Dinosaur Distribution

Abstract

This chapter summarizes all dinosaur locations currently known worldwide. Geographic location is based on primary administrative divisions (e.g., states, departments, provinces, counties) of the countries in which dinosaur material has been found. Locations are annotated for lithostratigraphic unit (bed, formation, group, series), faunal composition, and stratigraphic horizon. The members of the fauna are present in rank order, roughly based on their phylogenetic relationships. The chapter also includes locations yielding skeletal remains and those that have produced tracks and/or trackways, as well as locations where eggs and eggshell fragments have been found.

Full text

SECTION TWO
DINOSAUR DISTRIBUTION
AND BIOLOGY

Page 516 is blank

Research on dinosaurs, whether on systematics, biogeography,
paleoecology, or functional morphology, ultimately depends
on their discovery in the field. This update of known dinosaur
occurrences reflects an incredible 55% increase in the number
of entries over those provided by Weishampel (1992), indicat-
ing the astonishing growth in the underpinnings of our research
base in just over a decade. Not only has the number of dinosaur
localities increased, but the high-latitude faunas are better
known and the “midperiod” gaps are beginning to be filled in.
As will be seen in this chapter, there have been substantial in-
creases in the number of specimens from the Mesozoic in North
America and especially in Europe. Asia follows thereafter, with
signifivcant increases in the dinosaur record for all but the Late
Triassic. Increases in the number of of dinosaur localities dis-
covered are also significant in South America and to a lesser
extent in Africa in the Late Triassic and throughout the Creta-
ceous. Australasia and Antarctica lag behind other continents
in number of dinosaur occurrences, but they too provide new
Early Jurassic and Late Cretaceous localities.
This chapter summarizes all dinosaur locations currently
known worldwide. Geographic location is based on primary
administrative divisions (e.g., states, departments, provinces,
counties) of the countries in which dinosaur material has been
found. The only exception is Antarctica, for which there are no
subdivisions. Lithostratigraphic units are generally formational.
Note that, where there is more than one locality from a given
lithostratigraphic unit within one of the administrative divisions
of a county, the localities are lumped together; hence it is ap-
propriate to consider these grouped localities under the more
general term “locations.” Because of the ever-increasing number
of dinosaur discoveries around the world, the previous system
of numbering proved too unwieldy. A new system was devised
to accommodate this new list and provide for future growth. A
locality is identified by its symbol and three numbers: the first
corresponds to the country, the second to the administrative
district within the country, and the third to the formation in
which it was found. For example a ★★labeled 3.5-2 corresponds
to the Late Cretaceous Pen Formation of Texas, United States.
Locations are annotated for (1) lithostratigraphic unit (bed,
formation, group, series, svita), (2) faunal composition, and
(3) stratigraphic horizon (to stage where possible). Faunal com-
position is given to species-level taxa where possible, based on
information presented in this volume. The members of the fauna
are present in rank order, roughly based on their phylogenetic
relationships, although the full phylogenetic hierarchy is not
provided owing to space limitations.
Finally, locations yielding skeletal remains, as well as those
that have produced tracks and or trackways, have been included
(with higher taxonomic referrals; ichnological designations are
not used). The same applies to those locations that have yielded
eggs, eggshell fragments, or both (again, no ootaxa are indicated;
nests, isolated eggs, and eggshell fragments are listed only as
“dinosaur eggs”) and to those that have yielded coprolites. Age
assignments are based, as far as can be ascertained, on the stan-
dard European stages (see the introduction to the volume), but
these are often limited to best guesses about the temporal rela-
tionship between terrestrial and marine strata. For reasons of
length, no attempt has been made to characterize the distribu-
tion of faunas within lithologic units, nor are the sedimentology,
depositional environments, or taphonomic aspects of locations
indicated, although each of these sources of information is in
its own right important to an understanding of dinosaurian
distribution.
Symbols used in the maps refer to the eight series of the
Mesozoic for which there are dinosaur remains: ★, Late Triassic;
■■, Early Jurassic; ●●, Middle Jurassic; , Late Jurassic; ●, Early
Cretaceous; ★★, Late Cretaceous; and , uncertain age. The first
map indicates where larger-scale regional maps may be found
and gives a number of more poleward locations; successive
maps provide locations elsewhere in the world.
DINOSAUR DISTRIBUTION 517
TWENTY-FOUR
Dinosaur Distribution
DAVID B. WEISHAM PEL
PAUL M. BARRETT
RODOLFO A. C ORIA
JEAN LE LOEUF F
XU XING
ZHAO XIJIN
ASHOK SAHN I
ELIZAB ETH M. P. G OMANI
CHR ISTOPHER R. NOTO

Late Triassic
Late Triassic dinosaurs are known from over a hundred loca-
tions in North America, Europe, Asia, South America, Africa,
and Australasia.
North America
Information on Late Triassic dinosaurs in North America (33 lo-
cations, an increase of about 70%) comes to us primarily through
the work of E. H. Colbert on the faunas of the southwestern
United States and D. Baird on those of the eastern seaboard.
More recently, work on the faunas of the Southwest has been
conducted by a host of research groups led by K. Carpenter,
S. Chatterjee, A. P. Hunt, L. L. Jacobs, M. G. Lockley, R. A. Long,
S. G. Lucas, P. A. Murry, K. Padian, J. M. Parrish, T. Rowe, and
R. M. Sullivan. The faunas in the eastern United States and
518 DINOSAUR DISTRIBUTION AND BIOLOGY
FIGU RE 24.1. Global index map. Regional maps are indicated by boxes labeled with bold numbers and arrows. Boxed numbers indicate more
poleward localities not included in subsequent figures. Scale =3,000 km.
1.1.
NORTHWEST TERRITORIES
,
CANADA
1. Heiberg Formation (Adams 1875;
Lydekker 1889b; Galton and Cluver 1976)
?Theropoda indet. (=Arctosaurus osborni)
Age: Carnian-Rhaetian (Douglas et al.
1963)
1.7.
NOVA SCOTIA
,
CANADA
1. Wolfville Formation (Baird and Olsen
1983; Olsen and Baird 1986; Sues et al.
1994b; Weishampel and Young 1996)
Prosauropoda indet.
Ornithischia indet.
Ornithischian tracks
Age: late Carnian (Huber et al. 1993a)
2. Blomidon Formation (Olsen et al. 1982;
Weishampel and Young 1996)
Theropod tracks
Age: Norian (Olsen et al. 1982;
Lucas and Huber 1993)
2.1.
EAST GREENLAND
,
GREENLAND
1. Fleming Fjord Formation (Jenkins et al.
1993, 1995; Gatesy et al. 1999;
Clemmensen et al. 1998)
Theropoda indet.
Theropod tracks
Prosauropoda
Plateosaurus longiceps
Age: middle Norian (Jenkins et al. 1995)
3.1.
UTAH
,
UNITED STATES
1. Chinle Formation (Lockley 1986a)
Theropod tracks
Age: late Carnian–early Norian
(Olsen et al. 1982)
3.2.
ARIZONA
,
UNITED STATES
1. Petrified Forest Formation (Jacobs 1980;
Jacobs and Mur ry 1980; Padian 1986; Lucas
et al. 1992; Hunt and Lucas 1994; Kaye
and Padian 1994; Long and Murry 1995)
Maritime Canada have been studied most recently by P. E. Olsen,
H.-D. Sues, and R. E. Weems. Finally, F. A. Jenkins and collabo-
rators recently discovered an important dinosaur fauna from the
Late Triassic of eastern Greenland that is currently under study.
For the American Southwest, the lithostratigraphic nomencla-
ture used by Lucas and co-workers is used here, although some
(Dubiel 1994) regard these units as facies of the Chinle For-
mation. Figures 24.1–24.3 illustrate these North American Late
Triassic locations.

Theropoda
Saurischia incertae sedis
Chindesaurus bryansmalli
Ceratosauria
Coelophysis bauri
Prosauropoda indet. (=?Anchisauridae
indet.)
Ornithischia indet. (including
Revueltosaurus callenderi)
Age: late Carnian–early Norian (Olsen
et al. 1982; Murry 1990)
2. Bluewater Creek Formation (Hunt and
Lucas 1994; Kaye and Padian 1994;
Heckert and Lucas 1997)
Theropoda
Ceratosauria
Camposaurus arizonensis
Ornithischia indet. (=Tecovasaurus
murryi)
Age: late Carnian (Heckert and Lucas
1997)
3.3.
NEW MEXICO
,
UNITED STATES
1. Santa Rosa Formation (Hunt and Lucas
1995b)
Theropoda indet.
Undescribed ornithischian
Age: early Carnian (Lucas 1993a)
2. Garita Creek Formation (Hunt and
Lucas 1995b)
Theropoda indet.
Age: late Carnian (Lucas 1993a)
3. Bluewater Creek Formation (Hunt
and Lucas 1993a; Lucas 1993a;
Heckert 1997; Heckert and Lucas
1997)
Theropoda indet.
Ornithischia indet. (=Tecovasaurus
murryi)
Ornithischian tracks
Age: late Carnian (Heckert and Lucas
1997)
DINOSAUR DISTRIBUTION 519
FIGU RE 24.2. Late Tr iassic to Late Cretaceous localities in eastern Canada (1) and the United States (3). Scale =600 km.

4. Petrified Forest Formation (Hunt and
Lucas 1993a; Hunt 1996; Sullivan and
Lucas 1999; Heckert et al. 2000; Heckert,
pers. comm.)
Theropoda
Ceratosauria indet. (including
Eucoelophysis baldwini,
Coelophysis bauri,
C. longicollis, C. willistoni)
Theropoda indet.
Age: late Carnian–early Norian (Hunt
and Lucas 1993a)
5. Bull Canyon Formation (Hunt 1989;
Hunt and Lucas 1989, 1993a, 1993b,
1994; Lucas et al. 1985; Long and Murry
1995)
Saurischia incertae sedis
?Chindesaurus bryansmalli
Herrerasauridae indet.
Theropoda
Ceratosauria indet.
Ornithischia indet. (including
Revueltosaurus callenderi,
Lucianosaurus wildi)
Age: early Norian (Hunt and Lucas
1993a)
6. Cooper Canyon Formation (Carpenter
and Parrish 1985; Parrish and Carpenter
1986; Carpenter 1997a; ? =Bull Canyon
Formation)
Theropoda
Ceratosauria
Gojirasaurus quayi
Theropoda incertae sedis
Shuvosaurus inexpectatus
Theropoda indet.
Age: middle Norian (Carpenter 1997a)
7. Redonda Formation (Conrad et al. 1987;
Hunt et al. 1989; Lockley et al. 2000)
Theropod tracks
Prosauropod tracks
Age: late Norian (Hunt and Lucas 1993a)
8. Sloan Canyon Formation (Conrad et al.
1987; Hunt et al. 1989; Lockley and
Hunt 1993; Lockley et al. 2000)
Theropod tracks
Prosauropod tracks
Age: late Norian (Hunt and Lucas 1993a)
9. Sheep Pen Sandstone Formation
(Lockley et al. 1993, 2000)
Theropod tracks
Prosauropod tracks
Age: late Norian (Hunt and Lucas 1993a)
10. Rock Point Formation (Colbert 1974,
1989; Welles 1984; Farlow and Lockley
1993; Hunt and Lucas 1993a, 1995b)
Theropoda
Ceratosauria
Coelophysis bauri (=Rioarribasaurus
colberti)
Theropoda indet.
?Prosauropod tracks
Undescribed ornithischian
Ornithischia indet. (including
Revueltosaurus callenderi)
Age: late Norian (Hunt and Lucas
1993a)
3.4.
COLORADO
,
UNITED STATES
1. Popo Agie Formation (Hunt et al. 1998)
Theropoda indet.
Age: Carnian (Pipiringos and O’Sullivan
1978; Lucas 1993a)
2. Bell Springs Formation (Lockley 1986a;
Lucas et al. 1998a)
Theropod tracks
Age: Carnian (Pipiringos and O’Sullivan
1978)
3.5.
TEXAS
,
UNITED STATES
1. Tecovas Formation (Hunt and Lucas
1994; Heckert and Lucas 1997)
Saurischia incertae sedis
?Chindesaurus bryansmalli (including
Caseosaurus crosbyensis)
Theropod tracks
Ornithischia indet. (=Tecovasaurus
murryi)
Age: late Carnian (Heckert and Lucas
1997)
520 DINOSAUR DISTRIBUTION AND BIOLOGY
FIGU RE 24.3. Late Tr iassic to Late Jurassic localities in the southwestern United States (3). Scale =600 km.

2. Bull Canyon Formation (Jacobs and
Murry 1980; Chatterjee 1984, 1986b,
1993; Murry 1986; Sereno 1991b; Hunt
and Lucas 1994; Long and Murry 1995;
Heckert and Lucas 1997; Lucas et al.
1998a)
Theropoda
?Ceratosauria
Shuvosaurus inexpectatus
Theropoda incertae sedis
Protoavis texensis
?Prosauropoda indet.
Ornithischia
Technosaur us smalli
Age: early Norian (Lucas et al. 1998a)
3.19.
NEW YORK
,
UNITED STATES
1. Upper Stockton beds (Olsen 1980a;
Weishampel and Young 1996)
Theropod tracks
Age: late Carnian (Olsen 1980a)
3.22.
NEW JERSEY
,
UNITED STATES
1. Lockatong Formation (Bock 1952;
Weishampel and Young 1996)
Theropod tracks
Age: late Carnian (Olsen 1980a; Lucas
and Huber 1993)
2. Lower Passaic Formation (Baird 1957;
Weishampel and Young 1996)
Theropod tracks
Ornithischian tracks
Age: Norian-Rhaetian (Lucas and Huber
1993)
3.23.
PENNSYLVANIA
,
UNITED STATES
1. Lockatong Formation (Bock 1952;
Sullivan et al. 1994; Weishampel and
Young 1996)
Theropod tracks
Ornithischian tracks
Age: late Carnian (Olsen 1980a; Lucas
and Huber 1993)
2. New Oxford Formation (Huene 1921b;
Galton 1983a; Hunt and Lucas 1994;
Weishampel and Young 1996)
?Prosauropoda indet. (=Palaeosaurus
fraserianus)
Ornithischia indet. (=Thecodontosaurus
gibbidens)
Age: late Carnian (Lucas and Huber 1993)
3. Gettysburg Formation (Olsen and Baird
1986; Weishampel and Young 1996)
Theropod tracks
Ornithischian tracks
Age: Norian-Rhaetian (Lucas and Huber
1993)
4. Lower Passaic Formation (Olsen and
Baird 1986; Weishampel and Young
1996)
Theropod tracks
Ornithischian tracks
Age: Norian-Rhaetian (Lucas and Huber
1993)
3.25.
MARYLAND
,
UNITED STATES
1. Gettysburg Formation (Mitchell 1895;
Weishampel and Young 1996)
Theropod tracks
Age: Norian-Rhaetian (Lucas and Huber
1993)
3.26.
VIRGINIA
,
UNITED STATES
1. Cow Branch Formation (Olsen et al.
1978; Weishampel and Young 1996)
Theropod tracks
Ornithischian tracks
Age: late Carnian (Lucas and Huber
1993)
2. Manassas Sandstone (Weishampel and
Young 1996)
Theropod tracks
Age: late Carnian (Lucas and Huber
1993)
3. Balls Bluff Formation (Weems 1987;
Weishampel and Young 1996)
Ornithischian tracks
Theropod tracks
Age: ?early Norian (Weems 1987)
3.27.
NORTH CAROLINA
,
UNITED STATES
1. Pekin Formation (Hunt and Lucas 1994;
Weishampel and Young 1996)
Ornithischia indet. (=Pekinosaurus
olseni)
Age: late Carnian (Lucas and Huber
1993)
DINOSAUR DISTRIBUTION 521
Europe
The history of Late Triassic dinosaurs is well known from Europe
(55 locations, an increase of over 40%), in particular, in the
classic Upper Triassic sequence of the Neckar Valley of Baden-
Württemberg, Germany. These German localities were worked
originally by E. Fraas but most successfully by F. von Huene
and later by R. Seemann. Additional work on Upper Triassic sites
in Germany includes that by O. Jaekel at Halberstadt (Nieder-
sachsen) and by F. von Huene and most recently by P. Welln-
hofer in northern Bavaria. Similar dinosaur faunas are known
from Switzerland (from the work of B. Peyer at Hallau and
H. Rieber and M. Saunders at Frick) and France (by E. Buffetaut,
6.1.
MALMÖHUS LAN
,
SWEDEN
1. Lower Höganäs Formation (Bölau 1952;
Gierli´nski and Ahlberg 1994)
Theropod tracks
Age: Rhaetian (Gierli´nski and Ahlberg
1994)
7.1.
GRAMPIAN
,
SCOTLAND
1. Lossiemouth Sandstone Formation
(Benton 1984a; Benton and Walker
1985)
?Dinosauria
Saltopus elginensis
Age: late Carnian (Warrington et al. 1980;
Benton and Spencer 1995)
8.1.
MID
-
GLAMORGAN
,
WALES
1. Rhaetic Beds (Newton 1899; Galton
1985e)
?Theropoda indet.
Tetanurae incertae sedis
“Zanclodon” cambrensis
Age: Rhaetian (Warrington et al. 1980)
8.2.
SOUTH GLAMORGAN
,
WALES
1. Mercia Mudstone Group (Sollas 1879;
Tucker and Burchette 1977; Delair and
Sarjeant 1985; Lockley et al. 1996b)
Theropod tracks
Age: Norian (Delair and Sarjeant 1985)
2. Unnamed unit (Kermack 1984; Edwards
1998; Rauhut and Hungerbühler 2000)
Theropoda
Ceratosauria
?Syntarsus sp.
Theropoda indet.
Prosauropoda
Thecodontosaurus antiquus
Age: ?Norian (Fraser et al. 1985)
10.1.
SOMERSET
,
ENGLAND
1. Westbur y Formation (Seeley 1898;
Galton 1985f; Martill and Dawn 1986;
Storrs 1994)
Prosauropoda
Camelotia borealis
Age: Rhaetian (Warrington et al.
1980)
10.2.
AVON
,
ENGLAND
1. Magnesian Conglomerate (Riley and
Stutchbury 1836, 1840; Galton 1984b;
Benton and Spencer 1995)
Theropoda indet.
Prosauropoda
Thecodontosaurus antiquus
Thecodontosaurus sp.
S. Chopard, G. Cuny, P. Ellenberger, J. Pidancet, N. Theobald,
and G. Wouters). Other important Late Triassic faunistics from
elsewhere in Europe include research by C. Duffin and co-
workers in Belgium; by A. D. Walker and M. J. Benton at Elgin
in Scotland; by N. C. Fraser, K. A. Kermack, P. M. Robinson,
G. M. Walkden, and D. I. Whiteside on the fissure fillings in
England and Wales; by A. Ahlberg, E. Bölau, and G. Gierli´nski
in Sweden; by G. Gierli´nski in Poland; by P. Mietto in Italy;
and by J. Michalík and M. Kundrát in Slovakia. Locations for
Upper Triassic dinosaurs in Europe are given in figures 24.4
and 24.5.

Age: ?Norian-Rhaetian (Warrington
et al. 1980)
2. Unnamed unit (Seeley 1891;
Long 1998; Vickers-Rich et al.
1999b)
Prosauropoda indet. (=Agrosaurus
macgillivrayi)
Age: Late Triassic (Vickers-Rich et al.
1999b)
3. Unnamed unit of Fissure Fills (Fraser
et al. 2002)
?Dinosauria indet. (=Agnosphitys
cromhallensis)
Age: Late Triassic (Fraser et al. 2002)
10.3.
GLOUCESTERSHIRE
,
ENGLAND
1. Unnamed unit of Fissure Fills
(Fraser and Walkden 1983;
Fraser 1994)
Theropoda indet.
Prosauropoda
Thecodontosaurus sp.
Age: late Norian (Fraser and Walkden
1983)
11.1.
PROVINCE DU LUXEMBOURG
,
BELGIUM
1. ?HVL, Sl, S2, UDK I (Duffin et al. 1983)
Prosauropoda indet.
Age: Rhaetian (Duffin et al. 1983)
13.1.
GRAND DUCHY OF LUXEMBOURG
1. Unnamed unit (Godefroit et al. 1998a)
522 DINOSAUR DISTRIBUTION AND BIOLOGY
FIGU RE 24.4. Late Tr iassic to Early Jurassic localities in Sweden (6), Scotland (7), Wales (8), Nor thern Ireland (9), England (10), Belgium (11),
Luxembourg (13), France (14), Portugal (16), Germany (17), Switzerland (18), and Italy (19). Scale =600 km.

