A new Cretaceous lungfish (Dipnoi: Ceratodontidae) from the Rukwa Rift Basin, Tanzania

Abstract

'Ceratodontid' lungfishes have a wide Gondwanan distribution during the Mesozoic, and are well-known from a variety of Cretaceous sites in northern and western Africa. Despite this relatively broad occurrence, significant gaps remain in our knowledge of Mesozoic African lungfish palaeodiversity and palaeogeography, particularly from subequatorial Africa. Ongoing field research in the Cretaceous Galula Formation (Red Sandstone Group), which outcrops in the Rukwa Rift Basin (a segment of the greater East Africa Rift System) of southwestern Tanzania, has led to the discovery of a diverse vertebrate fauna, including a well-preserved lungfish toothplate. This specimen is described here as a new taxon, Lupaceratodus useviaensis gen. et sp. nov., on the basis of its unique combination of morphological features relative to other 'ceratodontids.' L. useviaensis represents the first Cretaceous record of a 'ceratodontid' lungfish from Tanzania, and more broadly from the southwestern portion of the East African Rift System. The new Tanzanian form adds further diversity and a new datum to the evolutionary history of lungfishes in Africa, and it suggests possible regional differentiation between the Cretaceous fishes of East Africa and the better-known fish faunas of the period from northern and western Africa, perhaps related to the Cretaceous establishment of the Trans-Sahara Seaway.

Full text

A new Cretaceous lungfish (Dipnoi: Ceratodontidae)
from the Rukwa Rift Basin, Tanzania
Michael D. Gottfried
Geological Sciences and Museum, Michigan State University, East Lansing, Michigan 48824, U.S.A.
E-mail: gottfrie@msu.edu
Nancy J. Stevens
Department of Biomedical Sciences, Ohio University, Athens, Ohio 45701, U.S.A.
E-mail: stevensn@ohio.edu
Eric M. Roberts
Department of Physical Sciences, Southern Utah University, Cedar City, Utah 84720, U.S.A.
E-mail: robertse@suu.edu
Patrick M. O’Connor
Department of Biomedical Sciences, Ohio University, Athens, Ohio 45701, U.S.A.
E-mail: oconnorp@ohio.edu
&
Remigius Chami
Tanzania Division of Antiquities, Box 2280, Dar es Salaam, Tanzania
(with 2 figures)
Received 27 July 2009. Accepted 14 September 2009
‘Ceratodontid’ lungfishes have a wide Gondwanan distribution during the Mesozoic, and are well-known
from a variety of Cretaceous sites in northern and western Africa. Despite this relatively broad occurrence,
significant gaps remain in our knowledge of Mesozoic African lungfish palaeodiversity and palaeogeography,
particularly from subequatorial Africa. Ongoing field research in the Cretaceous Galula Formation (Red
Sandstone Group), which outcrops in the Rukwa Rift Basin (a segment of the greater East Africa Rift System) of
southwestern Tanzania, has led to the discovery of a diverse vertebrate fauna, including a well-preserved
lungfish toothplate. This specimen is described here as a new taxon, Lupaceratodus useviaensis gen. et sp. nov.,
on the basis of its unique combination of morphological features relative to other ‘ceratodontids.’ L. useviaensis
represents the first Cretaceous record of a ‘ceratodontid’ lungfish from Tanzania, and more broadly from the
southwestern portion of the East African Rift System. The new Tanzanian form adds further diversity and a new
datum to the evolutionary history of lungfishes in Africa, and it suggests possible regional differentiation
between the Cretaceous fishes of East Africa and the better-known fish faunas of the period from northern and
western Africa, perhaps related to the Cretaceous establishment of the Trans-Sahara Seaway.
Key words:Lupaceratodus, lungfish, Ceratodontidae, Cretaceous, Rukwa Rift Basin, Tanzania.
