar); 3. left upper molar tentatively identified as M2. Not to scale. The cheek teeth are shown in occlusal (A). lingual (B), labial (C), mesial (D), and distal views (E). al-a3, lingual cusps of lower cheek teeth (cusp row a): bl-b4. labial cusps of lower cheek teeth (cusp row b); Al-A4, labial cusps of upper cheek teeth (cusp row A); Bl-B4. lingual cusps of upper cheek teeth (cusp row B); AA1, AA2, cingular cusps; pr, U-shaped posterior rim. 246 I-Ieinrich. W.-D.. New records of Jurassic mammals

IA 2A 3A

IB 3B

IC 3c

ID 3D

IE 2E 3E

Fig. 5. Srtrffrtr w/i~:~/iiu~ic~~i11-om the Cpper Jurassic of Tenciapru in Taniania. East Africa. 1. MB.Ma.50069, right lower posterior molar. tenta i\el! identified as m.3: 2. blB.-ISOSO. t! pe specimen. tentati\ely identified ;IS right lower posterior premolar ('? anterior molar) 3. blB.Xla.50070.left upper molar. tenlati\,eiy identified as M2. The cheek tccth are shown in occlusal (A). lingual (B).labial (C). niesini (U).and distal \.ie\vs (E).Scale bars = Imm. Mitt. Mus. Nat.kd. Berl., Geowiss. Reihe 4 (2001) 247 ally and located in the longitudinal midline of the posterior end of the preceding molar. The the tooth crown. When viewed in longitudinal root is not preserved, but the broken, tear- section, principal cusp a1 is remarkably asym- shaped outline of the base of the tooth crown metric, owing to its vaulted anterior and steep suggests that it extended longitudinally, with the posterior face (Fig. YlB-lC). Cusp a2 is sepa- greatest width just below the posterior portion of rated from the preceding main cusp a1 by a the central basin. Acessory roots are not present. large notch, open anterolingually, and with a The dimensions of the tooth crown are as fol- broad U-shaped floor (Fig. 5/1A-lB). Cusp a2 lows (in mm): maximum length (across cusps is much smaller than cusp a1 and distinctly se- al-b3): 0.92, maximum width (across cusps parated from cusp a3 by a well-developed trans- a2-bl): 0.70. verse notch. Cusp a3 is slightly lower than a2. The notch separating cusps a2 and a3 is distinctly smaller and shallower than that between Upper molar (MB.Ma.50070) a1 and a2. (Fig. 5/1B). Labial cusp row b is sligtly more curved than Even though it has suffered considerable wear the lingual row (a), and well-developed notches and corrosion, enough of the tooth crown is pre- separate the cusps. Among the three cusps mak- served to identify specimen MB.Ma.50070 as an ing up row b, bl is by far the largest. The cusp upper molar of a haramiyid. The tooth crown is height gradient of row b is: bl > b2 > b3. Cusp imperfectly preserved, heavily worn and cor- bl is separated from the labial slope of the prin- roded, with most of the original tooth crown pat- cipal cusp a1 by a deep notch, open anterola- tern obliterated. The posterolabial portion and bially. The basin wall cusp bl is displaced close the posterior margin of the crown are missing, so to the longitudinal midline of the tooth crown that nothing can be said about the structure of (Fig. 5/1E) and, as seen in occlusal view, drawn the tooth crown in this region. The roots have out anterolabially into a curved swollen ridge also been broken away. that tapers anteriorly and slopes towards the The tooth crown tapers anteriorly. Its occlusal notch bordered by cusps a1 and bl (Fig.S/lA). outline is asymmetrical because of the strong cin- Cusp b2 is markedly smaller than bl , barely gulum crest on the anterolabial flank of the higher than b3 and lies approximately opposite tooth crown (Fig. S/3A). The anterior margin, as cusp a3. Cusp b3, similar in girth and height to seen from above, is well-preserved, slightly in- b2, is displaced into the longitudinal mid-line of dented, and recedes towards the labial wall of the tooth crown. All cusps lack enamel ridges. the crown. The lingual margin is nearly The U-shaped posterior ridge bears the closely straight, but with faintly developed indentations spaced unworn cusps a3, b3, and b2. Cingulids (Fig. 5/3A). and single accessory cingular cusps are missing. The occlusal surface exhibits the basic hara- The central basin is deep and distinctly nar- miyid structure. There is an elongate central ba- rowed by the barrier-like projecting cusp bl sin, flanked on both sides by rows of mesiodis- (Fig. 5/1E), broader anteriorly than posteriorly, tally aligned cusps (row A and B), and a worn. and with the deepest point close to the rising U-shaped ridge that joins both cusp rows ante- basin face of cusp bl. The central basin extends riorly (Fig. 5/3A). From what can be seen of the an terolingually towards a broad trough-like badly preserved occlusal surface, the lingual cusp notch just posterior to principal cusp al, and it is row (B) is higher than the labial (A), but the also linked anterolabially to a deep notch, which two rows are of approximately the same length. has an approximately V-shaped cross section. The Cusp height gradients cannot be determined posterior portion of the central basin is smaller owing to the intensive wear that has truncated than the anterior, narrowed by cusps a3 and bl, most of the cusps, especially in the anterior half and opens externally through two small notches. of both cusp rows. The enamel surface of both the central basin The labial row of cusps (row A) consists of and the cusps is smooth: wrinkled (rugose) en- four approximately aligned cusps, of which the amel is not developed. The tip of the principal first, Al, is almost completely worn. The pre- cusp, al, is very slightly worn. No evidence of sence of this cusp is barely discernible, but the wear was found either on the tops of the other remnants of the cusp base stand well above the cusps or in the central basin. deeply excavated central basin, suggesting that a On the anterior edge of the tooth crown, there distinct cusp may have been present at an earlier is no indentation (recess) for the reception of stage of wear (Fig. 513E). There is no evidence 248 Hcinrich. W.-D.. New records of Jurassic mammals of a saddle-like ridge connecting the posterior- which the largest (AA2) is apparently at the most cusps A1 with BI. Cusp A1 is slightly dis- front. Accessory cusp AAl is situated somewhat placed medial1 y relative to A2. and separated labial to the notch between cusps A2 and A3, from the latter by an almost entirely obliterated and AA2 lies labial to the groove separating posterolabial iiotch. Cusp A2 appears to have cusps A3 and A4. Cusp row A and tlie cingular been the talkst and largest cusp of row A. ridge are separated by a shallow furrow, that be- though this is difficult to ;issess as the other comes slightly dccper postcriorly.