?Ornithischia indet.
Age: late Rhaetian (Godefroit et al.
1998a)
14.1.
DÉPARTEMENT DE LA VENDÉE
,
FRANCE
1. Niveaux 1-7 (Bessonat et al. 1965;
Lapparent and Montenat 1967)
Theropod tracks
Age: Rhaetian (Lapparent and
Montenat 1967)
14.2.
DÉPARTEMENT DE LA HAUTE
-
MARNE
,
FRANCE
1. “Grès de l’lnfralias” (Sauvage 1907;
Huene 1907–8; Galton 1985e; Buffetaut
et al. 1991; Cuny 1995)
?Theropoda indet. (including
Plateosaurus elizae)
Prosauropoda
Plateosaurus sp.
Prosauropoda indet.
Age: Rhaetian (Huene 1907–8)
14.3.
DÉPARTEMENT DE LA MEURTHE
-
ET
-
MOSELLE
,
FRANCE
1. Grès à Avicula contorta (Laugier 1971;
Buffetaut 1985; Buffetaut and Wouters
1986)
Theropoda indet.
Prosauropoda
Plateosaurus sp.
Age: late Norian (Cuny and Ramboer
1991); Norian (Buffetaut
1985; Buffetaut and Wouters
1986); Rhaetian (Sigogneau-
Russell 1983)
14.4.
DÉPARTEMENT DU DOUBS
,
FRANCE
1. Marnes Irisées Supérieures Formation
(Pidancet and Chopard 1862; Lapparent
1967; Buffetaut et al. 1991; Galton
1998a)
Prosauropoda
Plateosaurus longiceps
Age: ?Norian (=Keuper; Lapparent 1967)
14.5.
DÉPARTEMENT DU JURA
,
FRANCE
1. Marnes Irisées Supérieures Formation
(Pidancet and Chopard 1862; Théobald
et al. 1967; Buffetaut et al. 1991; Galton
1998a)
Prosauropoda
Plateosaurus longiceps
Prosauropoda indet.
(=Dimodosaurus poligniensis)
Age: late Norian (equivalent to the
Knollenmergel; Brenner
1973)
2. Marnes de Chalin Formation (Cuny and
Mazin 1993)
Undescribed prosauropod
Age: Norian (Cuny and Mazin 1993)
3. Unnamed unit (Henry 1876; Galton
1985e; Buffetaut et al. 1991)
?Theropoda indet. (=Megalosaurus
obtusus)
Age: Rhaetian (Lapparent 1967)
14.6.
DÉPARTEMENT DE L
’
AIN
,
FRANCE
1. Marnes Irisées Supérieures Formation
(Gervais 1861, 1869; Buffetaut et al.
1991; Bardet and Cuny 1993)
Prosauropoda
?Thecodontosaurus sp.
Age: Late Triassic (=Keuper;
Lapparent and Lavocat
1955)
14.7.
DÉPARTEMENT DU GARD
,
FRANCE
1. Grès supérieurs et Argilites bariolées
Formation (Gand et al. 2000)
Theropod tracks
Prosauropod tracks
Age: Norian (Gand et al. 2000)
2. Unnamed unit (Gauffre 1995b)
Prosauropoda indet.
Age: Norian (Gauffre 1995b)
3. Unnamed unit (Ellenberger 1965)
Theropod tracks
Age: Late Triassic (=Keuper; Ellenberger
1965)
14.8.
DÉPARTEMENT DU VAR
,
FRANCE
1. Unnamed unit (Ellenberger 1965)
Theropod tracks
Age: Rhaetian (Ellenberger 1965)
14.9.
DÉPARTEMENT DE L
’
ARDEÈCHE
,
FRANCE
1. Unnamed unit (Courel and Demathieu
2000)
DINOSAUR DISTRIBUTION 523
FIGU RE 24.5. Late Triassic to Late Cretaceous localities in Poland (20), the Czech Republic (23), Slovakia (24), Austria (25), Hungary (26), Slovenia (27),
Croatia (28), Romania (29), Ukraine (30), and Kurskaya Oblast, Russia (37.6). Scale =600 km.

Theropod tracks
Age: early Carnian (Courel and
Demathieu 2000)
17.1.
BIEDERSACHSEN
,
GERMANY
1. Knollenmergel (Huene 1907–8;
Weishampel and Westphal 1986)
Prosauropoda
Plateosaurus cf. longiceps
Age: late Norian (Brenner 1973)
17.2.
BADEN
-
WÜRTTEMBERG
,
GERMANY
1. Lower Stubensandstein (Berckhemer
1938; Galton 1984b; Haderer 1990)
Unnamed dinosaur (=Herrerasauria
incertae sedis)
Theropod tracks
Prosauropoda
Sellosaurus gracilis
Age: middle Norian (Brenner 1973)
2. Middle Stubensandstein (Huene 1907–8,
1932; Galton 1973b, 1985a, 1985b;
Galton and Bakker 1985; Sereno and
Wild 1992; Rauhut and Hungerbühler
1998)
Dinosauria incertae sedis (=Herrerasauria
incertae sedis)
Theropoda
Ceratosauria
Procompsognathus triassicus
partim
Ceratosauria indet. (including
Dolichosuchus cristatus,
Halticosaurus longotarsus)
Theropoda indet. (including
Tanystrophaeus posthumus,
Procompsognathus triassicus
partim)
Prosauropoda
Sellosaurus gracilis (including
Palaeosaurus diagnosticus,
Thecodontosaurus
hermannianus)
Prosauropoda indet. (including
Sellosaurus fraasi,
Teratosaurus minor,
T. trossingensis)
Age: middle Norian (Brenner 1973)
3. Trossingen Formation (Huene 1932;
Seemann 1933; Galton 1985e;
Weishampel and Westphal 1986; Sander
1992; Rauhut and Hungerbühler 1998;
=Knollenmergel)
Theropoda
Ceratosauria indet. (=cf. Liliensternus
liliensterni)
Prosauropoda
Plateosaurus longiceps (including
P. fraasianus, P. integer,
P. trossingensis)
Prosauropoda indet. (including
Plateosaurus erlenbergiensis,
P. plieningeri, P. quenstedti,
Gresslysaurus robustus,
Pachysaurus ajax, P. giganteus,
P. magnus, P. reiningeri,
P. wetzelianus)
Age: late Norian (Brenner 1973)
4. Rhätsandstein (Huene 1907–8; Galton
1985e)
Theropoda indet. (=Megalosaurus
cloacinus)
?Prosauropoda indet. (=Plateosaurus
ornatus)
Age: Rhaetian–?early Hettangian
(Clemens 1980)
17.3.
BAYERN
,
GERMANY
1. Benkersandstein (Rehnelt 1952)
Theropod tracks
Age: early Carnian (Laemmlen 1956)
2. Unterer Gipskeuper (Rehnelt 1950)
Theropod tracks
Age: early Carnian (Warrington et al. 1980)
3. Ansbachersandstein (Heller 1952; Olsen
and Baird 1986)
Theropod tracks
Ornithischian tracks
Age: late Carnian (Laemmlen 1956)
4. Blasensandstein (Weiss 1934)
Theropod tracks
Age: early Norian (Laemmlen 1956)
5. Plattensandstein (Kuhn 1938)
Theropod tracks
Age: early Norian (Laemmlen 1956)
6. Oberer Semionotensandstein (Beurlen
1950; Heller 1952)
Theropod tracks
Age: middle Norian (Laemmlen 1956)
7. Feuerletten (Dehm 1935; Urlichs 1966;
Wellnhofer 1993b)
Prosauropoda
Plateosaurus engelhardti
Prosauropoda indet. (=Plateosaurus
bavaricus)
Age: late Norian (Urlichs 1966)
8. Unnamed unit (Kuhn 1958a, 1958b)
Theropod tracks
Age: Rhaetian-?Sinemurian (=Rätolias;
Kuhn 1958a)
17.4.
SACHSEN
-
ANHALT
,
GERMANY
1. Knollenmergel (Jaekel 1913–14; Kuhn
1939; Sander 1992)
Theropoda
Ceratosauria indet. (including
Halticosaurus longotarsus)
Prosauropoda
Plateosaurus longiceps (including
cf. Palaeosaurus
?diagnosticus)
Age: late Norian (Brenner 1973)
2. Unnamed unit (Jaekel 1913–14)
Theropoda
Ceratosauria indet. (=Pterospondylus
trielbae)
Age: Rhaetian (Huene 1932)
17.5.
THÜRINGEN
,
GERMANY
1. Knollenmergel (Rühle von Lilienstern
et al. 1952; Welles 1984; Rauhut and
Hungerbühler 1998; Galton 2001a)
Theropoda
Ceratosauria
Liliensternus liliensterni
(=Halticosaurus liliensterni)
Prosauropoda
Ruehleia bedheimensis
Prosauropoda indet. (including
Gresslyosaurus torgeri)
Age: late Norian (Brenner 1973)
18.1.
KANTON BASELLAND
,
SWITZERLAND
1. Knollenmergel (Huene 1932)
Prosauropoda indet. (=Gresslyosaurus
ingens)
Age: late Norian (Brenner 1973)
18.2.
KANTON AARGAU
,
SWITZERLAND
1. Obere Bunte Mergel (Rieber 1985a,
1985b; Sander 1992; Rauhut and
Hungerbühler 1998)
Theropoda indet. (=cf. Liliensternus sp.)
Prosauropoda
Plateosaurus cf. longiceps
Age: late Norian (i.e., equivalent to the
Knollenmergel; Brenner
1973)
18.3.
KANTON SCHAFFHAUSEN
,
SWITZERLAND
1. Zanclodonmergel (=Knollenmergel;
Peyer 1944)
Prosauropoda
Plateosaurus sp.
Age: late Norian (Brenner 1973)
2. Unnamed unit (Peyer 1944; Tatarinov
1985)
Theropoda
?Tetanurae indet.
Prosauropoda
Plateosaurus sp.
?Ornithopoda
?Heterodontosauridae indet.
(=?Abrictosaurus sp.)
Age: ?late Norian–Hettangian (Clemens
1980)
19.1.
REGIONE TOSCANA
,
ITALY
1. Quarziti Viola Zonate Formation (Huene
1941; Tongiorgi 1980; Leonardi and
Lockley 1995; Leonardi 2000)
Theropod tracks
Age: late Carnian (Tongiorgi 1980)
19.2.
REGIONE TRENTINO
–
ALTO ADIGE
,
ITALY
1. Dolomia Principale Formation (Leonardi
and Avanzini 1994; Leonardi 2000)
Theropod track
Age: Norian (Leonardi and Avanzini
1994; Leonardi 2000)
19.3.
REGIONE VENETO
,
ITALY
1. Dürrestein Formation (Leonardi 2000)
?Prosauropod tracks
Age: late Carnian (Leonardi 2000)
2. Dolomia Principale Formation (Mietto
1988, 1990, 1992; Mietto and Dalla
Vecchia 2000)
Theropod tracks
Prosauropod tracks
Ornithopod tracks
Age: late Carnian (Mietto 1988)
524 DINOSAUR DISTRIBUTION AND BIOLOGY

3. Dolomia Principale Formation (Mietto
1992; Dalla Vecchia 1995)
Theropod tracks
Age: Norian (Mietto 1992; Dalla
Vecchia 1995; Leonardi
2000)
19.4.
REGIONE FRIULI
–
VENEZIA GIULIA
,
ITALY
1. Dolomia Principale Formation (Dalla
Vecchia 1997a, 2000a; Dalla Vecchia
and Mietto 1998)
Theropod tracks
Prosauropod tracks
Age: Norian (Dalla Vecchia and Mietto
1998)
19.5.
REGIONE LIGURIA
,
ITALY
1. Montemarcello Formation (Leonardi
2000)
Theropod tracks
Ornithopod tracks
?Prosauropod tracks
?Ornithischian tracks
Age: Carnian or Norian (Leonardi
2000)
20.1.
WOJEWÓDZTWO WROKLAW
,
POLAND
1. Lissauer Breccia (Huene 1932; Osmólska,
pers. comm.)
Theropoda
?Ceratosauria indet. (=Velocipes
guerichi)
Age: Norian (Wild, pers. comm.)
24.1.
PRESOVSKY KRAJ
,
SLOVAKIA
1. Tomanova Formation (Michalík and
Kundrát 1998)
Theropod tracks
Age: Norian-Rhaetian (Michalík and
Kundrát 1998)
27.1.
OBCINA IDRIJA
,
SLOVENIA
1. Unnamed unit (equivalent to the
Dolomia Principale; Dalla Vecchia
1997a)
?Prosauropod tracks
Age: Norian-Rhaetian (Dalla Vecchia
1997a)
DINOSAUR DISTRIBUTION 525
Asia
Upper Triassic sites that have yielded dinosaur remains are re-
stricted to Madhya Pradesh and Andhra Pradesh, India, and
Changwat Khonkaen, Thailand (four locations). Early studies
of the Indian fauna, yielding early theropod and prosauropod
remains, were carried out by R. Lydekker and F. von Huene, and
FIGU RE 24.6. Late Tr iassic to Late Cretaceous localities in Iran (36), Afghanistan (44), Pakistan (45), and India (46). Scale =600 km.
46.1.
MADHYA PRADESH STATE
,
INDIA
1. Tiki Formation (Chatterjee and
Majumdar 1987)
Theropoda indet.
Age: Carnian (Chatterjee 1978a)
46.2.
ANDHRA PRADESH STATE
,
INDIA
1. Lower Maleri Formation (Jain 1983;
Chatterjee 1987; Bandyopadhyay 1999;
Bandyopadhyay et al. 2002)
Saurischia incertae sedis
Alwalkeria maleriensis
Prosauropoda indet.
(=Massospondylus hislopi,
Plateosaurus sp.)
Age: Carnian-Norian (Kutty 1969)
later work was performed by S. Chatterjee, T. Roy-Chowdhury,
S. L. Jain, T. S. Kutty, and P. Robinson. The newly discovered Late
Triassic fauna from Thailand was described by V. Suteethorn
and E. Buffetaut. Figures 24.6 and 24.7 provide information on
the locations for Upper Triassic dinosaurs of Asia.

FIGU RE 24.7. Late Triassic to Early Cretaceous localities in Mongolia (47), People’s Republic of China (48), North Korea (49), South Korea (50), and Japan (53).
Scale =600 km.
FIGU RE 24.8. Late Tr iassic to Late Cretaceous
localities in Colombia (54), Ecuador (55), Peru
(56), Brazil (57), Bolivia (59), Guyana (60), and
Venezuela (61). Scale =600 km.

57.1.
ESTADO DO RIO GRANDE DO SUL
,
BRAZIL
1. Upper Santa Maria Formation
(Huene 1942; Colbert 1970; Bonaparte
1973; Galton 1977a; Kellner et al.
1999; Kischlat 1999; Langer et al.
1999)
?Dinosauria
Spondylosoma absconditum
?Dinosauria indet. (=Teywasu
barberennai)
Saurischia
Herrerasauridae
Staurikosaurus pricei
?Sauropodomorpha
Saturnalia tupiniquim (may be
the sister taxon to
Sauropodomorpha)
Unnamed prosauropod
Age: ?late Ladinian or early Carnian
(Galton 1977a)
2. Caturrita Formation (Kellner et al. 1999;
Bonaparte et al. 1999)
Eusaurischia
Guaibasaurus candelariensis
Unnamed prosauropod
Age: late Carnian (Bonaparte et al. 1999)
DINOSAUR DISTRIBUTION 527
FIGU RE 24.9. Late Tr iassic to Late Cretaceous localities in
Uruguay (58), Chile (62), and Argentina (63). Scale =600 km.
2. Dharmaram Formation (Kutty 1969;
Kutty and Roy-Chowdhury 1970; Jain
1996)
Prosauropoda indet.
Ornithischia indet.
Age: late Norian–Rhaetian
(Kutty 1969); Early
Jurassic (Bandyopadhyay
1999)
51.1.
CHANGWAT KHONKAEN
,
THAILAND
1. Nam Phong Formation (Buffetaut
et al. 1995c, 2002a, 2002b;
Suteethorn et al. 1995a; Buffetaut and
Suteethorn 1998b)
Prosauropoda indet.
Sauropoda
Isanosaurus attavipachi
Age: late Norian–Rhaetian (Racey et al.
1994)
South America
The earliest known and most basal dinosaur material from both
Saurischia and Ornithischia comes from South America (11 loca-
tions, an increase of nearly 60%). Collection of and work on these
early forms have been carried out by a number of paleontologists,
among them A. Arcucci, J. F. Bonaparte, R. M. Casamiquela, R. A.
Coria, F. von Huene, G. Leonardi, R. Martínez, F. Novas, O. A.
Reig, and P. C. Sereno. Late Triassic dinosaur locations of South
America are given in figures 24.8 and 24.9.