INTRODUCTION
African Mesozoic ‘ceratodontid’ lungfish records are concen-
trated in the northern and northwestern parts of the conti-
nent (e.g. Haug 1905; Peyer 1925; Weiler 1930; Arambourg
& Joleaud 1943; Tabaste 1963; Martin 1981, 1984; Churcher
1995; Churcher & De Iuliis 2001), reflective of a broader
pattern in the African Cretaceous terrestrial and freshwater
vertebrate fossil record. Apart from significant finds made in
South Africa (e.g. de Klerk et al. 2000) and Malawi (Jacobs
et al. 1990), our working knowledge of subequatorial African
Cretaceous vertebrates is, in general, notoriously poor – a
situation that we have previously referred to as the ‘African
Gap’ (e.g. O’Connor et al. 2006). With this in mind, we began
exploring the Rukwa Rift Basin (RRB) of southwestern Tan-
zania in 2002, and have conducted eight field seasons to
date, recovering both Cretaceous and Palaeogene verte-
brates, invertebrates, and plants from ca. 60 localities
(Stevens et al. 2008).
The Cretaceous fauna from the RRB is beginning to close
at least some portions of the aforementioned gap (Krause
et al. 2003; Gottfried et al. 2004; O’Connor et al. 2005, 2006),
and has the potential to provide data that are useful in assess-
ing competing Gondwanan biogeographical hypotheses. For
example, the ‘Africa First’ model (sensu Sampson et al.
1998; Krause et al. 1999, 2006) postulates that the (apparent)
absence of several Gondwanan vertebrate clades in the
CONTENTS
Abstract · · · · · · · · · · · · · · · · · · · · 31
Introduction · · · · · · · · · · · · · · · · · 31
Systematic palaeontology · · · · · 32
Lupaceratodus gen. nov. · · · · · · · · · · 32
Lupaceratodus useviaensis sp. nov. 32
Discussion· · · · · · · · · · · · · · · · · · · · · · 34
Acknowledgements · · · · · · · · · 35
References · · · · · · · · · · · · · · · · 35

African Cretaceous is the result of Africa being physically
isolated from Indo-Madagascar, Antarctica, and South
America and thus not part of a broadly distributed Gondwanan
Cretaceous biota. The alternative ‘Pan-Gondwana’ model
(sensu Sereno et al. 2004) predicts that certain clades will be
present in the Cretaceous of Africa and other Gondwanan
continents due to persistent, if somewhat intermittent,
physical connections during the Cretaceous. More regionally
focused biogeographical models (e.g. O’Connor et al. 2006)
suggest possible provinciality between Saharan and
subequatorial freshwater/terrestrial faunas, which could
have resulted in part due to the hypothesized trans-Saharan
seaway (Gebhardt 1999) acting as a marine barrier separating
northwestern Africa from more southern and eastern regions
of the continent. The primary challenge in testing these
biogeographical models remains the lack of Cretaceous fossil
data from subequatorial Africa.
The specimen reported here from the Cretaceous Galula
Formation (Red Sandstone Group) in the RRB of western
Tanzania (Fig. 1) establishes the presence of Mesozoic
lungfish for the first time in the southwestern portion of the
East African Rift System, and it expands the palaeodiversity
of ceratodontid lungfishes in Africa. More broadly, it under-
scores the palaeontological potential of the Tanzanian RRB
deposits in preserving diverse, and potentially biogeo-
graphically informative, Cretaceous freshwater and terres-
trial vertebrates from subequatorial Africa.
SYSTEMATIC PALAEONTOLOGY
SUBCLASS DIPNOI Müller, 1844
ORDER CERATODONTIFORMES Berg, 1940
Family CERATODONTIDAE Gill, 1872
Lupaceratodus
gen. nov.
Fig. 2
Type species
Lupaceratodus useviaensis
sp. nov.
Fig. 2
Derivation of name
Generic name refers to the Lupa Bounding Fault, a promi-
nent geological feature in the Rukwa Rift Basin, combined
with ‘ceratodus’ in reference to its affinities with other
32 African Natural History, Volume 5, 2009
Fig. 1. General geological setting and outcrop area of the Red Sandstone Group in the Rukwa Rift Basin, southwestern Tanzania. The specimen
described here was collected near Usevia (outcrop area 4), in the northwestern part of the Rukwa Rift Basin. Inset map shows Tanzania (shaded),
study area depicted is indicated by the line denoting the southwestern corner of Tanzania.

ceratodontid lungfishes. Specific epithet refers to the town
of Usevia, situated near the type locality.