?_ cusps of row ii have been worn almost entirely Iwo closely spaced contact facets are present away. Cusp 1.2 is markedly better preserved on the anterolabial face of the tooth crown, just than the other cusps of row A (Fig. Y3C. 3E). It below cusps A4 and AA2. The roots are bears a feeble. elongate crest and exhibits a stee- broken away, but there appear to have been ply inclined arcxa on its basin face caused. appar- two of them. a larger transversely broadened ently. b!, wear Nothing can he said about the root close to the posterior end and a smaller original morphology of the heavily worn cusps root close to the tapered anterior portion of the A-3 and A4. wk ich in oblique light are just distin- tooth crown. guishable on the occlusal surface. The dimensions of the fragmentary tooth B2 appears to have been the largest of the crown are as follows (in mm): maximum length: four cusps in tlie lingual row: the top of the cusp 1.52: maximum width (across cusps B3-AA1): has been replxed b!. a facet. The preceding 1.10. cusp. B1. is s ightly displaced medial to main cusp B2 and barely discernible in anterior aspect (Fig. 513D). Thg: top of cusp R1 is replaced by an Comparison and discussion inclined facet. facing posterolingually. Cusps B 1 and B2 arc very closely spaced and indistinctly Referal of both the upper and lower molar from separated by a notch (Fig. 5i3B). sloping towards the Tendaguru Beds to the Allotheria is based the basin floor. Cusp B3 is heavily worn and. on several diagnostic features of the tooth when viewed n lateral aspect. separated from crowns. in particular the two mesiodistally B2 by a shallcw transverse valley. The base of aligned rows of cusps bordering a central basin the anteriormoit cusp. B4. lies dinstinctly higher and the IJ-shaped ridge linking the two cusp than that of B.; (Fig. 513B). The cusps of row B rows. The presence of the labiolingual and me- lie slightly pos .erior to the corresponding cusps siodistal reversal of the tooth crown pattern in of I-ow A, exct.pt for cusp B4 which is approxi- opposing lower and upper cheek teeth, and the mately oppositts to A4. smooth enamel surface in the well-preserved Between the two cusp rous (A. B). there is an lower posterior molar exclude the posibility that elongate centrid basin that reaches its greatest the Tendaguru specimens belong to a multituber- depth close to the anterior basin slopes of cusps culate mammal and support the taxonomic allo- Al and B1. Tie floor of the central basin ap- cation to the non-multituberculate allotherian or- pears, in oblique light. as a deeply excavated. der Haramiyida. narrow. longitudinal groove. indicating extensive S e r i a 1 p o s i ti o n . Except for Haramiyaviu, the wear. This grol)ve is closer to cusp row B than known haramiyid genera and species were exclu- A. and thc lingual basin wall is distinctly steeper sively defined upon characters of isolated cheek than the labia. (Fig. 513A. 3E). Anteriorly. the teeth. Therefore the precise numbers of cheek basin floor risei more gradually than posteriorly teeth in Elei I th erodon , Theroteinus, Thomasin, and its mesial end is marked by the U-shaped and Striffia are unknown. Consequently, the ref- ridge which is \vorn to a barely discernible ridge. era1 of isolated haramiyid cheek teeth from the The posteriorniost end of the central basin is Tendaguru Beds to a particular serial position is brokcii off. The enamel surface of the central speculative. Assuming that Staffia possessed basin and cusps appears to have been smooth. three lower and upper molars, as found in Hara- but clear evidence is lacking due to the imper- t~z~,a~*ia(Jenkins et al. 1997) and suggested for fect state of prt scrvation. Tlzot7irisiri (Butler 2000), the following tentative There is a prominent. but imperfectly pre- conclusions regarding the serial position of the served cingular ridge on the anterolabial edge of Tendaguru specimens can be made: the tooth crowii. extending from the anterolabial The type specimen of S(affiia aenigmatica was corner of the tooth crown posteriorly. and with originally compared with presumed premolars of evidence of a least two accessory cusps, of Tliomasin (tooth groop 11: Sigogneau-Russell Mitt. Mus. Nat.kd. Bed. Geowiss. Reihe 4 (2001) 249