63.1.
PROVINCIA DE LA RIOJA
,
ARGENTINA
1. Los Rastros Formation (Arcucci et al.
1995)
Theropod tracks
Age: ?Ladinian (Arcucci et al. 1995)
2. Ischigualasto Formation (Casamiquela
1967b; Bonaparte 1973, 1976; Leonardi
1994)
?Theropod tracks
Ornithischia
Pisanosaurus mertii
Age: Carnian (Bonaparte 1973;
Stipanicic 1983)
3. Los Colorados Formation (Bonaparte
1973, 1978b; Galton 1985f; Arcucci and
Coria 1996, 1997, 1998, 2003; Bonaparte
and Pumares 1995)
Theropoda
Theropoda incertae sedis
Zupaysaurus rougieri
Theropoda indet.
Prosauropoda
Riojasaurus incertus
Coloradisaurus brevis
Lessemsaurus sauropoides
Prosauropoda indet. (including
Strenusaurus procerus)
Age: Norian (Bonaparte 1973; Stipanicic
1983)
63.2.
PROVINCIA DE SAN JUAN
,
ARGENTINA
1. Ischigualasto Formation (Reig 1963;
Bonaparte 1973; Brinkman and Sues
1987; Novas 1986 1993; Sereno 1993;
Sereno and Novas 1993; Sereno et al.
1993)
Saurischia
Eoraptor lunensis
Herrerasauridae
Herrerasaurus ischigualastensis
(including Ischisaurus cattoi,
Frenguellisaurus
ischigualastensis,
cf. Staurikosaurus sp.)
Dinosauria indet. (=Teyuwasu
barberenai)
Age: Carnian (Bonaparte 1973;
Stipanicic 1983)
2. Quebrada del Barro Formation (Bossi
and Bonaparte 1978)
Prosauropoda
Riojasaurus incertus
Age: Norian (equivalent to Los
Colorados Formation; Bossi
and Bonaparte 1978)
3. Cañon del Colorado Formation (Milana,
pers. comm.)
Undescribed prosauropod
Age: Late Triassic–Early Jurassic (Milana,
pers. comm.)
4. Los Colorados Formation. (Martínez,
pers. comm.)
Prosauropoda indet.
Age: Late Triassic (Martínez, pers.
comm.)
5. Portezuelo Formation (Marsicano and
Barredo 2004)
Theropod tracks
Sauropodomorph tracks
Age: Carnian (Mariscano and Barredo
2004)
63.3.
PROVINCIA DE SANTA CRUZ
,
ARGENTINA
1. “Complejo Porfirico” (Leonardi 1989)
Theropod tracks
Age: Late Triassic (Leonardi 1989)
2. Laguna Colorada Formation (Bonaparte
1973; Bonaparte and Vince 1979;
Casamiquela 1980; Baez and Mariscano
1998, 2001)
Prosauropoda
?Plateosaurus sp.
Mussaurus patagonicus
Ornithopoda
Heterodontosauridae
cf. Heterodontosaurus sp.
Dinosaur eggs
Age: ?Norian (Bonaparte and Vince
1979)
528 DINOSAUR DISTRIBUTION AND BIOLOGY
Africa
The majority of our knowledge about the Late Triassic and
Early Jurassic comes from the famous Stormberg Series, Karroo
Basin, of South Africa and Lesotho, primarily through the
early efforts of S. H. Haughton and E. C. N. van Hoepen and
the recent work of J. Attridge, A. W. Crompton, F. and P. Ellen-
berger, P. M. Galton, C. Gow, K. A. Kermack, J. W. Kitching, and
J. van Heerden. Additional locations from the Upper Triassic of
Africa are known from Morocco through work by J.-M. Dutuit
and F.-X. Gauffre, and from Zimbabwe via studies by M. R.
Cooper and M. A. Raath. These locations (12 in number, an in-
crease of approximately 20%) are given in figures 24.10 and
24.11.
64.1.
PROVINCE DE MARRAKECH
,
MOROCCO
1. Argana Formation (Dutuit 1972, 1974;
Biron and Dutuit 1981; Gauffre 1993a)
Theropoda indet.
Theropod tracks
Prosauropoda
Azendohsaurus laaroussii partim
Ornithischian tracks
Age: middle Carnian (Cousminer and
Manspeizer 1976)
77.1.
MASHONALAND NORTH
,
ZIMBABWE
1. Pebbly Arkose (Raath 1996)
Prosauropoda indet.
Age: early Carnian (Anderson and
Anderson 1984; Raath
1996)
77.4.
MATABELELAND SOUTH
,
ZIMBABWE
1. Mpandi Formation (Cooper 1980)
Prosauropoda
Euskelosaurus cf. browni
Age: Norian (Cooper 1980)
81.1.
TRANSVAAL
,
SOUTH AFRICA
1. Bushveld Sandstone (Haughton 1924)
Prosauropoda
Euskelosaurus browni
Prosauropoda indet. (=Gryponyx
transvaalensis, Gigantoscelus
molengraaffi)
Age: late Carnian or early Norian
(?=Lower Elliot Formation;
Kitching, pers. comm.)
81.2.
FREE STATE PROVINCE
,
SOUTH AFRICA
1. Lower Elliot Formation (Galton and
Cluver 1976; Kitching and Raath 1984;
Olsen and Galton 1984; Yates and
Kitching 2003)
Prosauropoda
cf. Euskelosaurus browni
Prosauropoda indet. (=Eucnemesaurus
fortis)
Sauropoda
Antetonitrus ingenipes
Age: late Carnian or early Norian (Olsen
and Galton 1984)
81.3.
CAPE PROVINCE
,
SOUTH AFRICA
1. Molteno Formation (Raath et al. 1990;
Raath 1996)
Theropod tracks
Age: early Carnian (Anderson and
Anderson 1984)
2. Lower Elliot Formation (Haughton 1924;
Kitching and Raath 1984; Olsen and
Galton 1984; Galton 1985f; Galton and
Heerden 1985; Heerden and Galton
1997; Galton et al. in press)
Dinosauria incertae sedis
Aliwalia rex
Prosauropoda
Massospondylus sp.
Melanorosaurus readi
Melanorosaurus sp.
Euskelosaurus browni
Euskelosaurus sp.
Prosauropoda indet. (=Orinosaurus
capensis, including
Plateosaurus stormbergensis,
P. cullingworthi,
Euskelosaurus
africanus)

?Prosauropod tracks
Sauropoda
Blikanasaurus cromptoni
?Ornithischia indet.
Age: late Carnian or early Norian
(Olsen and Galton
198 4)
82.1.
LERIBE DISTRICT
,
LESOTHO
1. Lower Elliot Formation
(Huene 1932; Kitching and
Raath 1984)
Prosauropoda
Melanorosaurus sp.
Age: late Carnian or early Norian
(Olsen and Galton
198 4)
82.2.
MASERU DISTRICT
,
LESOTHO
1. Lower Elliot Formation (Ellenberger
1970; Kitching and Raath 1984; Olsen
and Galton 1984)
Theropod tracks
Prosauropoda
cf. Melanorosaurus sp.
Prosauropod tracks
Age: late Carnian or early Norian (Olsen
and Galton 1984)
82.3.
MAFETENG DISTRICT
,
LESOTHO
1. Lower Elliot Formation (Ellenberger
1970; Kitching and Raath 1984)
Prosauropoda
Euskelosaurus sp.
Age: late Carnian or early Norian (Olsen
and Galton 1984)
82.4.
MOHALES HOEK DISTRICT
,
LESOTHO
1. Lower Elliot Formation (Ellenberger
1970; Kitching and Raath 1984; Olsen
and Galton 1984)
Theropod tracks
Prosauropoda
Euskelosaurus sp.
cf. Melanorosaurus sp.
Prosauropod tracks
Age: late Carnian or early Norian (Olsen
and Galton 1984)
83.1.
FARITANY TOLIARA
,
MADAGASCAR
1. Isalo II Group (Flynn et al. 1999b)
Unnamed prosauropods
Age: ?Middle Triassic–early Carnian
(Flynn et al. 1999b)
DINOSAUR DISTRIBUTION 529
FIGU RE 24.10. Late Triassic to Late Cretaceous localities in Morocco (64), Algeria (65), Tunisia (66), Mali (67),
Niger (68), Libya (70), and Cameroun (73). Scale =600 km.
Australasia
Only one location in Queensland, Australia, has yielded Upper
Triassic dinosaur material (fig. 24.12).
84.1.
QUEENSLAND
,
AUSTRALIA
1. Blackstone Formation (Molnar 1982;
Thulborn 1998)
Theropod tracks
Age: middle-late Carnian (Thulborn
1998)

Early Jurassic
Over the past decade, the dinosaur fossil record for the Early Juras-
sic has expanded appreciably. More than 80 locations are found
in North America, Europe, Asia, South America, and Africa, as
well as newly discovered sites in Australasia and Antarctica.
North America
Like those of the Late Triassic, North American Early Jurassic
dinosaurs (24 locations, an increase of over 30%) are known pri-
marily from the Maritimes of Canada (through work by D. Baird,
P. E. Olsen, N. Shubin, and H.-D. Sues), from the Southwest
of the United States (through studies by E. H. Colbert, A. W.
530 DINOSAUR DISTRIBUTION AND BIOLOGY
FIGU RE 24.11. Late Tr iassic to Late Cretaceous
localities in Ethiopia (72), Kenya (74), Tanzania
(75), Mozambique (76), Zimbabwe (77), Zambia
(78), Malawi (79), Namibia (80), South Africa (81),
Lesotho (82), and Madagascar (83). Scale =600 km.
Crompton, F. A. Jenkins, M. G. Lockley, M. Morales, K. Padian,
T. Rowe, and S. P. Welles), and again from the eastern seaboard
of the United States (primarily from studies by P. E. Olsen).
North American locations for the Lower Jurassic are given in fig-
ures 24.1–24.3 and 24.13.
1.7.
NOVA SCOTIA
,
CANADA
1. Scots Bay Formation (Olsen et al. 1982;
Olsen et al. 1987; Sues et al. 1987; Shubin
et al. 1994; Weishampel and Young
1996)
Theropod tracks
Age: Hettangian (Olsen et al. 1982)
2. McCoy Brook Formation (Olsen et al.
1982; Olsen et al. 1987; Sues et al. 1987;
Shubin et al. 1994; Weishampel and
Young 1996; Rainforth 2003)
Theropoda indet.
Theropod tracks
Prosauropoda
cf. Anchisaurus sp.
cf. Ammosaurus sp.
Prosauropod tracks
Ornithischia indet.
(?Scutellosaurus sp.)
Ornithischian tracks
Age: Hettangian (Olsen et al. 1982)

FIGU RE 24.12. Late Tr iassic to Late Cretaceous localities in Australia (84). Scale =600 km.
FIGU RE 24.13. Early Jurassic to Early Cretaceous localities in western Canada (1) and the nor thwest-central United States (3). Scale =600 km.

3.1.
UTAH
,
UNITED STATES
1. Wingate Sandstone (Brady 1960; Dubiel
et al. 1989)
?Theropod tracks
Age: Hettangian-Sinemurian (Olsen
et al. 1982)
2. Moenave Formation (Miller et al. 1989;
Hayden et al. 2001; Kirkland et al. 2002)
Theropod tracks
Prosauropod tracks
Age: Hettangian (Miller et al. 1989)
3. Navajo Sandstone (Stokes 1978;
Lockley et al. 1992a; Hamblin and
Bilbey 1999)
Theropod tracks
Age: Toarcian (Olsen et al. 1982)
3.2.
ARIZONA
,
UNITED STATES
1. Moenave Formation (Morales 1986,
1994; Irby 1996a, 1996b)
Theropoda indet.
Theropod tracks
Ornithischian tracks
Age: Hettangian (Miller et al. 1989)
2. Kayenta Formation (Colbert 1981;
Welles 1984; Attridge et al. 1985;
Morales 1986; Rowe 1989; Bulkley
1996; Morales and Bulkley 1996)
Theropoda
Ceratosauria
Dilophosaurus wetherilli
Syntarsus kayentakatae
Segisaurus halli
Unnamed coelophysoid
Theropod tracks
Prosauropoda
Massospondylus sp.
Ornithischia indet.
Ornithischian tracks
Ornithopoda
Undescribed heterodontosaurid
Thyreophora
Scutellosaurus lawleri
Scelidosaurus sp.
Age: Hettangian (Imlay 1952);
?Sinemurian (Rowe
and Gauthier 1990)
3. Wingate Sandstone (Morales 1996)
Theropod tracks
Age: Sinemurian (Olsen et al. 1982)
4. Navajo Sandstone (Camp 1936;
Galton 1976a; Winkler et al. 1987;
Nations et al. 1996; Rainforth and
Lockley 1996)
Theropoda
Theropoda indet.
Theropod tracks
Prosauropoda
Ammosaurus cf. major
?Prosauropod tracks
Ornithischian tracks
Age: Toarcian-Bajocian (Peterson
and Pipiringos 1979)
3.4.
COLORADO
,
UNITED STATES
1. Navajo Sandstone (Faul and Roberts
1951)
Theropod tracks
Age: Toarcian (Olsen et al. 1982)
2. Kayenta Formation (Stokes and Bruhn
1960; Lockley 1986a)
?Theropod tracks
Age: Hettangian (Imlay 1952);
?Sinemurian (Rowe
and Gauthier 1990)
3.10.
CALIFORNIA
,
UNITED STATES
1. Aztec Sandstone (Reynolds 1989)
Theropod tracks
Age: Early Jurassic (Reynolds 1989)
3.12.
WYOMING
,
UNITED STATES
1. Nugget Sandstone (Stokes 1978)
Theropod tracks
Age: Toarcian (Olsen et al. 1982)
3.20.
MASSACHUSETTS
,
UNITED STATES
1. East Berlin Formation (Ostrom 1972a;
Smith et al. 1996)
Theropod tracks
Age: Hettangian (Smith et al. 1996)
2. Portland Formation (Olsen 1980a;
Thulborn 1991; Gierli´nski 1996b;
Rainforth 2003; =Turner Falls
Formation)
Theropod tracks
Prosauropoda
Anchisaurus polyzelus
Prosauropod tracks
Ornithischian tracks
Dinosaur coprolites
Age: Pliensbachian-Toarcian (Olsen
1980a)
3. Unnamed unit (=?Portland Formation;
Galton 1980c)
Theropoda
Ceratosauria
Podokesaurus holyokensis
Age: Pliensbachian-Toarcian (Olsen
1980a)
3.21.
CONNECTICUT
,
UNITED STATES
1. Shuttle Meadow Formation (Olsen
1980a, 1980b; Weishampel and Young
1996)
Theropoda indet.
Theropod tracks
Ornithischian tracks
Age: Hettangian (Olsen 1980a)
2. East Berlin Formation (Olsen 1980a,
1980b)
Theropod tracks
Ornithischian tracks
Age: Sinemurian (Olsen 1980a)
3. Portland Formation (Olsen 1980a,
1980b; Rainforth 2003)
Theropoda indet. (=Podokesaurus sp.)
Theropod tracks
Prosauropoda
Anchisaurus polyzelus
Ammosaurus major
Prosauropoda indet. (including
Anchisaurus solus)
Prosauropod tracks
Ornithischian tracks
Age: Pliensbachian-Toarcian (Olsen
1980a)
3.22.
NEW JERSEY
,
UNITED STATES
1. Upper Passaic Formation (Olsen
1980a, 1980b; Olsen and Baird 1986;
Weishampel and Young 1996)
Theropod tracks
Ornithischian tracks
Age: early Hettangian (Olsen 1980a)
2. Feltville Formation (Olsen 1980a,
1980b)
Theropod tracks
Ornithischian tracks
Age: early Hettangian (Olsen 1980a)
3. Towaco Formation (Olsen 1980a, 1980b)
Theropod tracks
Ornithischian tracks
Age: Hettangian (Olsen 1980a)
4. Boonton Formation (Olsen 1980a,
1980b)
Theropod tracks
Ornithischian tracks
Age: Hettangian-Sinemurian (Olsen
1980a)
3.26.
VIRGINIA
,
UNITED STATES
1. Midland Formation (Pannell 1985)
Theropod tracks
Age: Pliensbachian–early Sinemurian
(Pannell 1985)
532 DINOSAUR DISTRIBUTION AND BIOLOGY
Europe
Much of what we know about the Early Jurassic of Europe (29
locations, an increase of more than two-fold) comes from the
early English work of R. Owen and A. Smith Woodward; from
later work by, among others, P. C. Ensom; and D. M. Martill;
and from the French studies of E. Buffetaut, G. Cuny, G.-R. De-
mathieu,. P. Ellenberger, A. F. de Lapparent, and C. Montenat.
Other studies include those by A. Ahlberg, E. Bölau, G. Gier-
li´nski, C. Pleijel, and on the small but important dinosaur fauna
of Sweden; by R. Wild and H. Haubold on German faunas; by
G. Gierli´nski and L. Kordos on a Hungarian ichnofauna; and
most recently by M. Lanziger and G. Leonardi, and G. Gierli´n-
ski, on new Italian and Polish track sites, respectively. (The
indeterminate theropod Macrodontophion [Zborzewski 1834] is
now known to be from the Jurassic of Nikolayevskaya Oblast,
Ukraine [location 30.2-1; Kurzanov, pers. comm.], but a bet-
ter age assignment is not possible at this time.) Lower Jurassic
dinosaur locations of Europe are given in figures 24.4 and
24.5.

6.1.
MALMÖHUS LAN
,
SWEDEN
1. Upper Höganäs Formation (Bölau 1952,
1954; Pleijel 1975; Gierli´nski and Ahlberg
1994)
?Theropoda indet.
Theropod tracks
Age: Hettangian (Gierli´nski and Ahlberg
1994)
7.2 .
SKYE
,
SCOTLAND
1. Broadford Beds Formation (Benton et al.
1995)
Theropoda
Ceratosauria indet.
Age: Sinemurian (Cope et al. 1980a)
8.1.
MID
-
GLAMORGAN
,
WALES
1. Unnamed unit (Evans and Kermack
1994)
Theropoda indet.
Age: Hettangian or Sinemurian (Fraser
1988)
9.1.
COUNTY ANTRIM
,
NORTHERN IRELAND
1. Lower Lias (Simms, pers. comm.)
Theropoda indet.
Thyreophora indet.
Age: late Sinemurian (Cope et al. 1980b)
10.4.
NORTH YORKSHIRE
,
ENGLAND
1. Whitby Mudstone Formation (Huene
1926b; Benton and Spencer 1995)
Theropoda indet.
Age: Toarcian (Benton and Spencer
1995)
10.5.
WARWICKSHIRE
,
ENGLAND
1. Lower Lias (Woodward 1908b; Huene
1932)
Theropoda
Ceratosauria indet. (=Sarcosaurus
andrewsi)
Age: late Sinemurian (Cope et al. 1980b)
10.6.
LEICESTERSHIRE
,
ENGLAND
1. Lower Lias (Andrews 1921; Huene 1932)
Theropoda
Ceratosauria
Sarcosaurus woodi
Age: late Sinemurian (Cope et al. 1980b)
10.7.
DORSET
,
ENGLAND
1. Lower Lias (Owen 1861a, 1863;
Lydekker 1888b; Rixon 1968; Ensom
1987, 1989; Martill 1991; Norman,
pers. comm.)
Theropoda indet. (=Megalosaurus
lydekkeri)
Thyreophora
Scelidosaurus harrisonii
Age: late Sinemurian (Cope et al. 1980b)
14.1.
DÉPARTEMENT DE LA VENDÉE
,
FRANCE
1. Niveaux 8-13 (Bessonat et al. 1965;
Lapparent and Montenat 1967)
Theropod tracks
Age: Hettangian (Lapparent and
Montenat 1967)
14.10.
DÉPARTEMENT DE LA MANCHE
,
FRANCE
1. Moon-Airel Formation (=Couches
d’Airel, Larsonneur and Lapparent 1966;
Cuny and Galton 1993; Rauhut and
Hungerbühler 1998)
Theropoda
Ceratosauria
“Liliensternus” airelensis
(=Halticosaurus sp.)
Age: early Hettangian (Cuny and Galton
1993)
2. Calcaire de Valognes (Rioult 1978;
Buffetaut et al. 1991)
Theropoda indet. (=Megalosaurus
cloacinus)
Age: early Hettangian (Buffetaut et al.
1991 )
14.11.
DÉPARTEMENT DE L
’
AVEYRON
,
FRANCE
1. “Dolomie Cubique” (Demathieu 1990;
Demathieu and Sciau 1992, 1999)
Theropod tracks
?Ornithischian tracks
Age: Hettangian (Demathieu and Sciau
1992)
2. Unnamed unit (Ellenberger and Fuchs
1965)
Theropod tracks
Age: ?Pliensbachian (=Lotharingian;
Ellenberger and Fuchs
1965)
14.12.
DÉPARTEMENT DE L
’
HÉRAULT
,
FRANCE
1. Unnamed unit (Thaler 1962; Buffetaut
et al. 1991)
Theropod tracks
Age: Hettangian (Buffetaut et al. 1991)
14.13.
DÉPARTEMENT DE LA DORDOGNE
,
FRANCE
1. Unnamed unit (Le Loeuff et al. 1998,
1999)
Theropod tracks
Thyreophoran tracks
Age: Hettangian (Le Loeuff et al. 1998,
1999)
14.14.
DÉPARTEMENT DE LA LOZÈRE
,
FRANCE
1. “Dolomie àdebit parallelepipedique”
(Thaler 1962; Ellenberger 1965;
Demathieu 1990)
Theropod tracks
Age: Hettangian (Ellenberger 1965)
14.15.
DÉPARTEMENT DE LA MOSELLE
,
FRANCE
1. Unnamed unit (Huene 1926c)
(not plotted)
Theropoda indet. (=Megalosaurus
terquemi)
Age: Hettangian (Huene 1926c)
16.1.
DISTRITO DO LEIRIA
,
PORTUGAL
1. Unnamed unit (Lapparent and
Zbyszewski 1957)
?Thyreophora indet. (=Lusitanosaurus
liasicus)
Age: ?Sinemurian (Lapparent and
Zbyszewski 1957)
17.2.
BADEN
-
WÜRTTEMBERG
,
GERMANY
1. Posidonienschiefer (Wild 1978a)
Sauropoda
Ohmdenosaurus liasicus
Age: middle Toarcian (Geyer and
Gwinner 1962)
17.6.
SCHLESWIG
-
HOLSTEIN
,
GERMANY
1. Unnamed unit (Huene 1966)
Theropoda indet.
Age: Toarcian (=Lias epsilon; Geyer
and Gwinner 1962)
17.7.
MECKLENBURG
,
GERMANY
1. Unnamed unit (Haubold 1990)
Thyreophora
Emausaurus ernsti
Age: early Toarcian (equivalent to the
Posidonienschiefer; Haubold
1990)
19.1.
REGIONE TRENTINO
–
ALTO ADIGE
,
ITALY
1. Lower Member of the Calcari Grigi
di Noriglio Formation (Lanziger and
Leonardi 1992; Dalla Vecchia 1994;
Avanzini et al. 1997; Avanzini, 1998,
2001; Leonardi 2000; Leonardi and
Mietto 2000; Avanzini et al. 2001;
Avanzini and Tomasoni 2001)
Theropod tracks
Sauropod tracks
Ornithischian tracks
?Ornithopoda tracks
Age: Simemurian (Fugagnoli and Loriga
Broglio 1998)
2. Upper part of the Lower Member of the
Calcari Grigi di Noriglio Formation
(Avanzini, 1997)
Theropoda tracks
Sauropoda tracks
Age: late Sinemurian (Leonardi 2000)
3. Tovel Member of the Calcari Grigi
di Noriglio Formation (Avanzini et al.
2001)
Prosauropod tracks
Tyreophoran tracks
Age: Pliensbachian (Avanzini et al. 2001)
19.3.
REGIONE VENETO
,
ITALY
1. Middle-Upper Member of the Calcari
Grigi di Noriglio Formation (Mietto and
Roghi 1994)
Theropod tracks
Age: Pliensbachian (Mietto and Roghi
1994)
19.6.
REGIONE LOMBARDIA
,
ITALY
1. Saltrio Formation (Dal Sasso, pers.
comm. in Dalla Vecchia 2001b)
Theropoda
Undescribed tetanuran
Age: middle Sinemurian (Dalla Vecchia,
pers. comm.)
DINOSAUR DISTRIBUTION 533