Holotype
Ruka Rift Basin Project (RRBP) 04289 (National Museum
of Tanzania), a moderately worn, complete upper left
(pterygoid) toothplate with a portion of the pterygoid bone
attached (Fig. 2).
Locality and geological setting
RRBP 04289 was collected on 30 June 2004 (by EMR) near
the town of Usevia, in the Rukwa District of western Tanzania,
at ca. 7°6’south latitude, 31°10’east longitude (see Fig. 1;
precise locality coordinates are on file with the authors). The
specimen was recovered from a sandstone outcrop designated
by us as locality TZ-22 and assigned to the Galula Formation
(Roberts et al., in press) (formerly ‘Unit I’ sensu Roberts et
al. 2004) of the Red Sandstone Group. The Galula Forma-
tion is Cretaceous in age based on its temporally distinctive
vertebrate fauna, which includes sauropod and theropod
dinosaurs, megaloolithid dinosaur eggshell, and osteo-
glossomorph fishes (Gottfried et al. 2004; Roberts et al.
2004; O’Connor et al. 2005, 2006). Detrital zircon analysis
provides some independent support for this by demonstrat-
ing that the depositional age must be Late Jurassic or youn-
ger (Roberts et al. 2007), and this age interpretation is con-
sistent with facies relationships and our geological mapping
of the field area. Taken together these data indicate a Creta-
ceous age assignment, and further suggest that the majority
if not all deposition of the Galula Formation transpired in
the lower to middle Cretaceous. Further refinement of
the age of the Galula Formation is the subject of ongoing
research.
The Galula Formation, consisting of red sandstones and
mudstones, accumulated in a freshwater fluvial/floodplain
continental rift setting. Deposition of these sediments was
along a northwest-flowing river system oriented parallel to
the axis of the RRB. The Galula Formation exposures near
Gottfried
et al.
: A new Cretaceous lungfish from the Rukwa Rift Basin, Tanzania 33
Fig. 2.
Lupaceratodus useviaensis
(gen. et sp. nov.) holotype – upper left toothplate and a portion of the pterygoid element (RRBP 04289), from
the Cretaceous Galula Formation (Red Sandstone Group), Rukwa region, southwestern Tanzania. Top image in occlusal view, with anterior to the
left and the lingual margin to the top; numbers 1–6 indicate ridges. Lower image shows holotype specimen rotated into lingual view.

Usevia are situated towards the northwestern end of the
basin (see Fig. 1), which is a roughly 300 km long by 50 km
wide northwest–southeast trending segment of the south-
western branch of the East African Rift System (EARS).
The overall Cretaceous fauna recovered to date includes
osteichthyans (osteoglossomorphs, fragments of an as yet
unidentified teleost, and a ‘ceratodontid’ lungfish (this
report)), turtles, crocodyliforms, at least four taxa of
saurischian dinosaurs, megaloolithid dinosaur eggshell, and
a (probable gondwanantherian) mammal (see Krause et al.
2003; Gottfried et al. 2004; O’Connor et al. 2005, 2006).
Diagnosis
Distinguished from other ‘ceratodontid’ lungfish upper
toothplates by the following unique combination of features:
Markedly slender (overall length:maximum width ratio
2.6:1) in proportion; first (anteriormost) ridge diverges at
an angle of ca. 55 degrees from the next ridge and has
distinct bend halfway along its length; second ridge slightly
shorter but projecting further lingually than first; second
and third ridges arcuate, fourth and fifth ridges sinusoidal
along ridge crests; sixth ridge much shorter (no more than
half the length of preceding ridges); labial margin of
toothplate essentially straight (rather than curved) from
second through sixth ridges and not strongly projecting at
base of second ridge as in other ceratodontids; third through
sixth ridges do not converge along labial margin.