1989) and provisionally regarded as a lower pos- teeth of Staffia aenigmatica should occur in the terior premolar (Heinrich 1999a). More recently, posterolabial portion of the tooth crown. Though Butler (2000) compared the cheek teeth of the imperfectly preserved, the Tendaguru specimen Thomasia tooth groop I1 (Sigogneau-Russell retains enough morphology to show that such a 1989) with the lower anterior molar (ml), sug- prominent notch appears to have been devel- gesting that the type specimen of Staffia aenig- oped exactly in the position where it would be matica might represent a ml. expected to occur (Fig. 5/3A). In view of this ob- By comparison with the lower dentition of servations referal of the ‘M2’ to Smffia aenigino- Haramiyavia and Thomasia, where the ultimate tica is warranted. molar is the smallest lower cheek tooth (Jenkins Ha r a m iy a via. Characters that distinguish the et al. 1997: 717, fig. 4; Butler 2000: 321, fig. 1), it m3 of Haramiyavia (Jenkins et al. 1997) from the is concluded here that the tiny isolated lower ‘m3’ of Staffia are: (1) the basically rectangular molar from Tendaguru is also likely to be an ulti- shape of the tooth crown; (2) a straight central mate lower molar (m3), referred to hereafter as basin bordered by approximately parallcl cusp ‘m3’. rows (a, b) and; (3) the position of main cusp al. As further outlined below, specimen MB.Ma. which is aligned with cusps a2 and a3. In con- 50070 from Tendaguru shows some similarities trast to Staffia, Haramiyavia has upper molars with upper cheek teeth from Thomasia, that with a roughly quadrate outline (Jenkins et al. were originally described as Haramiya tooth 1997: 416, fig. 1). In both genera, there are four group I by Sigogneau-Russell (1 989). According A cusps on M2, of which A2 is the largest. The to Butler (2000), it seems probable that this B row of M2 consists of four cusps in the Tenda- tooth group corresponds to the presumed second guru specimen, of which B2 is the highest, and upper molar in Thomasia, thus the Tendaguru five cusps in Haramiyavia, with B1 as the main specimen is tentatively regarded as M2, referred cusp in row B (Jenkins etal. 1997: 416, fig. 1). to hereafter as ‘M2’. Because of these differences, referal of the two A f f i n i t i e s. On the basis of the known materi- Tendaguru specimens to Haramiyavia appears al the affinities of the Tendaguru haramiyid can most unlikely. be outlined as follows: Tho m asin, The presumed m3 of Thornasin dif- Staffia. Despite the different size and serial fers from the ‘m3’ of Staffia in having: (1) a basi- position, the ‘m3’ from the Tendaguru Beds cally rectangular tooth crown and; (2) a double- shares several diagnostic features with the type peaked anterior profile (e.g., Thomasia sp: Par- specimen of Staffia aenigmatica: (1) a large and rington 1947: 714, fig. 4c, pl. 1, fig. 4a, b; Thointr- deep deep main notch between cusps a1 and a2; sia tooth group I Sigogneau-Russell 1989: (2) the length of the lingual cusp row a, which 140-151, fig. 7; Thomasia sp., Butler & MacIn- extends more anteriorly than labial row b; (3) a tyre 1994: 443, figs 4, 9h, i; Thomasia sp., Butler distinct anterolabial indentation of the buccal 2000: 321, fig. 1). Moreover, the supposed m3 of margin of the tooth crown; (4) a single-peaked Thomasia lacks the large notch separating cusps anterior profile and, more importantly; (5) the a1 and a2, a key character of Staffia that distin- medial position of main cusp a1 at the front of guishes it from Thomasia. the tooth crown, indicating the taxonomic affi- Owing to the fragmentary nature of the ‘M2’ nities of the compared molars. of Staffia comparison with Thornusio is tentative. Taxonomic evaluation of the haramiyid upper On the upper molars, there are three A cusps in molar from the Tendaguru Beds is more difficult, Thomasia, but four A cusps in Sk~ffa,although since most of the tooth crown pattern has been A2 is the principal cusp in both genera. In Stuf- worn away. However, there is a feature of the fia, cusp row A is as long as row B, as in the Tendaguru specimen, that suggests that it also Haramiya tooth group I sensu Sigogneau-Russell derives from Staffia aenigmatica. A key feature (1 989). A striking difference between these taxa of the lower cheek teeth in Staffia aenigmatica is the presence of a strong anterolabial cingular is the anterolingual main notch located just ridge in Staffia, a condition unlike that in othcr posterior to principal cusp a1 (Fig. 5/1A-lB, haramiyids, including Thomasia, where only 2A-2B). As haramiyid upper molars are both small cuspules or incipient cingula occasionally mesiodistal and labiolingual mirror images of the occur (Sigogneau-Russell 1989, Butler 2000), lowers (Clemens & Kielan-Jaworowska 1979, possibly foreshadowing the condition in Staff&. Butler 2000), a distinct notch in the upper cheek The differences in the dental morphology out- 250 Heinrich. W.