20.2.
WOJEWÓDZTWO KIELCE
,
POLAND
1. Zagaje Formation (Pienkowski and
Gierli´nski 1987; Gierli´nski 1990, 1991,
1994; Gierli´nski and Sawicki 1998;
Gierli´nski and Pienkowski 1999;
Gierli´nski et al. 2001)
Theropod tracks
Sauropod tracks
Ornithischian tracks
Age: early Hettangian (Gierli´nski 1990,
1994)
2. Przysucha Formation (Gierli´nski 1990,
1991, 1996a, 1997; Gierli´nski and
Pienkowski 1999; including Sk loby
Formation)
Theropod tracks
?Sauropod tracks
Ornithischian tracks
Age: late Hettangian (Gierli´nski 1990,
1991, 1996a)
26.1.
MEGYE BARANYA
,
HUNGARY
1. Mecsek Coal Formation (Kordos
1983)
?Ornithischian tracks
Age: Hettangian (Kordos 1983)
534 DINOSAUR DISTRIBUTION AND BIOLOGY
Asia
Twelve locations are known to have yielded Lower Jurassic dino-
saur material in Asia. These include sites in Iran (worked prin-
cipally by A. F. de Lapparent), India (known primarily through
studies by S. L. Jain and P. Yadagiri), and the People’s Republic
of China (known through studies by Cai K.-J., Dong Z.-M., He
X.-L., and C.-C. Young, among others). The last country has
produced the most extensive information on Early Jurassic
dinosaurs from this continent and indeed for the world. Asian
dinosaur locations from the Lower Jurassic are given in figures
24.6 and 24.7.
36.1.
OSTAN KERMAN
,
IRAN
1. Shemshak Formation (Lapparent and
Davoudzadeh 1972)
Theropod tracks
Age: ?Hettangian-Pliensbachian (=Lias;
Lapparent and Davoudzadeh
1972 )
36.2.
OSTAN MAZANDARAN
,
IRAN
1. Unnamed unit (Lapparent and Sadat 1975)
Theropod tracks
Age: ?Hettangian-Pliensbachian (=Lias;
Lapparent and Sadat 1975)
44.1.
FARYAB VELAYAT
,
AFGHANISTAN
1. Unnamed unit (Lapparent and Stocklin
1972 )
?Theropod tracks
?Sauropod tracks
?Ornithopod tracks
Age: ?Hettangian-Pliensbachian (=?Lias;
Lapparent and Stocklin
1972 )
46.2.
ANDRA PRADESH STATE
,
INDIA
1. Kota Formation (Jain 1980, 1996;
Yadagiri 1982, 1988; Bandyopadhyay
1999; Bandyopadhyay et al. 2002)
Theropoda indet. (including
Dandakosaurus indicus)
Sauropoda
Kotasaurus yamanpaliensis
Cetiosauridae
Barapasaurus tagorei
Thyreophora indet.
Age: Hettangian-Pliensbachian
(Bandyhopadhyay and
Roy-Chowdhury 1996);
Pliensbachian-Toarcian
(Bandyopadhyay 1999)
48.1.
XIZANG ZIZHIQU
,
PEOPLE
’
S REPUBLIC
OF CHINA
1. Daye Group (Wang et al. 1985; Zhao
unpubl. data)
Undescribed theropod
Prosauropoda
Lufengosaurus changduensis
Undescribed prosauropod
Undescribed sauropod
Undescribed ?thyreophoran
Age: Early Jurassic (Wang et al. 1985)
48.2.
SICHUAN
,
PEOPLE
’
S REPUBLIC OF
CHINA
1. Zhenzhunchong Formation (Chen et al.
1982; Dong et al. 1983; Dong 1984a)
Prosauropoda
cf. Lufengosaurus huenei
(=cf. L. magnus)
Prosauropoda indet. (=Anchisauridae
indet.)
Age: Toarcian-Bajocian (Wang and Sun
1983); Early Jurassic (Dong
et al. 1983)
2. Ziliujing Formation (Dong et al. 1983;
Wang et al. 1985; Dong 1992; He et al.
1998; Luo and Wang 1999, 2000; Luo
et al. 1999)
Theropoda indet.
Sauropoda
Gongxianosaurus shibeiensis
Sauropoda indet. (=Zizhongosaurus
chuanchengensis, Sanpasaurus
yaoi partim)
Ornithopoda indet. (=Sanpasaurus yaoi
partim)
Age: Early Jurassic (Wang et al. 1985)
48.3.
YUNNAN
,
PEOPLE
’
S REPUBLIC OF
CHINA
1. Dull Purplish Beds of the Lower Lufeng
Series (Young 1941a, 1941b, 1948a, 1948b,
1951; Sun et al. 1985; Zhang and Yang
1995; Zhao and Xu 1998; Xu et al. 2001a)
Theropoda
Therizinosauroidea
Eshanosaurus deguchiianus
Theropoda incertae sedis
Lukousaurus yini
Theropoda indet. (=Sinosaurus
triassicus)
Prosauropoda
Lufengosaurus huenei (including
Gyposaurus sinensis,
L. magnus)
Jingshanosaurus xinwaensis
Yunnanosaurus huangi (including
Y. robustus)
“Gyposaurus”sinensis
Age: ?Hettangian (Luo and Wu 1994)
2. Dark Red Beds of the Lower Lufeng
Series (Young 1941a, 1941b, 1948a, 1951,
1982b; Simmons 1965; Chen et al. 1982;
Galton 1985f; Sun et al. 1985; Hu 1993;
Luo and Wu 1994; Lucas 1996; Lamanna
et al. 1998; Wilson and Sereno 1998;
Barrett 1999, 2000a; Dong 2001; Zhao
unpubl. data)
Theropoda
Ceratosauria
“Dilophosaurus” sinensis
Theropoda incertae sedis
Lukousaurus yini
Theropoda indet. (=Sinosaurus
triassicus)
Prosauropoda
Lufengosaurus huenei (including
Gyposaurus sinensis,
L. magnus)
“Gyposaurus” capensis
Yunnanosaurus huangi (including
Y. robustus)
Prosauropoda indet. (=Fulengia
youngi, Tawasaurus minor)
Sauropoda indet. (including
Kunmingosaurus
wusdingensis)
Ornithopoda
Ornithopoda indet.
(=Dianchungosaurus
lufengensis)
Thyreophora incertae sedis
Bienosaurus lufengensis
Tatisaurus oehleri
Age: Sinemurian (Luo and Wu 1994)
3. Fengjiahe Formation (Zhen et al. 1986;
Bai et al. 1990; Zhao unpbul. data)
Theropod tracks
Prosauropoda
Yimenosaurus youngi
Sauropoda indet. (=Chinshakiangosaurus
chunghoensis)
Age: Early Jurassic (Zhen et al. 1986)

48.4.
LIAONING
,
PEOPLE
’
S REPUBLIC OF
CHINA
1. Heichengtzu Series (Yabe et al. 1940;
Young 1960)
Theropod tracks
Age: ?Rhaetic-Toarcian
(Zhen et al.
1989)
53.1.
NAGANO PREFECTURE
,
JAPAN
1. Kuruma Group (Hatakeyama 1995)
Theropod tracks
Age: Early Jurassic (Hatakeyama 1995)
DINOSAUR DISTRIBUTION 535
South America
Four locations in South America have produced dinosaur ichno-
faunas; all are from the Botucatu Formation of Brazil (worked
by G. Leonardi; fig. 24.8). Another, reported by D. E. Russell and
colleagues, and M. R. Sanchez-Villagra and J. M. Clark from
Venezuela, provides the first record of skeletal remains from the
Early Jurassic of South America.
57.1.
ESTADO DO RIO GRANDE DO SUL
,
BRAZIL
1. Botucatu Formation (Leonardi 1989)
Theropod tracks
Age: Hettangian-Toarcian (=Liassic;
Leonardi 1989)
57.2.
ESTADO DO SAO PAULO
,
BRAZIL
1. Botucatu Formation (Leonardi 1977,
1980a, 1981b, 1989; Leonardi and Godoy
1980)
Theropod tracks
?Ornithopod tracks
Age: Hettangian-Toarcian (=Liassic;
Leonardi 1989)
57.3.
ESTADO DO PARANÁ
,
BRAZIL
1. Botucatu Formation (Leonardi
1980a)
Theropod tracks
Age: Hettangian-Toarcian (=Liassic;
Leonardi 1989)
61.1.
ESTADO DO TÁCHIRA
,
VENEZUELA
1. La Quinta Formation (Russell
et al. 1992; Sanchez-Villagra and
Clark 1994)
Ornithischia indet.
(=Lesothosaurus?
sp. indet.)
Age: ?Early Jurassic (Schubert
1986)
Africa
Like the Upper Triassic of Africa, the Lower Jurassic is best known
from the Karroo Basin of southern Africa, primarily through the
efforts of J. Attridge, A J. Charig, A. W. Crompton, F. and P. Ellen-
berger, C. Gow, S. H. Haughton, K. A. Kermack, J. W. Kitching, and
E. C. N. van Hoepen. Additional information on the Lower Juras-
sic of Africa comes from work by J. Jenny and co-workers in Mo-
rocco and G. Bond, M. R. Cooper, and M. A. Raath in Zimbabwe.
Figures 24.10 and 24.11 provide information on these locations.
64.1.
PROVINCE DE MARRAKECH
,
MOROCCO
1. Aganane Formation (Jenny and Jossen
1982)
Theropod tracks
?Sauropod tracks
?Stegosaur tracks
Age: Pliensbachian (Jenny and Jossen
1982)
2. Unnamed unit (Jenny et al. 1980; Taquet
1985)
Theropoda indet.
Sauropoda indet.
Age: Toarcian (Jenny et al. 1980)
64.2.
PROVINCE D
’
OUARZAZATE
,
MOROCCO
1. Imi-n-Ifri Formation (Monbaron et al.
1985)
Ornithopod tracks
Age: Sinemurian (Monbaron et al.
1985)
2. Aganane Formation (Jenny and Jossen
1982)
Theropod tracks
?Sauropod tracks
?Stegosaur tracks
Age: Pliensbachian (Jenny and Jossen
1982)
64.3.
PROVINCE DE BENI MELLAL
,
MOROCCO
1. Aganane Formation (Jenny and Jossen
1982)
?Sauropod tracks
?Stegosaur tracks
Age: Pliensbachian (Jenny and Jossen
1982)
65.1.
WILAYA SAIDA
,
ALGERIA
1. Unnamed unit (Bassoulet 1971)
Theropod tracks
Age: ?Hettangian (Bassoulet 1971;
Taquet 1977)
77.1.
MASHONALAND NORTH
,
ZIMBABWE
1. Vulcanodon Beds (Raath 1972; Cooper
198 4)
Sauropoda
Vulcanodon karibaensis
Age: ?Hettangian (Raath 1972)
77.2.
MASHONALAND WEST
,
ZIMBABWE
1. Upper Karroo Sandstone (Raath et al.
1970 )
Prosauropoda
Massospondylus carinatus
Age: Early Jurassic (Raath et al. 1970)
77.3.
MATABELELAND NORTH
,
ZIMBABWE
1. Forest Sandstone (Raath 1969)
Theropoda
Ceratosauria
Syntarsus rhodesiensis
Prosauropoda
Massospondylus carinatus
Prosauropoda indet.
Age: Hettangian-Sinemurian
(Olsen and Galton
198 4)
80.1.
OTJIQARONGO
,
NAMIBIA
1. Etjo Sandstone (Huene 1925a; Gürich
1926; Rubidge, pers. comm.)
Theropod tracks
Undescribed prosauropod
Age: ?Sinemurian (=Clarens
Formation; Kitching,
pers. comm.)
81.1.
TRANSVAAL
,
SOUTH AFRICA
1. Bushveld Sandstone (Haughton
1924)
Prosauropoda
Massospondylus carinatus
Age: ?Sinemurian (?=Lower Clarens
Formation; Kitching, pers.
comm.)

81.2.
FREE STATE PROVINCE
,
SOUTH
AFRICA
1. Upper Elliot Formation (Haughton 1924;
Kitching 1979, 1981; Grine and Kitching
1987; Sereno 1991b; Zelenitsky and
Modesto 2002)
Theropoda
Ceratosauria
Syntarsus rhodesiensis
Prosauropoda
Massospondylus carinatus
Prosauropoda indet. (=Leptospondylus
capensis, Pachyspondylus
orpenii, Aetonyx palustris,
Gryponyx taylori, G. africanus,
Hortalosaurus skirtopodus,
Massospondylus browni,
Dromicosaurus gracilis)
Ornithopoda
Heterodontosauridae
Heterodontosaurus tucki
Abrictosaurus consors
Lycorhinus angustidens (=Lanasaurus
scalpridens)
Dinosaur eggs
Age: Hettangian-?Sinemurian (Olsen
and Galton 1984)
2. Clarens Formation (Haughton 1924)
Prosauropoda
Massospondylus carinatus
Prosauropoda indet.
(=Thecodontosaurus minor,
Aristosaurus erectus,
Thecodontosaurus dubius,
Gyposaurus capensis)
Ornithischia indet. (=Fabrosaurus sp.)
Ornithopoda
Heterodontosauridae
Heterodontosaurus tucki
Ornithopoda indet. (=Geranosaurus
atavus)
Age: Sinemurian (Olsen and Galton
198 4)
81.3.
CAPE PROVINCE
,
SOUTH AFRICA
1. Upper Elliot Formation (Haughton 1924;
Charig et al. 1965; Thulborn 1974; Gow
1975a; Hopson 1975a; Santa Luca et al.
1976; Raath 1980; Kitching and Raath
198 4)
Theropoda
Ceratosauria
Syntarsus rhodesiensis
Prosauropoda
Massospondylus carinatus
Massospondylus sp.
Prosauropoda indet. (including
Massospondylus browni,
M. harriesi, Thecodontosaurus
browni, T. minor,
Hortalotarsus skirtopodus,
Massospondylus schwarzi)
?Ornithischia indet.
Ornithopoda
Heterodontosauridae
Heterodontosaurus tucki
Lycorhinus angustidens
Abrictosaurus consors
Age: Hettangian-?Sinemurian (Olsen
and Galton 1984)
2. Clarens Formation (Haughton 1924;
Crompton and Charig 1962; Kitching
and Raath 1984)
Prosauropoda
Massospondylus carinatus
Prosauropoda indet. (=Hortalotarsus
skirtopodus)
Ornithopoda
Heterodontosauridae
Heterodontosaurus tucki
Heterodontosauridae indet.
(=Geranosaurus atavus)
Age: Sinemurian (Olsen and Galton
198 4)
82.1.
LERIBE DISTRICT
,
LESOTHO
1. Upper Elliot Formation (Haughton 1924;
Ellenberger 1970; Kitching and Raath
1984; Olsen and Galton 1984)
Theropod tracks
Prosauropoda
Massospondylus carinatus (=Gryponyx
sp., Aetonyx sp.)
Prosauropod tracks
Ornithischian tracks
Age: Hettangian-?Sinemurian (Olsen
and Galton 1984)
82.3.
MAFETENG DISTRICT
,
LESOTHO
1. Upper Elliot Formation (Ginsburg
1964; Galton 1978; Kitching and
Raath 1984; Olsen and Galton 1984;
Santa Luca 1984; Gauffre 1993c;
Knoll and Battail 2001; Knoll 2002a,
2002b)
Theropod tracks
Prosauropoda
cf. Massospondylus sp.
Melanorosaurus thabanensis
Prosauropod tracks
Ornithischia
Lesothosaurus diagnosticus
Unnamed ornithischian
Ornithischia indet. (=Fabrosaurus
australis)
Ornithischian tracks
Age: Hettangian-?Sinemurian
(Olsen and Galton
198 4)
82.4.
MOHALES HOEK DISTRICT
,
LESOTHO
1. Upper Elliot Formation (Ellenberger
1970; Kitching and Raath 1984; Olsen
and Galton 1984)
Theropod tracks
Prosauropoda indet.
(=cf. Gryponyx sp.)
Prosauropod tracks
?Ornithischia indet.
Ornithischian tracks
Age: Hettangian-?Sinemurian (Olsen
and Galton 1984)
82.5.
QACHAS NEK DISTRICT
,
LESOTHO
1. Upper Elliot Formation (Thulborn
1970a, 1974; Hopson 1975a; Kitching
and Raath 1984)
Prosauropoda
cf. Massospondylus sp.
Ornithopoda
Heterodontosauridae
Abrictosaurus consors (including
Lycorhinus consors)
Age: Hettangian-?Sinemurian (Olsen
and Galton 1984)
82.6.
QUTHING DISTRICT
,
LESOTHO
1. Upper Elliot Formation (Ellenberger
1970; Kitching and Raath 1984; Olsen
and Galton 1984)
Theropod tracks
Prosauropoda
Massospondylus carinatus (including
Gryponyx taylori)
Prosauropoda indet. (including
Thecodontosaurus browni)
Prosauropod tracks
Ornithopoda
Heterodontosauridae
Heterodontosaurus sp.
Age: Hettangian-?Sinemurian (Olsen
and Galton 1984)
536 DINOSAUR DISTRIBUTION AND BIOLOGY
Australasia
Two Early Jurassic dinosaur locations, both ichnofaunas, are
known from Queensland, Australia, through the work of J. A.
Long, T. H. V. Rich, R. A. Thulborn, and P. Vickers-Rich (fig. 24.12).
84.1.
QUEENSLAND
,
AUSTRALIA
1. Precipice Sandstone (Thulborn 1994;
Rich and Vickers-Rich 1997; Long 1998)
Theropod tracks
Ornithopod tracks
Age: Early Jurassic (Thulborn 1994)
2. Razorback Beds (Rich and Vickers-Rich
1997)
Theropod tracks
Ornithopod tracks
Age: Early Jurassic (Rich and Vickers-
Rich 1997)

Antarctica
From the heart of Antarctica, the Transantarctic Mountains have
yielded a surprisingly rich Early Jurassic dinosaur fauna, cur-
rently under study by W. R. Hammer and colleagues (fig. 24.1).
DINOSAUR DISTRIBUTION 537
86.0. (
NO ADMINISTRATIVE
-
LEVEL
DIVISIONS
)
1. Hanson Formation (Hammer and
Hickerson 1994, 1996)
Theropoda
Carnosauria
Cryolophosaurus ellioti
Allosauroidea indet.
Theropoda indet.
Prosauropoda indet.
Age: Sinemurian-Pliensbachian (Hammer
and Hickerson 1996)
Middle Jurassic
The fossil record of dinosaurs during the Middle Jurassic has im-
proved immensely over the past decade (they are now known
from nearly a hundred locations), although it still consists of
rather limited faunas. Scattered locations are now known on all
continents except Antarctica, but by far the best record comes
from Europe.
North America
The record of Middle Jurassic dinosaurs in North America (13 lo-
cations, an increase from 1) consists principally of ichnofaunas
from the southwestern United States and southern Mexico
(figs. 24.3, 24.13, and 24.14). These dinosaur occurrences come
to us principally from the research of A. P. Hunt, M. G. Lockley,
and S. G. Lucas in the southwestern United States and J. M.
Clark, D. E. Fastovsky, and I. Ferrusquia-Villafranca in central
Mexico.
3.1.
UTAH
,
UNITED STATES
1. Carmel Formation (Lockley and Hunt
1995a)
Theropod tracks
Age: Bajocian-Bathonian (Kowallis et al.
2001)
2. Entrada Sandstone (Lockley 1991;
Anderson and Lucas 1996; Lockley et al.
1996b)
FIGU RE 24.14. Middle Jurassic and Late Cretaceous localities in Mexico (4) and Honduras (5). Scale =600 km.