Description
Lupaceratodus useviaensis (RRBP 04289, Fig. 2) is repre-
sented by a well-preserved, relatively small and slender,
upper left toothplate, measuring 16.0 mm in length by
6.2 mm in (maximum) breadth (measured across the
occlusal surface of the toothplate and including the crests of
the ridges). As preserved, it is somewhat loosely ankylosed to
the underlying preserved portion of the pterygoid element
(Fig. 2). Five ridges (following the terminology of Churcher
& De Iuliis 2001) are well-preserved; a notably shorter sixth
posteriormost ridge (sometimes referred to as the posterior
heel) is present but incompletely preserved. The anterior-
most ridge diverges at an acute angle of ca. 55 degrees from
the next ridge, and has a distinct bend approximately half-
way along its length; the more posterior ridges are sub-
parallel to one another, with the fourth and fifth sinusoidal
along their crests. The four more posterior ridges do not
converge along the labial margin of the toothplate as is
common in many ‘ceratodontid’ toothplates, which thus
have a ‘palmate’ appearance that is lacking in L. useviaensis.
The angled anteriormost ridge is the longest at ca. 6.7 mm,
the other ridges (with the exception of the much shorter
sixth ridge) range from 6.3 to 5.2 mm in length and decrease
rather evenly in length from anterior to posterior. The crest
of the second ridge projects further lingually than does the
first. The crests in general are well-defined, with relatively
deep intervening sulci. The toothplate does not exhibit any
resorption on its occlusal surface, which (along with its small
size) suggests that it may be at a subadult stage of
development (Kemp 1977). Closely spaced simple round to
oval punctuations (~0.3 mm in diameter) are most apparent
along the ridges and crests, and along the lingual and buccal
margins, and are less distinctly developed in the deeper
parts of the furrows between ridges.
Lupaceratodus exhibits similarities to upper toothplates
of the extant Australian species Neoceratodus forsteri
(Kemp & Molnar 1981; Kemp 1997), including the presence
of a longer anteriormost divergent ridge followed by five
ridges and a short posteriormost sixth ridge (= posterior
heel). However, unlike Neoceratodus, Lupaceratodus has
subparallel and curved (rather than straight) posterior
ridges, and it differs in overall proportions. Toothplates that
have been assigned to the problematic taxon ‘Ceratodus’ are
broader in proportion than Lupaceratodus, and characteris-
tically bear fewer (4–5) and more radially divergent non-
parallel ridges, resulting in an overall palmate appearance
(see, e.g., C. humei in Churcher & De Iuliis 2001). The new
taxon does bear some similarities in its relatively slender
proportions and overall appearance to Retodus (formerly
Ceratodus)tuberculatus (Churcher et al. 2006) from the
Cretaceous of Egypt, Algeria, and Niger, but R. tuberculatus
has four or perhaps five ridges, rather than six as in
Lupaceratodus, and each of the ridges in Retodus has a
similar posteriorly angled orientation, different from the
situation in Lupaceratodus in which the anteriormost ridge
is divergent from the others, and not posteriorly oriented.
DISCUSSION
The phylogenetic interrelationships and taxonomy of
Mesozoic lungfishes remain problematic, and a comprehen-
sive synthesis of the group’s diversity and palaeogeographi-
cal distribution during the Mesozoic has yet to be achieved.
Many taxa are based on isolated toothplates, with more
complete material relatively rare, a situation that relates to
poor mineralization of the skeleton, but robust development
of petrodentine-reinforced toothplates in post-Palaeozoic
lungfishes (Cavin et al. 2007). As a result, many of the species
placed in the family Ceratodontidae have been assigned to
the ‘catch-all’ genus Ceratodus, and in some cases species
originally assigned to Ceratodus have more recently been
placed in Neoceratodus (e.g. Churcher & De Iuliis 2001),
Atlantoceratodus (Cione et al. 2007), or Retodus (Churcher
et al. 2006), illustrating the current instability in the
generic-level taxonomy of this group. Rather than placing
the morphologically distinctive toothplate described here
into Ceratodus, which is in need of a thorough revision, we
have erected the new genus and species, Lupaceratodus
useviaensis, to accommodate its distinctive and diagnosable
combination of morphological features.
The presence of Lupaceratodus in western Tanzania adds
a noteworthy new datum to dipnoan history in the Mesozoic
of Africa. Its occurrence is not startling given the long
geological time span and wide geographical distribution of
‘ceratodontid’ records in general (Obruchev 1964; Woodward
1906; Marshall 1986), including on Gondwanan continents.