-D., New records of Jurassic mammals lined above indicate that the Tendaguru speci- taxa. Staffin resembles more closely Thomusiu men are not referable to Thoinasia. than Haraniiyavia, Theroteinus or Eleutherodon. The dimensims of the type specimen of Staffla On the basis of the new material, the original (length: 2.1 mm: width: 1.0 nim) fall within the diagnosis for Stuffia uenigmatica (Heinrich size range of ihat of Thomasia (tooth group I1 1999a) can be modified as follows: sensu Sigogneau-Russell 1989), length: 1.36-2.17 Revised diagnosis. Lower posterior premo- mm, width: O.ti4-1.17 mm). This is also true for lar (? lower anterior molar): Tooth crown elon- the incomplett' 'M2' from Tendaguru (MB. Ma. gate, slightly pinched in the anterior portion, ta- 50070, length: 1.52 mm. width: 1.10 mm: Ham- pered posteriorly, with two approximately miyn tooth group I sensu Sigogneau-Russell aligned main cusp rows (a, b), bordering an 1989. length: 1.43-2.22 mm. width: 1.10-1.88 elongate central basin; double-rooted. Row a, mm). By contiast, the 'm3' from the Tendaguru higher than row b. with three cusps, of which a1 Beds (MB.Ma.50069: length 0.92 mm. width: 0.70 is the highest. Cusp height gradient: mm) is considcrably smaller than the size range a1 > a2 > a3. Main cusp a1 displaced medially, in Thomasia (tooth group I. Sigogneau-Russell and positioned at the front of the tooth crown. 1989, length: 1.09-2.49. width 0.74-1.61 mm). Row b with three cusps, cusp height gradient: 'The data suggests, that Stuffin had a cheek tooth bl > b2 > b3. Central basin closed by main cusp dentition simikir in size to that of Thomasia. a1 mesially, and a large U-shaped ridge distally, Eleurherodon. Cheek teeth of Eleirtherodori but open through the anterolingual main notch from the Middle Jurassic of England (Kermack and a deep anterolabial notch. The remaining et al. 1998. Butler 2000) differ markedly from notches are distinctly smaller, and noticeably the Tendaguru specimens in having: (1) upper narrower and shallower. The enamel surface is molars with a varying number of up to ten buc- smoo t 11. cal cusps: (2) lower molars with a labial cusp Lower posterior molar ('m3'): Tooth crown row that is higher than the lingual row. a varying noticeably reduced in size, pear-shaped in occlu- number of up to 21 cusps, and cusp bl located sal view. with two slightly curved cusp rows (a, mesially to cus 3 al: (3) the presence of accessory b): single-rooted. Cusp height gradients: cusps on the lingual margin of the upper molars bl > b2 > b3 and bl > b2 > b3. Row a higher and: (4) the presence of distinct enamel ridges than row b. Principal cusp a1 displaced into a (fluting) in lower and upper molars. Elleiithero- central position at the front of the tooth crown. don is one of tie most distinctive haramiyids and Central basin narrowed by cusp bl, but open a close relationship to Staffirr is unlikely. through the anterolingual main notch and a large anterolabial notch. Notches separating the Theroteinu.r. The cheek teeth of Theroteinus cusps in the posterior portion of the tooth crown from the Late Triassic of Europe (e.g.. Sigog- distinctly smaller and noticeably narrower and neau-Russell 1,483, Sigogneau-Russell et al. 1986. shallower. U-shaped ridge cusped. Enamel sur- Hahn et al. 19E9. Butler 2000) display several dif- face is smooth. ferences from the molars assigned to Staffia. in- Intermediate upper molar ('M2'): Tooth crown cluding: (1) lop, and blunt lower and upper molar tapered anteriorly, with an elongate central basin cusps; (2) the presence of additional lingual and two main rows of cups, of which cusp row cusps, instead if labial cusps. on the upper mo- B is higher than A. There are four A cusps, lars and: (3) the short central basin on the lower Al-A4, of which A2 is the highest, and four B molars bordered by a lingual cusp row with only cusps. Bl-B4, with B2 as main cusp in row B, two cusps (a1 and a2: Butler 2000). In view of and a strong anterolabial cusped cingular ridge, these differenc'zs, the specimens from the Tenda- that considerably broadcned the width of the guru Beds are .lot attributable to Theroteinus. tooth crown. The enamel surface is probably The number of cheek teeth from the Tenda- smooth, as in the lower cheek teeth. guru Beds is too small to allow detailed compar- isons, and mole definite statements of the affinities of Sruffiir must await the recovery of better Occlusion preserved mate rial. The available information on the dental morphology and size of the cheek Despite the discovery of two further molars, con- teeth suggests that the records from the Middle clusions concerning the pattern of occlusion and Saurian Bed ale best assignable to Stnffia nenig- chewing in Staffia remain uncertain, owing to the mritica. In com ,arison with the known haramiyid incompletely known dentition. The wear topo- Mitt. Mus. Nat.kd. Berl.. Geowiss. Rcihe 4 (2001) -.7i L