Theropod tracks
Age: late Bathonian–early Callovian
(Imlay 1952)
3. Summerville Formation (Gillette 1993;
Anderson and Lucas 1996; Lockley et al.
1996b, 2000)
Theropod tracks
Sauropoda incertae sedis
Dystrophaeus viaemalae
Age: Callovian-Oxfordian (Anderson
and Lucas 1996)
3.2.
ARIZONA
,
UNITED STATES
1. Undifferentiated Bluff-Summerville
Formation (Lockley et al. 1996b)
Theropod tracks
Sauropod tracks
Age: Callovian-Oxfordian (Anderson
and Lucas 1996)
3.3.
NEW MEXICO
,
UNITED STATES
1. Summerville Formation (Lockley et al.
1996b; Lucas et al. 1996)
Theropod tracks
Sauropoda indet. (=Camarasaurus sp.)
Age: Callovian-Oxfordian (Anderson
and Lucas 1992)
3.4.
COLORADO
,
UNITED STATES
1. Summerville Formation (Lockley et al.
1996b)
Theropod tracks
Age: Callovian-Oxfordian (Anderson
and Lucas 1996)
3.6.
OKLAHOMA
,
UNITED STATES
1. Bell Ranch Formation (Lockley et al.
1996b)
Theropod tracks
Age: Callovian-Oxfordian (Lockley et al.
1996b)
3.12.
WYOMING
,
UNITED STATES
1. Gypsum Spring Formation (Kvale et al.
2001)
Theropod tracks
Age: late Bajocian (Kvale et al. 2001)
2. Sundance Formation (Breithaupt et al.
1998; Kvale et al. 2001)
Theropod tracks
Age: middle-late Bathonian (Breithaupt
et al. 1998; Kvale et al. 2001)
4.1.
ESTADO DE MICHOACAN
,
MEXICO
1. Tecocoyunca Group (Ferrusquia-
Villafranca et al. 1978, 1980, 1996)
Theropod tracks
Sauropod tracks
Ornithopod tracks
Age: early Bajocian–early Bathonian
(Ferrusquia-Villafranca et al.
1978, 1980)
4.2.
ESTADO DE TAMAULIPAS
,
MEXICO
1. La Boca Formation (Clark et al.
1994a; Fastovsky et al. 1995;
Munter 1999)
Theropoda
Unnamed ceratosaur
Theropoda indet.
?Sauropoda indet.
Ornithischia indet.
Age: early Middle Jurassic (?Hettangian-
?Sinemurian; Fastovsky et al.
1995)
4.3.
ESTADO DE OAXACA
,
MEXICO
1. Zorrillo Formation (Ferrusquia-
Villafranca et al. 1995; Jiménez-Hidalgo
et al. 1998)
Theropod tracks
Sauropod tracks
Ornithopod tracks
Age: Bajocian-Bathonian (Jiménez-
Hidalgo et al. 1998)
4.4.
ESTADO DE PUEBLA
,
MEXICO
1. Unnamed unit (Ferrusquia-Villafranca
et al. 1995)
?Sauropoda indet.
Age: Jurassic (Ferrusquia-Villafranca
et al. 1995)
538 DINOSAUR DISTRIBUTION AND BIOLOGY
Europe
Middle Jurassic dinosaurs are best known from Europe (fig. 24.15;
54 locations, amounting to an increase of over 40%). Scottish
and English locations are known principally through the work
of W. Buckland, J. B. Delair, S. E. Evans, A. N. Leeds, D. M. Mar-
till, A. R. Milner, R. Owen, W. A. S. Sarjeant, H. G. Seeley, and
A. S. Woodward, and. French locations are best known through
the work of E. Buffetaut, F. de Broin, R. Hoffstetter, J. Kriwet and
collaborators, and J. Pivetaut. No less important, although rarer,
are the Portuguese, German, and Russian locations, which are
known through work by W. G. Kuhne, R. A. Thulborn, and V. F.
dos Santos; I. Michelis and collaborators; and V. P. Novikov and
co-workers, respectively.
7.2 .
SKYE
,
SCOTLAND
1. Valtos Sandstone Formation (Clark et al.
1995)
Sauropoda indet. (=?Cetiosauridae
indet.)
Age: Bathonian (Clark et al. 1995)
2. Ostracod Limestone (Great Estuarine
Series; Evans and Milner 1994)
Theropoda indet.
Age: middle Bathonian (Cope et al.
1980a)
7.3.
HIGHLAND
,
SCOTLAND
1. Lealt Shale Formation (Andrews and
Hudson 1984; Delair and Sarjeant 1985)
Ornithopod tracks
Age: Bathonian (Harris and Hudson
1980)
10.1.
SOMERSET
,
ENGLAND
1. Inferior Oolite (Woodward 1894; Benton
and Spencer 1995)
Theropoda indet. (=Megalosaurus sp.)
Age: Aalenian-Bajocian (Cope et al.
1980a)
10.3.
GLOUCESTERSHIRE
,
ENGLAND
1. Inferior Oolite (Lydekker 1888a;
Woodward 1894; Walker 1964; Benton
and Spencer 1995)
Theropoda indet. (=Megalosaurus
bucklandii, Megalosaurus sp.)
Age: Aalenian-Bajocian (Cope et al.
1980a)
2. Chipping Norton Formation (Galton
et al. 1980; Evans and Milner 1994;
Metcalf and Walker 1994; Benton and
Spencer 1995; Upchurch, pers. comm.)
Theropoda incertae sedis
Megalosaurus sp.
Theropoda indet. (=Megalosaurus sp.)
Sauropoda indet. (=Cetiosaurus sp.)
Ornithischia indet. (=cf. Alocodon sp.,
cf. Phyllodon sp.)
Stegosauria indet. (=Stegosaurus sp.,
?Lexovisaurus vetustus)
Age: early Bathonian (Cope et al. 1980a)
3. Cotswold Slate (Benton 1988; Evans
and Milner 1994; Benton and Spencer
1995)
Theropoda indet. (=Megalosaurus sp.)
Age: middle Bathonian (Cope et al.
1980a)
4. Forest Marble (Evans and Milner 1994;
Benton, pers. comm.)
Theropoda
Coelurosauria
Proceratosaurus bradleyi
Theropoda indet. (=Megalosaurus
bucklandii)
Sauropoda indet. (=Cetiosaurus
oxoniensis)
Ornithischia indet.
Age: late Bathonian (Cope et al. 1980a)
5. Great Oolite (Benton and Spencer 1995;
Upchurch, pers. comm.) (not plotted)
Sauropoda incertae sedis
Cardiodon rugulosus
Age: late Bathonian (Cope et al. 1980a)
10.4.
NORTH YORKSHIRE
,
ENGLAND
1. Saltwick Formation (Whyte and
Romano 1981, 1993, 2001; Romano et al.
1999; Romano and Whyte 2003)
Theropod tracks
Sauropod tracks
Stegosaur tracks

Ornithopod tracks
Age: Aalenian (Cope et al. 1980a)
2. Lower Deltaic Series (Sarjeant 1970; Hill
1976; Thulborn 1991)
Theropod tracks
Dinosaur coprolites
Age: late Aalenian (Sarjeant 1970)
3. Inferior Oolite (Owen 1840–45;
Brodrick 1909; Sarjeant 1970; Benton
and Spencer 1995; Upchurch and
Martin 2002)
Theropod tracks
Sauropoda indet. (including Cetiosaurus
hypoolithicus, C. longus,
C. medius)
Ornithopod tracks
Age: Aalenian-Bajocian (Cope et al.
1980a)
4. Upper Estuarine Series (Sarjeant 1970)
Theropod tracks
Age: early-middle Bathonian (Cope et al.
1980a)
10.6.
LEICESTERSHIRE
,
ENGLAND
1. Great Oolite (Phillips 1871; Benton and
Spencer 1995; Upchurch, pers. comm.)
Sauropoda indet. (Cetiosaurus sp.)
Age: late Bathonian (Benton and
Spencer 1995)
10.7.
DORSET
,
ENGLAND
1. Inferior Oolite (Huene 1923; Waldman
1974; Benton and Spencer 1995)
Theropoda incertae sedis
Magnosaurus nethercombensis
“Megalosaurus” hesperis
Stegosauria
Stegosauridae indet. (=“Stegosaurus” sp.)
Age: Aalenian-Bajocian (Cope et al. 1980a)
2. Forest Marble (Evans and Milner 1994)
Theropoda indet. (including
Megalosaurus sp.)
Ornithischia indet.
Age: late Bathonian (Cope et al. 1980a)
3. Lower Oxford Clay (Galton 1985c)
Stegosauria
Stegosauridae
Lexovisaurus durobrivensis
Age: middle Callovian (Cope et al.
1980a)
10.8.
NORTHAMPTONSHIRE
,
ENGLAND
1. Northampton Sands Formation (Reid
1984c)
Sauropoda indet. (=?Brachiosauridae
indet.)
Age: early Aalenian (Cope et al. 1980a)
DINOSAUR DISTRIBUTION 539
FIGU RE 24.15. Middle and Late Jurassic localities in Scotland (7), England (10), France (14), Spain (15), Por tugal (16), Germany (17), and Switzerland
(25). Position of England not to scale. Scale =600 km.

2. Inferior Oolite (Lydekker 1888b;
Woodward 1894)
Theropoda indet. (=?Megalosaurus
bucklandii)
Age: Aalenian-Bajocian (Cope et al. 1980a)
3. Great Oolite (Owen 1842a, 1842b;
Philips 1871; Benton and Spencer 1995;
Upchurch and Martin, 2003)
Sauropoda indet. (=Cetiosaurus oxoniensis)
Age: late Bathonian (Cope et al. 1980a)
4. Cornbrash Formation (Woodward 1894;
Benton and Spencer 1995; Upchurch
and Martin 2002)
Theropoda indet. (=Megalosaurus sp.)
Sauropoda indet. (=Cetiosaurus sp.)
Age: late Bathonian (Cope et al. 1980a)
10.9.
CAMBRIDGESHIRE
,
ENGLAND
1. Lower Oxford Clay (Hulke 1887;
Lydekker 1889b; Nopcsa 1911a; van
Straelen 1928; Charig 1980; Galton
1980c, 1983b, 1985c; Martill 1988;
Benton and Spencer 1995; Upchurch
1995; Upchurch and Martin 2002)
Sauropoda
Diplodocoidea incertae sedis
Cetiosauriscus stewarti
Sauropoda indet. (=Ornithopsis leedsi)
Stegosauria
Stegosauridae
Lexovisaurus durobrivensis
(=Stegosaurus priscus,
Omosaurus leedsi partim)
Ankylosauria incertae sedis
Sarcolestes leedsi
Ornithopoda
Euornithopoda indet.
?Iguanodontia indet. (=Camptosaurus
leedsi)
Dinosaur eggs
Age: middle Callovian (Martill 1988)
10.10.
BEDFORDSHIRE
,
ENGLAND
1. Oxford Clay (Lydekker 1888b; Benton
and Spencer 1995)
Stegosauria indet. (=?Lexovisaurus sp.)
Age: Callovian (Benton and Spencer 1995)
10.11.
OXFORDSHIRE
,
ENGLAND
1. Sharp’s Hill Formation (Phillips 1871;
Lydekker 1888a; Galton and Powell
1983; Galton 1985c; Boneham and
Forsey 1992; Evans and Milner 1994;
Benton and Spencer 1995; Upchurch,
pers. comm.)
Theropoda indet. (=Megalosaurus sp.)
Sauropoda indet. (=Cetiosaurus sp.)
Stegosauria
Stegosauridae indet. (=Omosaurus
vetustus)
Age: early Bathonian (Cope et al. 1980a)
2. Chipping Norton Formation (Owen
1842a; Phillips 1871; Hulke 1887; Galton
and Powell 1983; Benton and Spencer
1995; Upchurch, pers. comm.)
Theropoda indet. (=Megalosaurus sp.,
Streptospondylus cuvieri,
?Scrotum humanum)
Sauropoda indet. (including Cetiosaurus
sp., C. giganteus)
Stegosauria
Stegosauridae indet. (=Omosaurus
vetustus)
Age: late Callovian (Martill 1988); early
Bathonian (Cope et al.
1980a)
3. Hampen Marly Formation (Evans
and Milner 1994; Benton and Spencer
1995)
Sauropoda indet. (=Cetiosaurus sp.)
Age: middle Bathonian (Benton and
Spencer 1995)
4. Stonesfield Slate (Huxley 1869b; Huene
1926a, 1932; van Straelen 1928; Galton
1975, 1980c; Benton and Spencer 1995;
Upchurch, pers. comm.)
Theropoda
Megalosauridae
Megalosaurus bucklandii
Tetanurae incertae sedis
Iliosuchus incognitus
Sauropoda indet. (=Cetiosaurus
oxoniensis)
Ornithopoda
Euornithopoda indet.
?Dinosaur eggs
Age: middle Bathonian (Cope et al.
1980a)
5. White Limestone Formation (Day et al.
2002a, 2002b)
Theropod tracks
Sauropod tracks
Age: middle Bathonian (Cope et al.
1980a)
6. Forest Marble (Phillips 1871; Freeman
1976, 1979; Evans and Milner 1991,
1994; Benton and Spencer 1995;
Upchurch and Martin 2002)
Theropoda
Maniraptora indet.
Theropoda indet. (=Megalosaurus sp.)
Sauropoda
Cetiosauridae
Cetiosaurus oxoniensis
Diplodocoidea incertae sedis
“Cetiosaurus” glymptonensis
Sauropoda indet. (=Cetiosaurus sp.,
Bothriospondylus sp.)
Ornithischia indet. (=cf. Alocodon sp.)
Ornithopoda
Iguanodontia indet. (=?Iguanodon sp.)
Age: late Bathonian (Cope et al. 1980a)
7. Cornbrash Formation (Woodward 1894;
Galton and Powell 1983)
Theropoda indet. (=Megalosaurus
bucklandii)
Stegosauria
Stegosauridae indet. (=Omosaurus
vetustus)
Age: late Bathonian (Cope et al. 1980a)
8. Middle Oxford Clay (Martill 1988;
Benton and Spencer 1995)
Theropoda incertae sedis
Eustreptospondylus oxoniensis
Theropoda indet. (=Megalosaurus sp.)
Age: late Callovian (Martill 1988)
10.12.
RUTLAND
,
ENGLAND
1. “Rutland Clay” (Benton and Spencer
1995; Upchurch and Martin, in press)
Sauropoda
Cetiosauridae
Cetiosaurus oxoniensis
Age: late Bajocian (Upchurch and
Martin, 2002)
10.13.
BUCKINGHAMSHIRE
,
ENGLAND
1. Great Oolite (Phillips 1871; Woodward
1894)
Sauropoda indet. (=Cetiosaurus sp.)
Age: late Bathonian (Cope et al.
1980a)
2. Forest Marble (Delair and Sarjeant
1985)
Theropod tracks
Age: late Bathonian (Cope et al. 1980a)
10.14.
WILTSHIRE
,
ENGLAND
1. Inferior Oolite (Delair 1973; Upchurch,
pers. comm.)
Sauropoda indet. (=Cetiosaurus sp.)
Age: Aalenian-Bajocian (Cope et al.
1980a)
2. Great Oolite (Lydekker 1888a; Benton
and Spencer 1995; Upchurch, pers.
comm.)
Theropoda indet. (=Megalosaurus
bucklandii)
Sauropoda indet. (=Cetiosaurus sp.)
Age: late Bathonian (Cope et al. 1980a)
3. Forest Marble (Owen 1841, 1875; Delair
1973; Benton and Spencer 1995)
Theropoda indet. (=Megalosaurus sp.)
Sauropoda incertae sedis
Cardiodon rugulosus
Sauropoda indet. (=Bothriospondylus
robustus)
Ornithischia indet. (=?Hylaeosaurus sp.)
Age: late Bathonian (Cope et al. 1980a)
4. Oxford Clay (Delair 1973)
Sauropoda indet. (=Cetiosaurus sp.)
Age: Callovian (Cope et al. 1980)
14.4.
DÉPARTEMENT DU DOUBS
,
FRANCE
1. Unnamed unit (Broin et al. 1991)
Theropoda indet.
Age: Callovian (Broin et al. 1991)
2. Unnamed unit (Buffetaut 1994)
Theropoda
Megalosaurus sp.
Age: Callovian (Buffetaut 1994)
14.11.
DÉPARTEMENT DE L
’
AVEYRON
,
FRANCE
1. Unnamed unit (Kriwet et al. 1997)
Theropoda indet.
Euornithopoda indet.
Age: early Bathonian (Kriwet et al. 1997)
14.15.
DÉPARTEMENT DE LA HAUTE
-
SAÔNE
,
FRANCE
1. Unnamed unit (Buffetaut et al. 1995a)
Theropoda
Megalosaurus sp.
Age: Callovian (Buffetaut et al. 1995a)
540 DINOSAUR DISTRIBUTION AND BIOLOGY