Lupaceratodus comprises another example of a morphologi-
cally relatively conservative lungfish in the African Mesozoic,
conforming to the general ‘ceratodontid’ morphotype that
was ultimately supplanted in the Cenozoic of Africa by the
surviving lepidosirenid genus Protopterus, with its more
specialized and simplified toothplate morphology.
Ceratodontid lungfishes are a geologically ancient and
geographically widespread assemblage, the origin of which
predates Gondwanan fragmentation (see e.g. Apesteguía
34 African Natural History, Volume 5, 2009

et al. 2007). The new taxon from Tanzania may shed some
light on regional differentiation of the clade, in that
Lupaceratodus is taxonomically distinct from the more typi-
cal ‘ceratodontids’ recovered from northern and western
Africa. A possible abiotic mechanism that may account for at
least some of this geographical separation, as reflected by
the presence of Lupaceratodus in Tanzania, is the initial
transgression of the Trans-Saharan seaway that divided the
northwestern part of continental Africa from the rest of the
continent during the Cretaceous (Gebhardt 1999), perhaps
as early as pre-Albian (Al-Khashab 2000). Additional material
is required to discern whether other components of the
freshwater fish fauna in the southwestern portion of the
EARS are also distinctive enough from those of northern
and western Africa to posit intra-African biogeographical
provincialism. The discovery of Lupaceratodus does under-
score the continuing potential of the Rukwa Rift Basin and
the Galula Formation of southwestern Tanzania to provide
new and novel palaeontological data from the Cretaceous of
subequatorial Africa.
ACKNOWLEDGEMENTS
We thank our collaborators in the Department of Geology
at the University of Dar es Salaam, particularly E. Mbede,
S. Ngasala, and our late friend and colleague S. Kapilima; the
Tanzanian Division of Antiquities (especially D. Kamamba
and C. Msuya); and the Tanzania Commission for Science
and Technology. We also thank E. Johansen for logistical
assistance, and C.S. Churcher for helpful comments. The
paper was improved by comments from B. Grandstaff,
L. Cavin, and A. Cione. Z. Johanson and M. Richter at the
Natural History Museum (London) provided access to the
NHM collections to examine lungfish material. Finally, we
thank the following funding sources: the National Science
Foundation (EAR-0617561) of the U.S.A., National Geo-
graphic Society Committee for Research and Exploration,
the Office of the Vice President for Research and Graduate
Studies at Michigan State University, and the Ohio Univer-
sity Office of Research and Sponsored Programs and College
of Osteopathic Medicine.
REFERENCES
ALl-KHASAB, S.A. 2000. Lower Cretaceous ostracoda from selected
boreholes in Central Iraq. Cyris (International Ostracoda News-
letter) 18: 1–2.
APESTEGUÍA, S., AGNOLIN, F. & CLAESON, K. 2007. Review of
Cretaceous dipnoans from Argentina. Revista del Museo
Argentino de Ciencias Naturales 9: 27–40.
ARAMBOURG, C. & JOLEAUD, L. 1943. Vertébrés fossiles du
bassin du Niger. Bulletin du Service des Mines de l’Afrique
Occidentale Française 7: 27–84.
CAVIN, L., SUTEETHORN, V. BUFFETAUT, E. & TONG, H.
2007. A new Thai Mesozoic lungfish (Sarcopterygii, Dipnoi) with
an insight into post-Palaeozoic dipnoan evolution. Zoological
Journal of the Linnean Society 149: 141–177.
CHURCHER, C.S. 1995. Giant Cretaceous lungfish Neoceratodus
tuberculatus from a deltaic environment in the Qusair (= Baris)
Formation of Kharga Oasis, western desert of Egypt. Journal of
Vertebrate Paleontology 15: 845–849.
CHURCHER, C.S. & DE IULIIS, G. 2001. A new species of
Protopterus and a revision of Ceratodus humei (Dipnoi: Cerato-
dontiformes) from the Late Cretaceous Mut Formation of eastern
Dakleh Oasis, western desert of Egypt. Palaeontology 44:
305–323.