graphy of the ‘M2’ is difficult to decipher. Ex- Considering the specific cusp configuration of cept for A2, nearly all cusps have been worn al- the lower cheek teeth as well as the condition most entirely away and most indications of the of the central basin of the ‘M2’ from Tenda- the original tooth crown pattern have been oblit- guru, it is tentatively concluded here, that. in erated. B1 has a well-defined patch of wear, in- contrast to Thomasia, the main cusp a1 of the clined posterolingually. There is a triangular lower molars occluded in the central basin of wear facet on cusp B2, tapered anteriorly and the opposing upper molars, as illustrated in Fig- inclined slightly lingually. There are possible ure 6B. If so, the main cusp a1 must have un- traces of abrasion on the cingulum ridge, due dergone a remarkable evolutionary change. In perhaps to contact with food. Further evidence haramiyids, cusp rows a and A have mainly a of wear is visible in the central basin, wherein cutting function, whilst cusp rows b and B pri- the floor has been deeply planed, forming an marily crush and grind the food items accumu- elongate groove. There also seems to be an elon- lated in the central basins (Fig. 6A). If properly gate strip of wear just below the external (lin- interpreted, Staffia differs considerably from gual) wall of the worn cusp row B. By contrast, this pattern. Owing to its central position at the ‘m3’ is almost unworn and displays the original front of the tooth crown, cusp a1 might have crown pattern in all details. Only the top of main been able to travel, with the cusps of row b, cusp a1 exhibits faint traces of wear. along the central basin of the opposing upper As mentioned above, the crown pattern of the molar (Fig. 6B), suggesting a functional shift of Tendaguru specimens resembles more closely main cusp a1 from a cutting to a crushing and that of Thomasia than the remaining haramiyid grinding action. genera, suggesting that occlusion in Staffia might The model presented here was criticised by P. have been also similar to that of Thomasia. Butler (pers. comm. of July 4, 2001), who pro- In Thonznsia, occlusal movements were mainly posed that the large anterior cusp on the ‘m3’ restricted to a longitudinal (palinal) direction due interpreted here as the a1 cusp (Figs4/1A-lE, to the specific cusp configuration of both lower 5/1A-lE) might represent cusp b2 (sensu Butler and upper cheek teeth (Butler & MacIntyre 2000) which, during occlusion, follows the rest of 1994). According to Butler (2000), labial cusp the b row along the central basin of the oppos- row b of the lower molars of Thomasia occluded ing upper molar, whilst cusps a1 and a2 pass lin- in the central basin between rows A and B of the gual to the upper B row. Further, Butler suggests upper molars, and lingual row B of the upper that the link between cusp a1 and a2 on the ‘m3‘ molars occluded in the central basin between is the “saddle”, and that the large anterior cusp cusp rows a and b of the lower molars (Fig. 6A). of the type specimen (Figs 4/2A-2E, .5/2A-2E) Consequently, transverse masticatory movements would have to be identified as cusp b2 (sensu could not occur when the cheek teeth were in Butler 2000) or b2 fused with al, in that it is a occlusion (Butler & MacIntyre 1994). crushing-grinding cusp and not a shearing cusp The distinct, straight groove in the central ba- like a1 of Thomasia. If the large anterior cusp is sin of the ‘M2’ of Staffin is interpreted here as really b2 (sensu Butler 2000), the suggested func- strong evidence of wear owing to longitudinal tional change discussed above would not be ne- occlusal (palinal) movements, with cusp row B cessary. fitting into the central basin of the opposing low- My putative identification of the large anterior er tooth and cusp row b fitting into the central cusp of the lower molariforms in Staffia is pri- basin of the opposing upper molar (Fig. 6B), as marily based on observations of features of the previously postulated for Thomasia by Butler & type specimen (Fig. 5/2A-2E), in particular, the MacIntyre (1994) and Butler (2000). presence of a low rounded ridge connecting cusp However, there are some striking differences b2 (sensu Butler 2000) with principal cusp a1 in the dental pattern of Staffia. In particular, the that might have resulted from fusion of cusp a1 central position of the main cusp a1 which is lo- with bl (Heinrich 1999). This “saddle” is posi- cated at the front ot’ the cheek tooth crown. This tioned in the anterolabial portion of the tooth suggests that the occlusion pattern in the Tenda- crown and not in the anterolingual region which guru haramiyid must have been somewhat differ- is dominated by the main notch between cusps ent from that in Thomasia, where the main cusp a1 and a2. A similar condition is found in ‘m3’ a1 is only displaced medially on the supposed in that cusp bl is drawn out to a rounded ridge- lower posterior premolar (tooth groop I1 sensu like structure (Fig. .5/1A), more likely to be a Sigogneau-Russell 1989). “saddle” than that between cusps a1 and a2. 252 Heinrich. W.-D., New records of Jurassic mammals