14.16.
DÉPARTEMENT DE L
’
ORNE
,
FRANCE
1. Calcaire d’Ecouché (Rioult 1978;
Buffetaut et al. 1991)
Theropoda indet.
Age: early Bathonian (Rioult 1978;
Buffetaut et al. 1991)
14.17.
DÉPARTEMENT DU CALVADOS
,
FRANCE
1. Ecouche Limestone (Mercier 1937;
Buffetaut et al. 1991; Buffetaut 1994)
Theropoda indet. (=Megalosaurus sp.)
Age: early Bathonian (Buffetaut et al.
1991 )
2. Calcaire de Caen (Caumont 1828;
Eudes-Deslongchamps 1838; Gervais
1859; Rioult 1978; Buffetaut et al. 1991;
Allain 2002)
Theropoda
Megalosauridae
Poekilopleuron bucklandii
Unnamed eustreptospondyline
(=“Poekilopleuron”
valesdunensis)
Theropoda indet. (=Megalosaurus sp.)
Sauropoda indet.
Age: middle Bathonian (Buffetaut et al.
1991 )
3. Marnes à Belemnopsis latesulcata
Formation (Hoffstetter and Brun 1956;
Hoffstetter 1957; Buffetaut 1995b)
Sauropoda indet.
Stegosauria
Stegosauridae
Lexovisaurus durobrivensis
Age: early Callovian (Buffetaut et al.
1991 )
4. Marnes de Dives (Taquet and Welles
1977; Buffetaut et al. 1991; Buffetaut
and Enos 1992)
Theropoda
Tetanurae incertae sedis
Piveteausaurus divesensis
Theropoda indet.
Age: late Callovian (Taquet and Welles
1977; Buffetaut et al. 1991)
5. Unnamed unit (Gaudry 1890; Lapparent
1967; Taquet, pers. comm.)
Theropoda indet. (listed as Ornithopoda
indet. in Lapparent 1967)
Age: ?Middle Jurassic (Taquet, pers.
comm.)
6. Unnamed unit (Allain 2001) (not
plotted)
Theropoda
Tetanurae incertae sedis
Streptospondylus altdorfensis
Age: late Callovian or early Oxfordian
(Allain 2001)
14.18.
DÉPARTEMENT DE L
’
INDRE
,
FRANCE
1. Unnamed unit (Sauvage 1900; Buffetaut
et al. 1991; Buffetaut 1994)
Theropoda indet. (=Megalosaurus
bucklandii)
Sauropoda indet. (=Pelorosaurus sp.)
Age: Bathonian (Buffetaut 1994)
14.19.
DÉPARTEMENT DE LA CHARENTE
-
MARITIME
,
FRANCE
1. Unnamed unit (Lapparent 1967; no
further information on this entry, and it
may be in error)
Theropoda indet. (=Megalosaurus sp.)
Age: Bathonian (Lapparent 1967)
16.1.
DISTRITO DO LEIRIA
,
PORTUGAL
1. Unnamed unit (Thulborn 1973a)
Ornithischia indet. (=Alocodon kuehnei)
Age: late Callovian (Thulborn 1973a)
16.2.
DISTRITO DO SANTARÉM
,
PORTUGAL
1. Calcários Micriticos da Serra de Aire
(Santos et al. 1994)
Sauropod tracks
Age: late Bajocian–early Bathonian
(Santos et al. 1994)
17.8.
NORDRHEIN
-
WESTPHALEN
,
GERMANY
1. Unnamed unit (Michelis et al. 1996)
Stegosauria
Stegosauridae
Lexovisaurus sp.
Ornithopoda
Iguanodontia indet.
Age: middle Callovian (Michelis et al.
1996)
20.2.
WOJEWÓDZTWO KIELCE
,
POLAND
1. Gl ´nowaczów Formation (Gazdzicka et
al. 2001; Gierli´nski et al. 2001)
Theropod tracks
Stegosaur tracks
Euornithopod tracks
Age: early Kimmeridgian (Gazdzicka et
al. 2001; Gierli´nski et al.
2001)
37.1.
MOSKVA
,
RUSSIA
1. Unnamed unit (Novikov et al. 1998;
Alifanov 2000; Alifanov and Sennikov
2001)
Theropoda incertae sedis
cf. Ricardoestesia sp.
Theropoda indet. (including
Dromaeosauridae indet.)
Age: late Bathonian (Alekseev et al.
2001)
DINOSAUR DISTRIBUTION 541
Asia
Thirteen locations are known for the Middle Jurassic of Asia (a
threefold increase), most from the People’s Republic of China
(fig. 24.7). The faunas from Sichuan have yielded the most di-
verse, complete, and abundant dinosaur material known from
this timespan. The has the further advantage of currently being
exploited by Cai K.-J., He X.-L., and Peng G.-Z., as well as by
Dong Z.-M., P. J. Currie, Zhao X.-J., and other participants in the
Sino-Canadian Dinosaur Project. Although known only from a
brief mention by Wang S. and collaborators, the Xinjiang Uygur
Zizhiqu fauna is yielding new and important Middle Jurassic
dinosaur material. Farther to the west in Tadzikistan and Uzbek-
istan, L. K. Gabuniya, M. G. Lockley, and V. P. Novikov have re-
corded several new important ichnofaunas.
43.1.
VILOYATI LENINOBOD
,
TADZIKISTAN
1. Unnamed unit (Romanovsky 1882;
Lockley et al. 1996c)
Theropod tracks
Age: Bathonian (Lockley et al. 1996c)
43.2.
SOUTHWESTERN TADZIKISTAN
(
NO OBLAST
-
LEVEL DIVISION EXISTS
FOR THIS REGION
)
1. Zarbiz Svita (Novikov and Dzhalilov 1993)
?Euornithopod tracks
Age: Callovian-Oxfordian (Novikov
et al. 1993)
46.3.
GUJARAT
,
INDIA
1. Kaladongar Formation (Satyanarayana
et al. 1999)
Dinosauria indet.
Age: Aalenian-Bathonian
(Satyanarayana et al.
1999)
2. Patcham Formation (Ghevariya and
Srikarni 1994; Jana and Das 2002)
Sauropoda indet.
Dinosaur eggs
Age: Bathonian (Jana and Das 2002)
3. Chari Formation (Jana and Das 2002)
Sauropoda indet.
Age: Callovian (Jana and Das 2002)
48.1.
XINJIANG UYGUR ZIZHIQU
,
PEOPLE
’
S REPUBLIC OF CHINA
1. Dapuka Group (Wang et al. 1985; Zhao
unpubl. data)
Theropoda
Ceratosauria
Ngexisaurus dapukanensis
?Tetanurae
?Megalosaurus cachuensis
Undescribed sauropod
Undescribed ?stegosaur
Age: Middle Jurassic (Wang et al. 1985)

48.2.
SICHUAN
,
PEOPLE
’
S REPUBLIC
OF CHINA
1. Lower Shaximiao Formation (He 1979;
Dong et al. 1983; He and Cai 1984; He
et al. 1984; Zhang et al. 1984; Dong and
Tang 1985; Ouyang 1989; Peng and Shu
1992; Gao 1993; Zhang and Chen 1996)
Theropoda
Tetanurae
Xuanhanosaurus qilixiaensis
“Szechuanosaurus” zigongensis
Tetanurae incertae sedis
Kaijiangosaurus lini
Tetanurae indet.
(=Chuandongocoelurus
primitivus)
Avetheropoda incertae sedis
Gasosaurus constructus
Sauropoda
Eusauropoda
Shunosaurus lii
Omeisaurus junghsiensis
Omeisaurus tianfuensis
Mamenchisaurus fuxiensis
Macronaria
Abrosaurus dongpoi
Sauropoda incertae sedis
Datousaurus bashanensis
Sauropoda indet. (including
Protognathosaurus oxyodon,
Omeisaurus luoquanensis)
Ornithischia
Ornithischia indet. (=Xiaosaurus
dashanpensis)
Stegosauria
Huayangosaurus taibaii
Ornithopoda
Euornithopoda
Agilisaurus louderbacki
Agilisaurus multidens
Yandusaurus hongheensis
Age: Bathonian-Callovian (Dong and
Tang 1984); Bajocian (Chen
et al. 1982)
48.3.
YUNNAN
,
PEOPLE
’
S REPUBLIC
OF CHINA
1. Chuanjie Formation (Fang et al. 2000)
(not plotted)
Sauropoda incertae sedis
Chuanjiesaurus anaensis
Age: early Middle Jurassic (Fang et al.
2000)
48.5.
XINJIANG UYGUR ZIZHIQU
,
PEOPLE
’
S REPUBLIC OF CHINA
1. Wucaiwan Formation (Dong 1990a,
1992; Zhao 1993; Zhao and Currie
1993)
Theropoda
Carnosauria
Monolophosaurus jiangi
Sauropoda
Sauropoda incertae sedis
Klamelisaurus gobiensis
Macronaria
Bellusaurus sui
Stegosauria indet.
Ankylosauria indet.
Ornithopoda indet.
Age: Middle Jurassic (Zhao 1993)
2. Toutunhe Formation (Dong 1993d;
Maisch et al. 2003)
Theropoda
Ceratosauria indet.
Carnosauria indet.
?Thyreophora indet.
Ankylosauria indet. (=Tianchiasaurus
nedegoaperferima)
Age: Bathonian-Callovian (Eberth
et al. 2001a)
48.6.
SHAANXI
,
PEOPLE
’
S REPUBLIC
OF CHINA
1. Lower Chiloo Group (Zhen et al. 1989)
Theropod tracks
Age: Middle Jurassic (Zhen et al. 1989)
48.7.
JILIN
,
PEOPLE
’
S REPUBLIC
OF CHINA
1. Unnamed unit (Zhen et al. 1989)
Theropod tracks
Age: Middle Jurassic (Zhen et al. 1989)
542 DINOSAUR DISTRIBUTION AND BIOLOGY
South America
Work by A. Cabrera, R. M. Casamiquela, and, most recently, by
J. F. Bonaparte, R. A. Coria, and O. Gimenez has introduced a
few Middle Jurassic sites in South America. These are restricted
to northern Chile and southern Argentina (fig. 24.9).
62.1.
PROVINCIA DE ANTOFAGASTA
,
CHILE
1. Unnamed unit (Biese 1961; Casamiquela
1967a)
Theropoda indet.
Age: early Callovian (Biese 1961)
63.3.
PROVINCIA DE SANTA CRUZ
,
ARGENTINA
1. La Matilde Formation (Casamiquela
1964b, 1987)
Theropod tracks
Age: early-middle Callovian
(Casamiquela 1987)
63.4.
PROVINCIA DE CHUBUT
,
ARGENTINA
1. Cerro Carnerero Formation (Cabrera
1947; Casamiquela 1963)
Sauropoda
Eusauropoda incertae sedis
Amygdalodon patagonicus
Age: Bajocian (Bonaparte 1979a)
2. Cañadon Asfalto Formation (Bonaparte
1979b, 1986c; Coria 1989; Gímenez
1996; Rich et al. 1999b; Rauhut et al.
2001)
Theropoda
Tetanurae
Piatnitzkysaurus floresi
Undescribed coelurosaurian
Theropod tracks
Sauropoda
Sauropoda incertae sedis
Volkheimeria chubutensis
Eusauropoda
Tehuelchesaurus benitezii
Cetiosauridae
Patagosaurus fariasi
Undescribed sauropod
Age: Callovian (Bonaparte 1986c)
Africa
Six Middle Jurassic dinosaur locations are known from Africa
(figs. 24.10 and 24.11): five in Morocco and Algeria from the
early work of A. F. de Lapparent and the continuing studies of
S. Ishigaki, M. Monbaron, P. Taquet, and one in Madagascar from
the research of J. J. Flynn and collaborators.
64.3.
PROVINCE DE BENI MELLAL
,
MOROCCO
1. Guettioua Sandstone (Monbaron and
Taquet 1981; Monbaron et al. 1999)
Sauropoda
Macronaria
Atlasaurus imelakei (=Cetiosaurus
mogrebiensis)
Age: late Bathonian (Monbaron
and Taquet 1981;
Monbaron et al.
1999)

2. Unnamed unit (Bourcart et al. 1942;
Monbaron 1978; Debénath et al. 1979;
Taquet 1985)
Sauropoda
?Cetiosauridae indet.
Age: Bathonian (Taquet 1985)
64.4.
PROVINCE DE FES
,
MOROCCO
1. Unnamed unit (Lapparent and Lavocat
1955)
?Theropoda indet. (=Megalosaurus
mersensis; may be teleosaurid
crocodilian; Chabli 1985)
Sauropoda indet. (=Cetiosaurus
mogrebiensis)
Age: late Bathonian (Monbaron and
Taquet 1981)
64.5.
PRÉFECTURE DE AZILAL
,
MOROCCO
1. Iouaridène Formation (Ishigaki 1985,
1989)
Theropod tracks
Sauropod tracks
Age: Bathonian (Ishigaki 1995)
65.2.
WILAYA TLEMCEN
,
ALGERIA
1. Unnamed unit (Lapparent and Lucas
1957 )
Sauropoda indet.
Age: middle Callovian (Lapparent and
Lucas 1957)
83.2.
FARITANY MAJUNGA
,
MADAGASCAR
1. Isalo III Formation (Taquet 1977;
Bonaparte 1986b; Flynn et al. 1997)
Theropoda
?Allosauroidea indet.
Sauropoda
Titanosauriformes
Lapparentosaurus madagascariensis
Sauropoda indet. (=Bothriospondylus
madagascariensis)
Age: Bathonian (Taquet 1977)
DINOSAUR DISTRIBUTION 543
Australasia
Australasian dinosaur remains from the Middle Jurassic are
known from only four locations found in Australia, one in West-
ern Australia described by J. A. Long and three in Queensland
known through the research of R. E. Molnar (fig. 24.12).
84.1.
QUEENSLAND
,
AUSTRALIA
1. Hutton Sandstone Formation
(Longman 1926, 1927; Long 1998)
Sauropoda incertae sedis
Rhoetosaurus brownei
Age: Bajocian (Long 1998)
2. Walloon Group (Molnar 1982)
Theropod tracks
?Stegosaur tracks
Euornithopod tracks
Age: Middle Jurassic (Molnar 1982)
3. Unnamed unit (Molnar 1982; Long
1998)
Theropod tracks
Age: Middle Jurassic (Molnar
1982)
84.2.
WESTERN AUSTRALIA
,
AUSTRALIA
1. Colalura Sandstone (Long 1992, 1998;
Long and Molnar 1998)
Theropoda
Avetheropoda incertae sedis
Ozraptor subotaii
?Sauropoda indet.
Age: Bajocian (Long 1992)
Late Jurassic
More than any other interval of dinosaur history, the Late Ju-
rassic represents the public’s conception of dinosaur life: long-
necked sauropods, predatory theropods like Allosaurus, and
low-browsing stegosaurs and camptosaurs. These animals and
their relatives are known more extensively—from nearly 120
locations—during the Late Jurassic than previously, particularly
in North America but also in Europe, Asia, South America, and
Africa. There is no present record of Late Jurassic dinosaurs in
Australasia or Antarctica.
North America
Of the 12 North American locations known for the Upper Ju-
rassic, all but three are known from the Western Interior of the
United States and are restricted to the Morrison Formation (figs.
24.1, 24.3, and 24.13). Our knowledge of the Morrison fauna is
derived from the often arduous work of North America’s pio-
neers in vertebrate paleontology. Names such as J. B. Hatcher,
O. C. Marsh, and S. W. Williston; and E. D. Cope; E. Douglass
and C. W. Gilmore; H. F. Osborn and B. Brown; E. S. Riggs; and
J. W. Stovall are intimately associated with the exploitation of
the Upper Jurassic fossil fields of the western United States (see
Ostrom and McIntosh 1966 and Colbert 1968 for the history of
dinosaur discoveries in the Morrison Formation). Work on Late
Jurassic dinosaur faunas has continued unabated by a host of in-
vestigators, among them R. T. Bakker, G. Callison, K. Carpenter,
D. Chure, P. J. Currie, D. D. Gillette, J. R. Horner, J. A. Jensen, J. I.
Kirkland, M. G. Lockley, S. G. Lucas, .J. H. Madsen Jr., W. E. Miller,
W. L. Stokes, and T. E. Williamson.
1.3.
BRITISH COLUMBIA
,
CANADA
1. Mist Mountain Formation (Currie
1989a; McCrea et al. 2001b)
Theropod tracks (including avialan
tracks)
Ornithischian tracks
Age: late Tithonian–Berriasian (Currie
1989a)
3.1.
UTAH
,
UNITED STATES
1. Morrison Formation (Gilmore 1914a,
1936; Holland 1915b; Stokes 1944;
Ellinger 1950; Galton and Jensen 1973b;
Shepherd et al. 1973; Madsen 1974,
1976a, 1976b; Galton 1982a, 1983e;
Berman and McIntosh 1986; Chure and
Engelmann 1989; Hirsch et al. 1989;
Barnes and Lockley 1994; Chure 1994;
Lockley et al. 1994b, 1998c; Bray and
Hirsch 1998; Chure and Madsen 1998;
Turner and Peterson 1999; Madsen and
Welles 2000)
Theropoda
Ceratosauria
Ceratosaurus nasicornis

Ceratosaurus dentisulcatus
Ceratosaurus sp.
Tetanurae
Torvosaurus tanneri
Allosauroidea
Allosaurus fragilis
Unnamed Allosaurus species
Avetheropoda incertae sedis
Marshosaurus bicentesimus
Tyrannosauroidea
Stokesosaurus clevelandi
Coelurosauria incertae sedis
Coelurus fragilis
Ornitholestes hermanni
?Troodontidae indet. (=Koparion
douglassi)
Theropoda indet.
Theropod tracks
Sauropoda
Diplodocidae
Apatosaurus louisae
Barosaurus lentus
Diplodocus longus
Diplodocus carnegiei
Diplodocus sp.
?Diplodocidae indet.
Brachiosauridae
Brachiosaurus cf. altithorax
Camarasauromorpha
Camarasaurus lentus (including
C. annae, C. supremus)
Camarasaurus sp.
?Haplocanthosaurus sp.
Sauropoda indet.
Sauropod tracks
Ornithischia indet.
Stegosauria
Stegosauridae
Stegosaurus armatus
(=S. ungulatus)
Stegosaurus stenops
?Stegosaurus longispinus
Stegosaurus sp.
Stegosauridae indet.
(=Stegosaurus affinis)
Stegosaur tracks
Ornithopoda
Euornithopoda
Othnielia rex
Euornithopod tracks
Iguanodontia
Dryosaurus altus
Camptosaurus dispar
Camptosaurus sp.
Dinosaur eggs
Age: Kimmeridgian-Tithonian (Kowallis
et al. 1998)
3.2.
ARIZONA
,
UNITED STATES
1. Morrison Formation (Lockley et al.
1996c)
Theropod tracks
Age: Kimmeridgian-Tithonian (Kowallis
et al. 1998)
3.3.
NEW MEXICO
,
UNITED STATES
1. Morrison Formation (McIntosh 1981;
Rigby 1982; Gillette 1987, 1991; Hunt
and Lucas 1993b; Lucas 1993b; Lucas
et al. 1996; Williamson and Chure 1996;
Bray and Lucas 1997; Turner and
Peterson 1999)
Theropoda
Allosauroidea
Allosaurus fragilis
Allosauroidea indet.
Theropod tracks
Sauropoda
Camarasauromorpha
Camarasaurus cf. supremus
Camarasaurus sp.
Diplodocidae
Seismosaurus halli
Diplodocus carnegiei
Diplodocus sp.
Apatosaurus sp.
Stegosauria
Stegosauridae
Stegosaurus sp.
Ornithopoda
Iguanodontia indet.
Dinosaur eggs
Age: Kimmeridgian-Tithonian (Kowallis
et al. 1998)
3.4.
COLORADO
,
UNITED STATES
1. Entrada Sandstone (Eriksen 1979a, 1979b)
Theropod tracks
Age: Oxfordian (Hintze 1973)
2. Morrison Formation (Marsh 1884a,
1884d; Riggs 1901a, 1901b, 1903a,
1903b, 1904; Hatcher 1903c; Galton and
Jensen 1979b; Callison and Quimby
1984; Jensen 1985a, 1985b; Lockley
1986a; Lockley et al. 1986, 1994b;
Bartleson and Jensen 1988; McIntosh
and Williams 1988; Jensen and Padian
1989; Britt 1991; Miller et al. 1991;
Bakker et al. 1992; Alf and Carpenter
1993; Hirsch 1994; Kirkland and
Carpenter 1994; Lockley and Hunt
1995a; Alf 1998; Bray and Hirsch 1998;
Carpenter and Tidwell 1998; Chin and
Kirkland 1998; Harris 1998b; Turner and
Peterson 1999; Madsen and Welles 2000;
Brill and Carpenter 2001; Chure 2001a)
Theropoda
Ceratosauria
Ceratosaurus nasicornis
Ceratosaurus magnicornis
Ceratosaurus sp.
Elaphrosaurus sp.
Allosauroidea
Allosaurus fragilis (including
Labrosaurus ferox)
Tetanurae
Torvosaurus tanneri
Avetheropoda incertae sedis
Marshosaurus bicentesimus
Coelurosauria
cf. Coelurus sp.
?Dromaeosauridae (=Palaeopteryx
thomsoni)
Theropod tracks
Sauropoda
Sauropoda incertae sedis
“Morosaurus” agilis
Diplodocoidea incertae sedis
Amphicoelias altus
Diplodocidae
Diplodocus longus
Diplodocus lacustris
Diplodocus sp.
Barosaurus sp.
Apatosaurus excelsus (=Brontosaurus
amplus)
Apatosaurus ajax
Apatosaurus sp.
Supersaurus vivianae (including
Ultrasauros macintoshi
partim)
Brachiosauridae
Brachiosaurus altithorax
Brachiosaurus sp. (including
Ultrasauros macintoshi
partim)
Dystylosaurus edwini
Camarasauromorpha
Camarasaurus supremus
Camarasaurus grandis
Camarasaurus lewisi
Camarasaurus sp.
Haplocanthosaurus priscus
Haplocanthosaurus delfsi
Sauropoda indet. (=Titanosaurus
montanus)
Sauropod tracks
Ornithischia indet.
Stegosauria
Stegosauridae
Stegosaurus armatus
Stegosaurus stenops
Stegosaurus sp.
Stegosauridae indet. (=Hypsirhophus
discurus)
Ankylosauria incertae sedis
Mymoorapelta maysi
Ornithopoda
Euornithopoda
Othnielia rex
Euornithopoda indet. (including
Nanosaurus agilis, Tichosteus
aequifacies, T. lucasanus)
Euornithopod tracks
Iguanodontia
Dryosaurus altus
Camptosaurus dispar
Undescribed iguanodontian
Dinosaur coprolites
Dinosaur eggs
Age: Kimmeridgian-Tithonian (Kowallis
et al. 1998)
3.5.
TEXAS
,
UNITED STATES
1. ?Morrison Formation (McIntosh 1981)
Sauropoda
Camarasauromorpha
Camarasaurus sp.
Age: Kimmeridgian-Tithonian (Kowallis
et al. 1998)
3.6.
OKLAHOMA
,
UNITED STATES
1. Morrison Formation (Stovall 1938;
Langston 1974; Lockley 1986a;
Carpenter and McIntosh 1994; Chure
1996; Turner and Peterson 1999; Madsen
and Welles 2000)
544 DINOSAUR DISTRIBUTION AND BIOLOGY