CHURCHER, C.S., DE IULIIS, D. & KLEINDIENST, M.R. 2006. A
new genus for the Dipnoan species Ceratodus tuberculatus
Tabaste, 1963. Geodiversitas 28: 635–647.
CIONE, A.L., CAVALLI, G., GOIN, S. & POIRE. D.F. 2007. Atlanto-
ceratodus, a new genus of lungfish from the upper Cretaceous of
South America and Africa. Revista del Museo de la Plata, Paleon-
tologia 10: 1–12.
DE KLERK, W.J., FORSTER, C.A. SAMPSON, S.D. CHINSAMY,
A. & ROSS, C.F. 2000. A new coelurosaurian dinosaur from the
early Cretaceous of South Africa. Journal of Vertebrate Paleontol-
ogy 20: 324–332.
GEBHARDT, H. 1999. Cenomanian to Coniacian biogeography and
migration of North and West African ostracods. Cretaceous
Research 20: 215–229.
GOTTFRIED, M.D., O’CONNOR, P.M. JACKSON, F. ROBERTS,
E.M. & CHAMI, R. 2004. Dinosaur eggshell from the Red Sand-
stone Group of Tanzania. Journal of Vertebrate Paleontology 24:
494–497.
HAUG, É. 1905. Paléontologie. Documents scientifiques de la
Mission saharienne (Mission Foureau-Lamy). Publication de la
Société de géographie, Paris: 751–832.
JACOBS, L.L., KAUFULU, Z.M. & DOWNS, W.R. 1990. The
Dinosaur Beds of northern Malawi. National Geographic
Research 6: 196–204.
KEMP, A. 1977. The pattern of tooth plate formation in the Austra-
lian lungfish Neoceratodus forsteri (Krefft). Zoological Journal of
the Linnean Society 60: 223–258.
KEMP, A. 1997. A revision of Australian Mesozoic and Cenozoic
lungfish of the Family Neoceratodontidae (Osteichthyes: Dipnoi)
with a description of four new species. Journal of Paleontology
71: 713–733.
KEMP, A. & MOLNAR, R. 1981. Neoceratodus forsteri from the
Lower Cretaceous of New South Wales, Australia. Journal of
Paleontology 55: 211–217.
KRAUSE, D.W., GOTTFRIED, M.D., O’CONNOR, P.M. &
ROBERTS, E.M. 2003. A Cretaceous mammal from Tanzania.
Acta Palaeontologica Polonica 48: 321–330.
KRAUSE, D.W., O’CONNOR, P.M., CURRY ROGERS, K.,
SAMPSON, S.D., BUCKLEY, G.A. & ROGERS, R.R. 2006. Late
Cretaceous terrestrial vertebrates from Madagascar: implica-
tions for Latin American biogeography. Annals of the Missouri
Botanical Garden 93: 178–208.
KRAUSE, D.W., ROGERS, R.R., FORSTER, C.A., HARTMAN,
J.H., BUCKLEY, G.A. & SAMPSON, S.D. 1999. The Late
Cretaceous vertebrate fauna of Madagascar: implications for
Gondwana biogeography. GSA Today 9: 1–7.
MARSHALL, C. 1986. A list of fossil and extant dipnoans. In:
BEMIS, W.E., BURGGREN, W.W. & KEMP, N. (eds) The Biology
and Evolution of Lungfishes. Journal of Morphology Supplement
1: 15–23.
MARTIN, M. 1981. Les Ceratodontiformes (Dipnoi) de Gadoufaoua
(Aptien supérieur, Niger). Bulletin Museum National Histoire
Naturelle, C: Sciences Terre (Séries 4)3: 267–283.
MARTIN, M. 1984. Révision des Arganodontidés et des Néocéroto-
dontidés (Dipnoi, Ceratodontiformes) du Crétacé africain. Neues
Jahrbuch für Geologie und Paläontologie, Abhandlungen 169:
225–260.
OBRUCHEV, D.V. 1964. Fundamentals of Paleontology. Vol. XI.
504 pp. Izdatelstvo Nauka, Moscow.
O’CONNOR, P.M., GOTTFRIED, M.D., ROBERTS, E.M.,
STEVENS, N.J. & NGASALA, S. 2005. New dinosaurs and other
vertebrate fossils from the Cretaceous Red Sandstone Group,
Rukwa Rift Basin, southwestern Tanzania. Journal of Vertebrate
Paleontology (Supplement)25: 97A.