A Thomasia B Staffia

Fig. 6. Diagramrnaiic represcntation of putati\ c molar occlusal relations for 77iorrmio (A) and Srrrffzn (B). The tccth are >hewn in cronn vi:\\. To emphasize the relative po3itioriF of main cusp a! during the occlusal movcrnents, the drawings of a 1oueI molar are si pcrimposcd on di-a\ving of upper molars (stippled). Lingual to the left, mesial above. 'The diagram shows the relative positicn of the lo\vcr molars in relation to t\vo opposing upper molars at the beginning (left) and at the end of tiic occlusal powcr stroke (right). During the occlusal moven1ents. thc lo\\ cr molar moved postcriorly (arrows). In Tlmrznsia, cusp row n of the l*.)\vermolars closes with main cusp a1 lingnlly to cuq7 row B of the opposing upper molar. In Staffin, main cuyp a1 of thc lo~ei-molars occluded with ro\\ h in the central hasin between row A and B oE the opposing upper molar, bccause of its central position at the front of the tooth crown. For further explanations see text. Fig. A modificd l'roni Butler (2000).

P. Butler (pers. comm. of July 3. 2001) but also corroded. due to transport and sorting doubted that there was ;i transitional condition prior to burial. wherein cusp $11 "jumped" across the upper B The new allotherian records are interpreted as row. 1 agree aid P. Butler's objection is impor- representing a single haramiyid species and re- tant as it migl-t indicate that Thonirrsiri did not ferred to Srriffia aenigmatica, which was prc- give rise to S,ciffici. More likely is that Sroffia viously known only from an isolated lower cheek evolved from E lineage that originally had lower tooth of Tendaguru Site dy (Heinrich 1999a). molariforms w th main cusp a1 in the anterior The dental morphology of Sta,ffia is compared central positioc. Better preserved and more com- with that of the known haramiyid genera and plete material is needed before definite conclu- shows some similarities with Thornasia. How- sions regarding occlusion in Srrrffio can be made. ever. Stoffia displays several features, that clearly separate it from Thonzasia and the other haramiyid genera. A conspicous difference in the Conclusions preserved lower cheek teeth of Staffiri is the ccn- tral position of principal cusp a1 at the front of Two non-mu1 Lituberculate allotherian cheek the tooth crown and the presence of a large teeth, tentatively identified as a lower posterior main notch between cusps a1 and a2. The upper molar ('m3') and an intermediate upper molar cheek teeth of Staffio differ from Thomasin and ('M2'). are reported from the Upper Jurassic of other haramiyids, as far as can bc judged from Tendaguru. Taiizania. East Africa. Both records an imperfectly preserved upper molar, by the were obtained from matrix of bone bed Wj of presence of four A and four B cusps, and the the Middle Saurian Bed at Tendaguru Site dy strong anterolabial cusped cingular ridge. This through the q'plication of the acid technique. ridge broadens the width of the tooth crown con- Unfortunately, the 'M2' is not only heavily worn siderably and increases the number of cusps. A Mitt. Mus. Nat.kd. Bcrl., Geowiss. Reihe 4 (2001) 253

comparison of dental dimensions of Staffia with mammals, even though a land connection (e.g.. those in Thomasia shows no substantial differ- Smith et al. 1994, Galonka 2000) and limited fau- ences, except for the ‘m3’ that seems to be dis- nal interchange of land vertebrates between tinctly smaller in the Tendaguru taxon. Africa and Laurasia occurred sometime during The fragmentary nature of the Tendaguru spe- the Late Jurassic (e.g., Galton 1977, 1982; Sigog- cimens does not permit definite conclusions con- neau-Russell 1991a, b; Kielan-Jaworowska 1992. cerning occlusion in Staffia. In contrast to Tho- Goodwin et al. 1999). However, multituberculate mnsia, the main cusp a1 in Staffia might have mammals have been reported from the Lower been able to occlude in the central basin of the Cretaceous of mainland Africa (Sigogneau-Rus- opposing upper molar, indicating a functional sell 1991a), thus the absence of these allother- shift of this cusp mainly from a cutting to a ians in the Tendaguru fauna may simply reflect crushing and grinding action. local ecological conditions. Owing to the differences described above, The presence of calcretes in the mammal-bear- Staffia may represent a new lineage of hara- ing strata of Tendaguru indicate semi-arid to arid miyids and might belong to a new family of climatic conditions, interrupted by seasonal rain- these non-multituberculate allotherians, however, falls when the sediments were deposited (Hein- erecting new taxa above the genus level should rich et al. 2001). Therefore, the Tendaguru hara- await the discovery of more complete material. miyid Staffia aenigmatica and the associated Until the discovery of Staffia aenigmatica in terrestrial vertebrate assemblage is presumed to 1999, haramiyids were unknown outside Laura- have existed in a distinct East African faunal sia. The findings from Tendaguru are the only province under a dry climate with seasonally known records of haramiyids from the Late Jur- rainy intervals, allowing the persistence of al- assic and from Gondwana. Staffia aenigmatica lotherian clades, such as the haramiyids. from the Upper Jurassic of Tendaguru is younger than any other haramiyid yet discovered, and it has, like other haramiyids, no known descen- Acknowledgements dents. The origin and phylogenetic relationships I am very grateful to Prof. Dr. P. M. Butler. University of of Staffia are poorly understood. London, and Prof. Dr. W. A. Clemens, Museum of Paleontol- In comparison with the diverse dinosaur as- ogy, Berkeley, for the opportunity to demonstrate and dis- semblage, the knowledge of mammals from the cuss the Tendaguru mammals during the International Sym- posium “Origin and Evolutionary Transformation of