Theropoda
Allosauroidea
Allosaurus fragilis
Saurophaganax maximus
Theropoda indet. (=Ceratosaurus sp.)
Sauropoda
Camarasauromorpha
Camarasaurus sp.
Diplodocidae
Apatosaurus excelsus
Apatosaurus sp.
Sauropod tracks
Stegosauria
Stegosauridae
Stegosaurus sp.
Ornithopoda
Iguanodontia
Camptosaurus dispar
Euornithopod tracks
Age: Kimmeridgian-Tithonian (Kowallis
et al. 1998)
3.11.
MONTANA
,
UNITED STATES
1. Morrison Formation (McIntosh 1981;
Horner 1989; Curry 1994; Turner and
Peterson 1999)
Sauropoda
Diplodocidae
Diplodocus sp.
Diplodocidae indet.
Camarasauridae
Camarasaurus grandis
Camarasaurus sp.
Sauropoda indet.
Ornithischia indet.
Stegosauria
Stegosauridae indet.
Age: Kimmeridgian-Tithonian (Kowallis
et al. 1998)
3.12.
WYOMING
,
UNITED STATES
1. Morrison Formation (Marsh 1878a,
1878b; Mook 1917a; Brown 1935;
Ostrom and McIntosh 1966; Galton
1982c; Bakker et al. 1992; Filla and
Redman 1994; Bakker 1996, 1998a,
1998b; McIntosh et al. 1996<1996a>;
Makovicky 1997a; Carpenter et al. 1998,
2001b; Kirkland et al. 1998c; Lockley et
al. 1998c; Miles et al. 1998; Britt et al.
1999; Turner and Peterson 1999)
Theropoda
Ceratosauria
Ceratosaurus sp. (including
Labrosaurus sulcatus)
Tetanurae
Torvosaurus tanneri (=Edmarka rex)
Tetanurae indet. (=Coelurus
gracilis)
Allosauroidea
Allosaurus fragilis (including
Labrosaurus lucaris)
Coelurosauria incertae sedis
Coelurus fragilis
Ornitholestes hermanni
Theropod tracks
Sauropoda
Sauropoda incertae sedis
“Apatosaurus” minimus
Camarasauromorpha
Camarasaurus grandis
Camarasaurus supremus
Camarasaurus lentus
?Eobrontosaurus yahnahpin
cf. Haplocanthosaurus priscus
Diplodocidae
Diplodocus carnegiei
Diplodocus hayi
Diplodocus sp.
Apatosaurus excelsus
Apatosaurus sp.
Sauropod tracks
Stegosauria
Stegosauridae
Hesperosaurus mjosi
Stegosaurus armatus (=S. ungulatus)
Stegosaurus stenops
Stegosaurus longispinus
Stegosaurus sp.
Stegosauria tracks
Ankylosauria
Ankylosauridae
Gargoyleosaurus parkpini
Undescribed ankylosaurian
Ornithopoda
Euornithopoda
Othnielia rex
Drinker nisti
Euornithopoda indet.
(=Nanosaurus agilis,
Laosaurus celer)
Iguanodontia
Dryosaurus altus
Camptosaurus dispar
Camptosaurus amplus
?Camptosaurus cf. depressus
Iguanodontia tracks
Dinosaur eggs
Age: Kimmeridgian-Tithonian (Kowallis
et al. 1998)
3.15.
SOUTH DAKOTA
,
UNITED STATES
1. Morrison Formation (Marsh 1899; Lull
1919; Foster 1996; Turner and Peterson
1999)
Theropoda
Allosauroidea
Allosaurus fragilis
Theropod tracks
Sauropoda
Camarasauromorpha
Camarasaurus sp.
Diplodocidae
Apatosaurus sp.
Barosaurus lentus
Age: Kimmeridgian-Tithonian (Kowallis
et al. 1998)
3.33.
ALASKA
,
UNITED STATES
1. Naknek Formation (Rich et al.
1997)
Theropod tracks
Age: middle Kimmeridgian–late
Tithonian (Imlay 1952;
Rich et al. 1997)
DINOSAUR DISTRIBUTION 545
Europe
Abundant yet scattered locations pertaining to the Late Jurassic
are known throughout Europe (64 locations, an increase of
42%; fig. 24.15). English faunas were originally revealed by J. W.
Hulke, A. Leeds, R. Lydekker, R. Owen, H. G. Seeley, and, most
recently, J. B. Delair and P. M. Galton. A. Bidar and co-workers,
E. Buffetaut, J. Kriwet, J.-M. Mazin, F. Nopcsa, and E. Sauvage,
among others, have been the principal researchers on the Late
Jurassic faunas of France. Late Jurassic Portuguese dinosaurs has
been investigated by W. G. Kuhne, A. F. de Lapparent, M. Telles
Antunes, R. A. Thulborn, and G. Zbyszewski as well as a host of
current workers who have documented a rich track and skeletal
record, including P. Dantas, V. F. dos Santos, M. G. Lockley,
O. Mateus. O. W. M. Rauhut, and J. Zinke. Thanks to C. A. Meyer,
the dinosaurs of the Late Jurassic from Switzerland are becom-
ing well known, and particularly their track record. Finally, the
Late Jurassic dinosaurs of Germany are known through the work
of H. Friese, M. Kaever, and A. F. de Lapparent.
10.4.
NORTH YORKSHIRE
,
ENGLAND
1. Corallian Oolite Formation (Fox-
Strangways 1892; Galton 1985c)
Theropoda indet. (=Megalosaurus
bucklandii)
Stegosauria
Stegosauridae indet. (=Omosaurus
phillipsi)
Age: early-middle Oxfordian (Cope et al.
1980a)
2. ?Lower Calcareous Grit (Seeley 1875;
Galton 1980e)
Ankylosauria
Nodosauridae indet. (=Iguanodon
phillipsii)
Age: ?early Oxfordian (Cope et al. 1980a)
10.7.
DORSET
,
ENGLAND
1. Corallian Oolite Formation (Lydekker
1890a; Huene 1923; Delair 1959; Walker
1964; Galton 1983e; Benton and Spencer
1995)
Theropoda
Tetanurae incertae sedis
Metriacanthosaurus parkeri

?Theropoda indet. (=?Cryptodraco sp.)
Age: early-middle Oxfordian (Cope et al.
1980a)
2. Kimmeridge Clay (Hulke 1874a;
Hoffstetter 1957; Galton 1975; H. P.
Powell 1987; Brokenshire and Clarke
1994; Benton and Spencer 1995; Galton
1999b; Upchurch, pers. comm.)
Theropoda
Undescribed theropod
Ornithomimidae indet.
Theropoda indet. (=Megalosaurus sp.)
Sauropoda
Unnamed brachiosaurid
(=“Cetiosaurus”
humerocristatus)
Sauropoda indet. (=Ischyrosaurus
manseli, Cetiosaurus sp.,
Ornithopsis leedsi)
Stegosauria
Stegosauridae
Dacentrurus armatus
Stegosauridae indet.
Ornithopoda
Euornithopoda indet.
(=cf. Bugenasaura sp.)
Age: Kimmeridgian (Cope et al.
1980a)
3. Portland Stone (Delair 1958, 1991,
1993)
Theropoda indet. (=Megalosaurus sp.)
Sauropoda indet. (=Ornithopsis sp.)
Age: late Tithonian (=middle
Portlandian; Cope et al.
1980a)
10.9.
CAMBRIDGESHIRE
,
ENGLAND
1. Ampthill Clay (Seeley 1869; Galton
1980c)
Ankylosauria indet. (=Cryptosaurus
eumerus)
Age: late Oxfordian (Cope et al. 1980a)
2. Kimmeridge Clay (Seeley 1869; Lydekker
1889b, 1890a; Galton 1985c; Benton and
Spencer 1995; Upchurch, pers. comm.)
Sauropoda
Unnamed brachiosaurid
(=“Cetiosaurus”
humerocristatus)
Sauropoda indet. (=Gigantosaurus
megalonyx, Ornithopsis sp.)
Stegosauria
Stegosauridae
Dacentrurus armatus
Age: Kimmeridgian (Cope et al. 1980a)
10.11.
OXFORDSHIRE
,
ENGLAND
1. Corallian Oolite Formation (Benton and
Spencer 1995)
Theropoda indet. (=Megalosaurus sp.)
Sauropoda indet. (=Cetiosaurus sp.)
Age: early-middle Oxfordian (Cope et al.
1980a)
2. Kimmeridge Clay (Galton and Powell
1980)
Ornithopoda
Iguanodontia
Camptosaurus prestwichii
Age: Kimmeridgian (Cope et al. 1980a)
3. Portland Stone (Phillips 1871;
Delair 1993; Benton and Spencer
1995)
Sauropoda indet. (=Cetiosaurus longus,
C.?longus, C. sp.)
Age: early Tithonian (=early
Portlandian; Cope et al.
1980b)
10.13.
BUCKINGHAMSHIRE
,
ENGLAND
1. Portland Stone (Delair 1993)
Sauropoda indet. (=Cetiosaurus longus)
Age: early Tithonian (=early
Portlandian; Cope et al.
1980b)
2. Purbeck Limestone Formation
(Huddleston 1887; Lydekker 1893b;
Woodward 1895; Delair 1993; Benton
and Spencer 1995; Norman and Barrett,
2003)
Theropoda indet. (=Megalosaurus sp.)
Sauropoda indet. (=Pelorosaurus
humerocristatus)
Ornithopoda
Iguanodontia indet.
Age: Tithonian (Allen and Wimbledon
1991 )
10.14.
WILTSHIRE
,
ENGLAND
1. Kimmeridge Clay (Owen 1877a, 1877b;
Galton 1975, 1980c, 1980d, 1985c;
Taylor et al. 1993; Upchurch,
pers. comm.)
Theropoda indet. (=?Megalosaurus
insignis)
Sauropoda indet. (=Bothriospondylus
suffosus)
Thyreophora indet.
Stegosauria
Stegosauridae
Dacentrurus armatus (including
Omosaurus armatus,
O. hastiger)
Stegosauridae indet.
Ankylosauria
Nodosauridae indet.
Age: early Kimmeridgian (Cope et al.
1980a)
2. Portland Stone (Delair 1993; Benton and
Spencer 1995)
Theropoda indet. (=Megalosaurus sp.)
Sauropoda indet. (=Camarasaurus sp.,
Diplodocus sp.)
Ornithischia incertae sedis
Echinodon sp.
Ornithischia indet.
Undescribed thyreophoran
Stegosauria indet.
Ankylosauria
Nodosauridae indet.
Age: early Tithonian (=early
Portlandian; Cope et al.
1980b)
10.15.
NORFOLK
,
ENGLAND
1. Kimmeridge Clay (Benton and Spencer
1995)
?Sauropoda indet. (=?Ornithopsis sp.)
Age: Kimmeridgian (Cope et al. 1980a)
10.16.
BERKSHIRE
,
ENGLAND
1. Corallian Oolite Formation (Benton and
Spencer 1995)
Undescribed dinosaur
Age: early-middle Oxfordian (Cope et al.
1980a)
14.2.
DÉPARTEMENT DE LA HAUTE
-
MARNE
,
FRANCE
1. Unnamed unit (Buffetaut 1990;
Buffetaut et al. 1991)
Sauropoda indet. (=?Camarasauridae
indet.)
Age: late Tithonian (Buffetaut et al.
1991 )
14.4.
DÉPARTEMENT DU DOUBS
,
FRANCE
1. Unnamed unit (Buffetaut 1995b)
Theropoda indet.
Age: Oxfordian (Buffetaut 1995b)
14.5.
DÉPARTEMENT DU JURA
,
FRANCE
1. Unnamed unit (Lapparent 1943; Enay
1980)
Theropoda indet. (=?Megalosaurus
insignis)
Sauropoda indet. (=?Bothriospondylus
madagascariensis)
Age: late Oxfordian (Buffetaut 1988)
14.6.
DÉPARTEMENT DE L
’
AIN
,
FRANCE
1. Calcaires Lithographiques de Cerin
(Bernier 1984; Bernier et al. 1984)
Theropod tracks
Age: late Kimmeridgian (Bernier 1984;
Bernier et al. 1984)
14.8.
DÉPARTEMENT DU VAR
,
FRANCE
1. Lithographic Limestone (Bidar et al.
1972 )
Theropoda
Compsognathidae
Compsognathus longipes
(=C. corallestris)
Age: late Tithonian (Buffetaut et al.
1991 )
14.17.
DÉPARTEMENT DU CALVADOS
,
FRANCE
1. Unnamed unit (Douville 1885;
Lapparent 1967)
Theropoda indet. (=Streptospondylus
cuvieri)
Age: Kimmeridgian (Buffetaut et al.
1991 )
2. Les Sables de Glos (Buffetaut et al.
1985c)
Theropoda indet.
Sauropoda indet. (=?Camarasauridae
indet.)
Age: late Oxfordian (Buffetaut et al.
1985c)
3. Unnamed unit (Buffetaut et al. 1991)
Theropoda indet.
Age: Oxfordian (Buffetaut et al. 1991)
546 DINOSAUR DISTRIBUTION AND BIOLOGY

14.19.
DÉPARTEMENT DE LA CHARENTE
-
MARITIME
,
FRANCE
1. Unnamed unit (Lapparent and Oulmi
196 4)
Theropoda indet. (=Megalosaurus sp.)
Age: late Tithonian (=Portlandian;
Lapparent 1967)
2. Unnamed unit (Lapparent and Oulmi
196 4)
Theropod tracks
Age: late Tithonian (Lapparent and
Oulmi 1964; Buffetaut et al.
1991 )
14.20.
DÉPARTEMENT DE LA CHARENTE
,
FRANCE
1. Unnamed unit (Vignaud et al. 1994;
Le Loeuff et al. 1996)
Theropoda indet.
Sauropoda indet. (=Camarasauridae
indet.)
Age: Tithonian (Le Loeuff et al. 1996)
14.21.
DÉPARTEMENT DE LA SEINE
-
MARITIME
,
FRANCE
1. Argiles d’Octeville (Nopcsa 1911b;
Lapparent 1967; Galton and Boiné 1980;
Galton 1991)
Sauropoda indet. (=Pelorosaurus sp.)
Stegosauria
Stegosauridae
Dacentrurus armatus (=D. lennieri)
Age: late Kimmeridgian (Buffetaut et al.
1991 )
2. Unnamed unit (Eudes-Deslongchamps
1870; Buffetaut et al. 1991)
Theropoda indet. (=Megalosaurus
insignis)
?Sauropod indet.
Age: early Kimmeridgian (Buffetaut et al.
1991 )
3. Marnes de Bléville (Buffetaut and
Cacheleux 1997)
Ornithopoda
Iguanodontia
Dryosaurus sp.
Age: early Kimmeridgian (Rioult 1980)
14.22.
DÉPARTEMENT DU PAS
-
DE
-
CALAIS
,
FRANCE
1. Unnamed unit (Sauvage 1888, 1897;
Galton and Powell 1980; Buffetaut and
Martin 1993)
Theropoda
?Ceratosauria
?Elaphrosaurus sp.
Theropoda indet. (including
?Megalosaurus insignis,
Teinurosaurus sauvagei)
Sauropoda indet. (=Iguanodon praecursor,
Morinosaurus typus)
Ornithopoda
Iguanodontia indet.
Age: Kimmeridgian (Galton 1982c;
Buffetaut et al. 1991)
2. Argiles de Chatillon Formation (Buffetaut
and Martin 1993; Buffetaut 1995b)
Theropoda indet.
Sauropoda indet.
?Stegosauria indet.
Age: late Kimmeridgian (Buffetaut
1995b)
3. Unnamed unit (Cuny et al. 1991)
Theropoda indet.
Sauropoda indet. (Pelorosaurus sp.)
Ankylosauria indet. (=?Nodosauridae
indet.)
Ornithopoda indet.
Age: late Tithonian (Cuny et al. 1991)
4. Sables et Grès à Trigonia gibbosa (Bonte
et al. 1985; Cuny et al. 1991; Buffetaut
and Martin 1993; Buffetaut 1995b)
Theropoda
Tetanurae
Megalosaurus sp.
Theropoda indet.
Sauropoda indet. (=Neosodon sp.)
?Ornithischia indet.
Ankylosauria
Nodosauridae indet.
Ornithopoda
Iguanodontia indet. (=Camptosaurus
cf. prestwichii)
Age: late Tithonian (Cuny et al. 1991)
5. Unnamed unit (Sauvage 1876b;
Moussaye 1884; Lapparent 1967;
Buffetaut et al. 1991)
Theropoda indet. (including
Megalosaurus insignis,
M. oweni)
Sauropoda indet. (=“Cetiosaurus”
humerocristatus)
Ornithopoda
Iguanodontia
Camptosaurus cf. prestwichii
Age: late Tithonian (Buffetaut et al.
1991 )
14.23.
DÉPARTEMENT DE LA SARTHE
,
FRANCE
1. Sables de Cherré (Bigot 1904; Buffetaut
et al. 1991)
Sauropoda indet.
Age: late Oxfordian (Buffetaut et al.
1991 )
14.24.
DÉPARTEMENT DE LA VIENNE
,
FRANCE
1. Unnamed unit (Lapparent 1967; dubious
occurrence)
Theropoda indet. (=Megalosaurus sp.)
Age: Oxfordian (Lapparent 1967)
14.25.
DÉPARTEMENT DE LA NIÈVRE
,
FRANCE
1. Unnamed unit (Lapparent 1967)
?Theropoda indet. (=Megalosaurus sp.;
may be sauropterygian;
Buffetaut et al. 1991)
Age: Kimmeridgian (Lapparent 1967)
14.26.
DÉPARTEMENT DE LA MEUSE
,
FRANCE
1. Unnamed unit (Huene and Maubeuge
1954; Galton and Powell 1980)
Theropoda indet.
Sauropoda indet.
Ornithopoda
Iguanodontia indet.
Age: late Oxfordian (Buffetaut et al.
1991 )
14.27.
DÉPARTEMENT DU LOT
,
FRANCE
1. Unnamed Unit (Mazin et al. 1997)
Theropod tracks
Sauropod tracks
Euornithopod tracks
Age: early Tithonian (Mazin et al. 1997)
15.1.
PROVINCIA DE SORIA
,
SPAIN
1. Oncala Group (Aguirrezabala and Viera
1980, 1983; Sanz et al. 1992; Sanz Pérez
1993; Casanovas-Cladellas et al. 1995c;
Fuentes Vidarte 1996, Moratalla 2000b,
2000d, 2000f, 2000h)
Theropod tracks (including avialan
tracks)
Sauropoda
Sauropoda indet.
Sauropod tracks
Ornithopoda
Euornithopoda
Hypsilophodon sp.
Euornithopod tracks
Age: Tithonian-Berriasian (Wright et al.
1998)
15.2.
PROVINCIA DE ASTURIAS
,
SPAIN
1. Lastres Formation (Egozcue 1873;
Garcia-Ramos and Valenzuela 1977,
1981; Mensink and Mertmann 1984;
Valenzuela et al. 1988; Lires 2000; Lires
et al. 2000, 2001a; Martínez et al. 2000a,
2000b, 2001; Piñuela 2000; García-
Ramos et al. 2001)
Theropoda indet. (including
Megalosaurus sp.)
Theropod tracks (including ?avialan
tracks)
Sauropoda
Camarasauridae indet.
?Diplodocidae indet.
Sauropod tracks
Thyreophoran tracks
Euornithopod tracks
Age: Kimmeridgian (García-Ramos and
Valenzuela 1981)
2. Tereñes Formation (García-Ramos et al.
2001; Lires et al. 2001b)
Theropod tracks
Sauropod tracks
Euornithopod tracks
Age: Kimmeridgian (Martínez et al.
2001)
3. Vega Formation (Lires 2000; Martínez
et al. 2000b, 2000c, 2001; Piñuela
2000)
Theropoda indet. (including
Megalosaurus sp.)
Theropod tracks
Sauropoda
?Camarasauridae indet.
Sauropod tracks
DINOSAUR DISTRIBUTION 547