O’CONNOR, P.M., GOTTFRIED, M.D., STEVENS, N.J.,
ROBERTS, E.M., NGASALA, S., KAPILIMA, S. & CHAMI, R..
2006. A new vertebrate fauna from the Cretaceous Red Sand-
stone Group, Rukwa Rift Basin, southwestern Tanzania. Journal
of African Earth Sciences 44: 277–288.
PEYER, B. 1925. Ergebnisse der Forschungsreisen Prof. E.
Stromers in den Wüsten Ägyptens. II. Wirbeltier-Reste der Bahar
Gottfried
et al.
: A new Cretaceous lungfish from the Rukwa Rift Basin, Tanzania 35

je-Stufe (Unterstes Cenoman): 6. Die Ceratodus-Funde.
Abhandlungen der Bayerischen Akademie Wissenschaften 30:
5–32.
ROBERTS, E.M., O’CONNOR, P.M., ARMSTRONG, R.A.,
STEVENS, N.J. & GOTTFRIED, M.D. 2007. U-PB geochronol-
ogy of detrital zircons from the Rukwa Rift Basin, Tanzania: new
data on the pre-Neogene tectonic and sedimentary evolution of
the western branch of the East African Rift System. Geological
Society of America Abstracts with Program 39: 505.
ROBERTS, E.M., O’CONNOR, P.M., GOTTFRIED, M.D.,
STEVENS, N.J., KAPILIMA, S. & NGASALA, S. 2004. Revised
stratigraphy and age of the Red Sandstone Group in the Rukwa
Rift Basin, Tanzania. Cretaceous Research 25: 749–759.
ROBERTS, E.M., O’CONNOR, P.M., STEVENS, N.J.,
GOTTFRIED, M.D., JINNAH, Z.A., NGASALA, S., CHOH, A.M.
& ARMSTRONG, R.A. In press. Sedimentology and depositional
environments of the Red Sandstone Group, Rukwa Rift Basin,
southwestern Tanzania. New insights into Cretaceous and
Paleogene terrestrial ecosystems and tectonics in sub-equatorial
Africa. Journal of African Earth Sciences.
SAMPSON, S.D., WITMER, L.M., FORSTER, C.A., KRAUSE,
D.W., O’CONNOR, P.M., DODSON, P.D. & RAVOAVY, F. 1998.
Predatory dinosaur remains from Madagascar: implications for
the Cretaceous biogeography of Gondwana. Science280:
1048–1051.
SERENO, P.C., WILSON, J.A. & CONRAD, J.L. 2004. New dino-
saurs link southern landmasses in the mid-Cretaceous. Proceed-
ings of the Royal Society of London, Series B,271: 1325–1330.
STEVENS, N.J., GOTTFRIED, M.D., ROBERTS, E.M.,
KAPILIMA, S., NGASALA, S. & O’CONNOR, P.M. 2008.
Paleontological exploration in Africa. A view from the Rukwa Rift
Basin in Tanzania. In: J.G. FLEAGLE & C.C. GILBERT (eds)
Elwyn Simons – A Search for Origins, pp. 159–183. Springer
Science, New York.
TABASTE, N. 1963. Étude des restes de poissons du Crétacé
saharien. Mélanges ichthyologiques à la mémoire d’Achille
Valenciennes. Mémoires de l’Institut Français d’Afrique Noire
68: 437–485.
WEILER, W. 1930. Beschreibung von Wirbeltier-Resten aus dem
nubischen Sandsteine Oberägyptens und aus ägyptischen
Phosphaten nebst Bemerkungen über die Geologie der
Umgegend von Mahamîd in Oberägypten. Abhandlungen der
Bayerischen Akademie Wissenschaften (Neue Folge) 7: 1–42.
WOODWARD, A.S. 1906. On a tooth of Ceratodus and a dinosaur-
ian claw from the Lower Jurassic of Victoria, Australia. Annals
and Magazine of Natural History 17: 1–3.
36 African Natural History, Volume 5, 2009