Upper Jurassic of Tendaguru is still limited. Mammals ~ Using biological signals in understanding earth However, the recovery of Brancatherirlum tenda- history” held in May 2000 in Berlin. I am most indebted to gurense (Dietrich 1927, Simpson 1928, Heinrich Prof. Dr. I? M. Butler, University of London. Dr. D. Sigop- neau-Russell, Museum National d’Histoire Naturelle Parib. 1991), Tendagurodon janenschi, Tendaguruthe- the late Prof. Dr. B. Krebs (formerly Freie Ilniversitat Ber- riiun dietrichi (Heinrich 1998, 1999a), and, in lin), and Dr. Th. Martin (Freie Universitat Berlin) for helpful particular, the unexpected discovery of Staff ae- comments. Prof. E Jenkins, Harvard University Cambridge. is ia thanked for slides of Hartmziyuviu and remarks on the hara- nigrnatica, reveal that the associated mammalian miyid specimens from Tendaguru. My sincere thanks are due assemblage appears to have been correspond- to Dr. D. Unwin, Museum fur Naturkunde der Humboldt- ingly diverse, even though mammals are among Universitat zu Berlin, Institut fur Palaontologie, for criticism on the manuscript and improvements to the English. My sir]- the rarest terrestrial vertebrates in the Tenda- cere thanks are extended to Mr. J. Mendau. Museum fiir guru local fauna. Naturkunde der Humboldt-Universitat zu Berlin, Institut fur The mammalian assemblage from the Upper Palaontologie, for drawings and to Mrs. E. Siebert. from the same institution, for the camera lucida drawings in Figure 5. Jurassic of Tendaguru shows some peculiarities, Financial support from the Deutsche Forschungsgcmeinschaft suggesting the existence of a distinct East Afri- (He 27.5711-3) is gratefully acknowledged. can faunal province (Heinrich 1999a). For exam- ple, the Tendaguru local fauna lacks multituber- References culate mammals, which are common elements in Late Jurassic faunas of the Northern Hemi- Aitken, W. G. 1961. Geology and Palaeontology of the Juras- sphere (e.g., Clemens & Kielan-Jaworowska sic and Cretaceous of Southern Tanganyika. - Bulletin of 1979, Kielan-Jaworowska & Ensom 1992, Kielan- the Geological Survey of Tanganyika 31: 1 - 144. Branca, W. 1914: Kurzer Bericht uber die von Dr. Reek er- Jaworowska & Hurum ZOOl), but it contains har- zielten Ergebnisse im vierten Grabungsjaht-e 1912. - Ar- amiyids, which, in turn, are so far unrecorded in chiv fur Biontologie 3 (1): 59-63, the Upper Jurassic of Laurasia. If collecting bias - 1915. Einige Bctrachtungen uber die ailtesten SBuger der Trias- und Liaszeit. - Abhandlungen der Koniplich Prcu- can be excluded, this absence may indicate bar- Bischen Akademie dcr Wissenschalten. Physikalisch-Ma- riers (filters) to the dispersal of multituberculate thematische Klasse 1915 (5): 1-77. 254 Heinrich. W.-Il.? New records of Jurassic mamnials

Broschinski. A. 15W. Ein tilier (Scincomopha. Parama- - 199s. Late Jurassic mammals from Tendaguru, Ximania. cellodidae) iiut den1 0 Jura \.[in 'kndaguru (Tansa- East Africa. - Journal of Mammalian Evolution 5 (4): nia). - Mittei'ungen aus den1 Museum fiir Natui-hunde 269-290. Berlin. Geo~vi~ienscliaftliclicKeihe 2: 155- 1%. - 19Wi. First haraniiyid (Mammalia. Allotheria) from the Butler. P. M. 2000 Re\ie\v of he early allotherian mammals. hlesomic of Gondwana. - Mitteilungen aus dcm Mu- - Acta Palaeo itologica I'olonica 45 (4): .; 17-.3.17. semi fur Naturkunde in Berlin. Geowissenschaftliche Butler. P. M. bt Maclnt!re. G. T. 1994. Re\.ie\\ of the British Reihe 2: 159-170. Haramiyidae ( ? Mammalia. Allotheria). their molar oc- - 19L)9h.The taphonomy of dinosaurs from the Upper JLIMS- clusion and relationships. - Philosophical Transaction\ of sic of Tendapru. Tanzania (East Africa), based on field the Royal Soci:ty of London 345: 43.3-45S. sketches of the German Tcndaguru expedition Cleniens. W. A. 1'180. Rhaeto-Lias\ic mamnials from S\\it- (l9W-101~~).- Mitteilungen aus dem Museum fur Nat- land and West-Gernian!. - Zitteliana 5: 51-91, urkunde in Berlin. Geowissenschaftliche Rcihc 2: 25-61. Clemcns. W. A. 6. Kirlan-Ja\soro~\ska.Z. 1979. Muhituber- flcini-ich. W-D.. Bussort. R.. Aberhan. M., Hampe. 0..Kapi- culata. Ill Lillc gra\en. J. A,. Kielan-Ja\\oro\\~ka.Z. and lima S.. Schrank. E.. Schultka. St., Maicr,G., Msaky, E., Clcmens. W. 4. (eds). bfeso7oic hlanimals. The Fir\t Sames. B. Chami. R. 2001 . The Gernian-Tai7anian Ten- T\vo-thirds of :\laninialian History: 9% l4<). I.ni\ ersit! of dag~iruEspedition 2000. - Mitteilungen aus dem Mu- California Prcsy. Berkely. wum tur Naturkunde in Berlin. Geowissenschaftliche DacquC. E. A Krinhcl. E. IW). Jura und Kreide iii Ostafri- Reihe 4: 223-237. ka. - Neues .:alirhuch fur Mincralogit.. Geololopir und Hennig. E.. 1914. Beitriige zur Geologie und Stratigraphie Paldontologic. Beil-Bd. 28: I90-2.~2. Deutscli-C)stafrikas. - I. Geologisch-stratigraphisclie E3e- Delsate. D. 1995. Unc noui elk dent d'Hai-anii!-idac ( 7710- obachtunrgen im Kiistcngebiete des sudlichen Deutsch- /?imiuii.o//r

Parrington, F. R. 1947. On a collection of Rhaetic mammal- Sigogneau-Russell, D. 1983. Nouveaux taxons de manimifk- ian teeth. - Proceedings of the Zoological Society of res rhetiens. - Acta Palaeontologica Polonica 28: London 116 (314): 707-728. 233 -249. Peyer, B. 1956. Uber Zahne von Haramiyiden, von Tricono- - 1989. Haramiyidae (Mammalia, Allotheria) en provenance donten und von wahrscheinlich synapsiden Reptilien aus du Trias superieur de Lorraine (France). - Palaeontogra- dem Rhat von Hallau, Kt. Schaffhausen, Schweiz. - phica, Abt. A 206: 137-198. Schweizerische Palaontologische Abhhandlungen 72: - 1991a. First evidence of Multituberculata (Mamnialia) in 1-72. the Mesozoic of Africa. - Ncues Jahrbuch fur Gcologie Plicninger. W. H. von. 1847. Abbildungen von Zahnen aus und Palaontologie, Mh. 1991 (2): 119-125. der oberen Grenzbreccic des Keupers bei Degerloch und - 1991b. Dkcouverte du premier Mammifkre tribosphknique Steinenbronn. - Jahreshefte des Vereins fur vaterlandi- du Mksozoique africain. - Comptes Rendus de I' AcadC- schc Naturkunde in Wurttemberg 2: 164-167. mie des Sciences de Paris. 11 13: 405-414. Russell, D., Btland, P. & McIntosh, J. S. 1980. Paleoecology Sigogneau-Russell, D. Frank, R. M. & HemmerlC, J. IY86. A of the dinosaurs of Tendaguru (Tanzania). - MCmoirs de new family of mammals from the lower part of the la SociCtC Geologique de France, N. S. 139: 169-175. French Rhaetic. In Padian, K. (ed.). The Beginning of the Schrank, E. 1999. Palynology of the dinosaur beds of Tenda- Age of Dinosaurs. Faunal Change Across the Triassic-Ju- guru (Tanzania) - preliminary results. - Mitteilungen rassic Boundary: 99-108, Cambridge University Press. aus dem Museum fur Naturkunde Berlin, Geowissen- Cambridge. schaftliche Reihe 2: 171-183. Sigogneau-Russell, D. & Hahn, G. 1994. Late Triassic micro- - 2000. Age and biogeographic links of palynomorph assem- vertebrates from central Europe. In Fraser. N. & Sues. blages from dinosaur beds of Tendaguru (Late Jurassic, H.-D. (eds). In the Shadow of the Dinosaurs. Early Meso- Tanzania). Actes du 4Cme Symposium de Palynologie zoic Tetrapods: 197-213. Cambridge University Press. africaine (Sousse, Tunesie) 25-30 avril 1999, Geo-Eco- Cambridge. Trop, Numtro SpCcial 22: 141-146. Simpson, G. G. 1928. Mesozoic Mammalia. XI. Bmncnrhenr- Schudack, M. 1999. Some charophytes from the Middle Di- lum tendagurense Dietrich. - American Journal of Sci- nosaur Member of the Tendaguru Formation (Upper Ju- ence 15: 303-308. rassic of Tanzania). - Mitteilungen aus dem Museum fur Smith, A. G., Smith, D. G., Funel B. M. 1994. Atlas of Meso- Naturkunde Berlin, Geowissenschaftliche Reihe 2: zoic and Cenozoic coastlines. 99 pp, Cambridge Universi- 201 -205. ty Press, Cambridge. Schudack, M. E., Schudack, U. 2001. Ostracods from the Unwin, D. M. & Heinrich, W.-D. 1999. On a pterosaur jaw Middle Dinosaur Member of the Tendaguru Formation from the Upper Jurassic of Tendaguru, Tanzania. - Mit- (Upper Jurassic of Tanzania). - Neues Jahrbuch fur Geo- teilungen aus dem Museum fur Naturkunde in Berlin. logic und Palaontologie, Mh. 2001: (in print). Geowissenschaftliche Reihe 2: 121-134.