Age: Kimmeridgian (Martínez et al.
2000c)
4. Unnamed unit (Royo y Gómez and
Gómez Llueca 1927)
Dinosauria indet.
Age: Kimmeridgian (Martínez et al.
2001)
15.3.
PROVINCIA DE VALENCIA
,
SPAIN
1. Unnamed unit (Lapparent et al. 1965)
?Theropod tracks
Age: Kimmeridgian (Lapparent et al.
1965)
2. Unnamed unit (Casanovas et al. 2001)
Sauropoda
Diplodocoidea incertae sedis
Losillasaurus giganteus
Age: Late Jurassic–Early Cretaceous
(Casanovas et al. 2001)
15.4.
PROVINCIA DE TERUEL
,
SPAIN
1. Higueruelas Formation (Pérez-Lorente
and Romero-Molina 2001)
Theropod tracks
Age: Tithonian (Pérez-Lorente and
Romero-Molina 2001)
2. Higueruelas or Villar del Arzobispo
Formation (Blanco et al. 2000)
Sauropod tracks
Age: Tithonian (Blanco et al. 2000)
3. Villar del Arzobispo Formation (Pérez
Onata et al. 1994; Alcalá and Martín
1995; Canudo et al. 1996c, 1997; Ruiz
Omeñaca et al. 1998a, 1998b; Barco
1999; Canudo et al. 1999; Barco and
Ruiz-Omeñaca 2001; Pérez-Lorente and
Romero-Molina 2001; Pérez-Lorente
et al. 2001)
Theropoda
Theropoda indet.
Coelurosauria indet.
Sauropoda
Diplodocidae indet.
Sauropoda indet.
Sauropod tracks
Ornithopoda
Euornithopod tracks
Age: Tithonian-Berriasian (Cuenca-
Bescós et al. 1999a, 1999b;
Pérez-Lorente and Romero-
Molina 2001)
16.1.
DISTRITO DO LEIRIA
,
PORTUGAL
1. Camadas de Montejunto or Cabaços
(Lapparent and Zbyszewski 1957)
Theropoda
Tetanurae indet. (=?Megalosaurus
insignis)
Age: middle-late Oxfordian (Manuppela
et al. 2000)
2. Camadas de Montejunto (Lapparent
and Zbyszewski 1957; Lockley et al.
1998b)
Theropoda indet. (=Megalosaurus
insignis)
Theropod tracks
Age: late Oxfordian (Manuppela et al.
2000)
3. Camadas de Alcobaça (Sauvage 1897–98;
Lapparent and Zbyszewski 1957;
Pérez-Moreno et al. 1999; Mateus,
pers. comm.)
Theropoda
Allosauroidea
Allosaurus sp.
Sauropoda indet.
Stegosauria
?Stegosauridae indet. (=Iguanodon
prestwichi)
Dinosaur tracks
Age: Kimmeridgian (Manuppela et al.
2000)
4. Camadas de Guimarota (Thulborn
1973a; Zinke and Rauhut 1994; Weigert
1995; Zinke 1998; Carpenter 1999;
Rauhut 2001, 2003; Mateus,
pers. comm.)
Theropoda
Ceratosauria indet.
?Allosauroidea indet.
Compsognathidae
Compsognathus sp.
Tyrannosauroidea
Aviatyrannus jurassica
Stokesosaurus sp.
Tyrannosauridae indet.
Maniraptora incertae sedis
cf. Ricardoestesia sp.
?Troodontidae indet. (including cf.
Paronychodon sp.)
Dromaeosauridae indet.
Avialae
Archaeopterygidae
Archaeopteryx sp.
Sauropoda indet. (including
?Bothriospondylus sp.)
Ornithopoda
Euornithopoda indet. (=Phyllodon
henkeli)
Iguanodontia indet.
Dinosaur eggs
Age: Kimmeridgian (Schudack 2000)
5. Unidade Amoreira-Porto Novo
(Lapparent and Zbyszewski 1957;
Mateus, pers. comm.)
Theropoda
Carnosauria
Lourinhanosaurus antunesi
Theropoda indet.
Sauropoda
Eusauropoda
Lourinhasaurus alenquerensis
Sauropoda indet.
Stegosauria
Stegosauridae
Dacentrurus armatus
Dacentrurus sp.
Ornithopoda indet.
Age: late Kimmeridgian–Tithonian
(Manuppela et al. 2000)
6. Unidade Sobral (Mateus, pers. comm.)
Stegosauria
Stegosauridae
Dacentrurus armatus
Ornithopoda
?Dryosauridae indet.
Age: late Kimmeridgian–Tithonian
(Manuppela et al. 2000)
7. Unidade Bombarral (Sauvage 1897;
Lapparent and Zbyszewski 1957;
Mateus, pers. comm.)
Theropoda
Tetanurae
Torvosaurus sp.
Theropoda indet. (including
Megalosaurus insignis)
Sauropoda
Eusauropoda
Lourinhasaurus alenquerensis
Brachiosauridae
Brachiosaurus atalaiensis
Sauropoda indet. (including
Pelorosaurus
humerocristatus)
Stegosauria
Stegosauridae
Dacentrurus armatus
Stegosauridae indet. (=Dacentrurus
armatus)
Age: Tithonian (Manuppela et al. 2000)
8. Arranhó unit (Lapparent and
Zbyszewski 1957)
Theropoda indet. (including
Megalosaurus pombali)
Age: Tithonian (Leinfelder 1993)
16.3.
DISTRITO DO COIMBRA
,
PORTUGAL
1. Unnamed unit (Gomes 1916; Lapparent
et al. 1951)
Theropod tracks
Age: middle-late Oxfordian (Manuppela
et al. 2000)
16.4.
DISTRITO DO LISBOA
,
PORTUGAL
1. Camadas de Alcobaça (Lapparent and
Zbyszewski 1957; Thulborn 1973a;
Galton 1980a, 1980b, 1994; Dantas et al.
1992; Bonaparte and Mateus 1999;
Mateus 1999; Pérez-Moreno et al. 1999;
Mateus and Antunes 2000)
Theropoda
Ceratosauria
Ceratosaurus sp.
Theropoda indet. (including
Megalosaurus insignis)
Sauropoda
Eusauropoda
Lourinhasaurus alenquerensis
Brachiosauridae incertae sedis
Brachiosaurus atalaiensis
Diplodocoidea incertae sedis
Dinheirosaurus lourinhanensis
Sauropoda indet.
Ornithischia indet. (including
Trimucrodon cuneatus)
Stegosauria
Stegosauridae
Dacentrurus armatus
Stegosauridae indet.
Ornithopoda
Euornithopoda
Hypsilophodon sp.
Euornithopoda indet.
Dinosaur eggs
548 DINOSAUR DISTRIBUTION AND BIOLOGY

Age: Kimmeridgian (Manuppela et al.
2000)
2. Unnamed unit (Mateus and Antunes
2001)
Ornithopod tracks
Age: Kimmeridgian (Mateus,
pers. comm.)
3. Unidade Sobral (Lapparent and
Zbyszewski 1957; Mateus 1998,
pers. comm.)
Theropoda
Carnosauria
Lourinhanosaurus antunesi
Theropoda indet.
Stegosauria
Stegosauridae
Dacentrurus armatus
Age: late Kimmeridgian–Tithonian
(Manuppela et al. 2000)
4. Unidade Amoreira-Porto Novo or
Castelhanos (Lapparent and
Zbyszewski 1957; Galton 1981b;
Mateus, pers. comm.)
Stegosauria
Stegosauridae
Dacentrurus armatus (including
Astrodon pusillus)
Age: late Kimmeridgian–Tithonian
(Manuppela et al. 2000)
5. Unidade Sobral or Amoreira Porto-Novo
(Lapparent and Zbyszewski 1957)
Sauropoda
Eusauropoda
Lourinhasaurus alenquerensis
Age: late Kimmeridgian–Tithonian
(Manuppela et al. 2000)
6. Unidade Amoreira-Porto Novo
(Lapparent and Zbyszewski 1957;
Mateus, pers. comm.)
Sauropoda
Eusauropoda
Lourinhasaurus alenquerensis
Brachiosauridae
Brachiosaurus atalaiensis
Stegosauria
Stegosauridae
Dacentrurus armatus
Age: late Kimmeridgian–Tithonian
(Manuppela et al. 2000)
7. Unnamed unit (Mateus et al. 1997;
Mateus, pers. comm.)
Theropoda
Carnosauria
Lourinhanosaurus antunesi
Theropod tracks
Sauropoda indet.
Dinosaur eggs
Age: late Kimmeridgian–Tithonian
(Mateus, pers. comm.)
8. Unnamed unit (Mateus, pers. comm.)
Sauropoda
Brachiosauridae
Brachiosaurus atalaiensis
Age: late Kimmeridgian–Tithonian
(Mateus, pers. comm.)
9. Unnamed unit (Lapparent and
Zbyszewski 1957; Rauhut and Kriwet
1994; Mateus, pers. comm.)
Theropoda indet. (including Megalosaurus
insignis, M. pombali)
Theropod tracks
Sauropoda
Eusauropoda
Lourinhasaurus alenquerensis
Age: late Kimmeridgian–Tithonian
(Mateus, pers. comm.)
10. Unidade Bombarral (Lapparent and
Zbyszewski 1957; Galton 1980a; Mateus
and Antunes 2001; Mateus, pers.
comm.)
Theropoda indet. (including
Megalosaurus insignis)
Theropod tracks
Sauropoda
Eusauropoda
Lourinhasaurus alenquerensis
Brachiosauridae
Brachiosaurus atalaiensis
Ornithopoda
Iguanodontia
Draconyx loureiroi
Age: Tithonian (Manuppela et al. 2000)
11. Unidade Arranhó (Mateus,
pers. comm.)
Sauropoda
Eusauropoda
Lourinhasaurus alenquerensis
Age: Tithonian (Leinfelder 1993)
12. Unnamed unit (Lapparent and
Zbyszewski 1957)
Sauropoda
Brachiosauridae
Brachiosaurus atalaiensis
Age: Tithonian (Mateus, pers. comm.)
16.5.
DISTRITO DO SETÚBAL
,
PORTUGAL
1. Unnamed unit (Lapparent and
Zbyszewski 1957)
Stegosauria
Stegosauridae
Dacentrurus armatus
Age: late Kimmeridgian–Tithonian
(Mateus, pers. comm.)
2. Azóia Limestone (Lockley et al. 1992b;
Santos et al. 1995)
Sauropod tracks
Age: Kimmeridgian (Lockley and Santos
1993)
3. Unnamed unit (Dantas et al. 1994)
Theropod tracks
Age: late Tithonian (=early Portlandian;
Lockley et al. 1994d)
17.1.
NIEDERSACHSEN
,
GERMANY
1. Unnamed unit (Windolf 1999, pers.
comm.)
Sauropoda
Undescribed brachiosaurid
Sauropoda indet. (including
Camarasauridae indet.,
Brachiosauridae indet.)
Age: Late Jurassic (Windolf 1999)
17.3.
BAYERN
,
GERMANY
1. Ober Solnhofen Plattenkalk
(Wagner 1861; Ostrom 1978a;
Griffiths 1993, 1999, 2000; Elz˙´anowski
2001)
Theropoda
Compsognathidae
Compsognathus longipes
Avialae
Archaeopterygidae
Archaeopteryx lithographica
Archaeopteryx bavarica
Wellnhoferia grandis
?Dinosaur eggs
Age: Kimmeridgian (Ostrom 1978a)
17.8.
NORDRHEIN
-
WESTPHALEN
,
GERMANY
1. Malm (Friese 1972, 1979; Kaever and
Lapparent 1974)
Theropod tracks
Sauropod tracks
Age: early Kimmeridgian (Kaever and
Lapparent 1974)
18.1.
KANTON BASELLAND
,
SWITZERLAND
1. Unnamed unit (Meyer, pers. comm.)
Theropoda indet.
Age: ?Oxfordian (Meyer, pers. comm.)
18.4.
KANTON BERN
,
SWITZERLAND
1. Unnamed unit (Meyer et al. 1997)
Theropoda indet.
Age: Oxfordian (Meyer et al. 1997)
2. Unnamed unit (Meyer et al. 1997)
Stegosauria indet.
Age: late Oxfordian (Meyer et al. 1997)
3. Unnamed unit (Huene 1925b; Meyer
et al. 1997)
Theropoda indet.
Age: early Kimmeridgian (Meyer et al.
1997)
4. Reuchenette Formation (Meyer 1990,
1993; Meyer and Hauser 1994; Meyer
and Lockley 1996; Meyer et al. 1997)
Theropod tracks
Sauropod tracks
Age: middle-late Kimmeridgian (Meyer
1993; Meyer and Lockley
1996)
18.5.
KANTON SOLOTHURN
,
SWITZERLAND
1. Villigen Formation (Meyer et al. 1997;
Meyer and Hunt 1998)
Stegosauria indet.
Age: late Oxfordian (Meyer et al. 1997)
2. Reuchenette Formation (Greppin 1870;
Huene 1922; Meyer 1990, 1993; Meyer
and Hauser 1994; Meyer and Lockley
1996; Meyer et al. 1997; Madsen and
Welles 2000)
Theropoda
Tetanurae indet. (=Megalosaurus
meriani)
Theropoda indet.
Theropod tracks
Sauropoda indet. (=Ornithopsis greppini)
Sauropod tracks
Age: middle-late Kimmeridgian (Meyer
1993; Meyer and Lockley
1996)
DINOSAUR DISTRIBUTION 549

Asia
Asian records of Late Jurassic dinosaurs (29 locations, an 80%
increase) have been scattered and often fragmentary (figs. 24.6,
24.7, and 24.16). Most of the material comes from the People’s
Republic of China, originally studied by Cai K.-J., P. J. Currie,
Dong Z.-M., He X.-L., C.-C. Young, Zhao Z.-C., and co-workers
are currently studying the still-burgeoning Late Jurassic dino-
saurs from Sichuan, many of which provide us with a new per-
spective on the fauna of this time interval. Records of dinosaur
ichnofaunas from Tadzhikistan, Turkmenistan, and Georgia have
been studied by L. K. Gabuniya, M. G. Lockley, V. P. Novikov,
and colleagues. Dinosaur faunas from Thailand have also been
described by E. Buffetaut, R. Ingavat, and V. Suteethorn. The first
Late Jurassic fauna from Mongolia (otherwise known only for
its Cretaceous dinosaur record) has recently been announced
by Y. M. Gubin and S. M. Sinitza. Finally, the dinosaur faunas of
M. Matsukawa, I. Obata, and Y. Tomida.
550 DINOSAUR DISTRIBUTION AND BIOLOGY
FIGU RE 24.16. Late Triassic to Late Cretaceous localities in Thailand (51)
and Laos (52). Scale =600 km.
35.1.
MUHAFAZAH SA
’
DAH
,
REPUBLIC
OF YEMEN
1. Mabdi Formation (Jacobs et al. 1999)
Sauropoda indet.
Age: Kimmeridgian (Jacobs et al. 1999)
37.2.
YAKUTSKAYA AVTONOMNAYA OBLAST
,
RUSSIA
1. Suntar Series (Rozhdestvensky 1973b;
Rich et al. 1997; Kurzanov et al. 2001)
Theropoda
Carnosauria indet.
?Coelurosauria indet.
Theropoda indet.
Sauropoda indet. (=Camarasauridae
indet.)
Stegosauria
Stegosauridae
cf. Stegosaurus sp.
Age: Late Jurassic–Early Cretaceous
(Rozhdestvensky 1973b)
38.2.
TYVA RESPUBLIKA
,
RUSSIA
1. Unnamed unit (Kudryavtseva and
Kudryavtsev 2003) (not plotted)
Unidentifed Dinosauria
Age: Late Jurassic (Kudryavtseva and
Kudryavtsev 2003)
38.1.
AP
’
KHAZET
’
IS AVTONOMIURI
RESPUBLIKA
,
GEORGIA
1. Unnamed unit (Gabunia et al. 1998)
(not plotted)
Sauropoda indet.
Age: middle Oxfordian (Gabuniya et al.
1998)
39.1.
ONGTUSTIK QAZAQSTAN OBLYSY
,
KAZAKHSTAN
1. Balabansai Svita (Rautian 1978;
Kurochkin 2000)
Theropoda
Avialae incertae sedis
Praeornis sharovi
Age: Late Jurassic (Kurochkin 2000)
40.1.
QASHQADARYO WILOYATI
,
UZBEKISTAN
1. Kurek Svita (Gabuniya and Kurbatov
1982, 1988; Lockley et al. 1996d, 1998a)
Theropod tracks
Sauropod tracks
Age: late Oxfordian–early Kimmeridgian
(Lockley et al. 1998a)
40.2.
SURKHONDARYO WILOYATI
,
UZBEKISTAN
1. Unnamed unit (Lockley, pers. comm.)
Theropod tracks
Age: Oxfordian-Kimmeridgian (Lockley,
pers. comm.)
41.1.
CHARDZHOU OBLAST
,
TURKMENISTAN
1. Unnamed unit (Lockley et al. 1996d)
Theropod tracks
Age: Oxfordian-Kimmeridgian
(Lockley et al. 1996d,
1998a, 1998b)
43.1.
VILOYATI LENINOBOD
,
TADZHIKISTAN
1. Kugitang Svita (Dzhalilov and Novikov
1987; Lockley et al. 1994e)
?Theropod tracks
Age: Oxfordian (Lockley et al. 1994e)
2. Karabil Svita (Novikov and Dzhalilov
1993)
Theropod tracks
Age: Tithonian (Lockley, pers. comm.)
46.4.
RAJASTHAN
,
INDIA
1. ?Jabalpur Formation (Mathur et al. 1985)
Dinosauria indet.
Age: Tithonian-Hauterivian (Dogra et al.
1994)
47.1.
GOV
’
ALTAI
,
MONGOLIA
1. Shar Teeg Beds (Gubin and Sinitza 1996)
Theropoda indet.
Age: Late Jurassic (Gubin and Sinitza
1996)
2. Ulaan Malgait Beds (Gubin and Sinitza
1996)
Sauropoda indet. (=?Mamenchisaurus sp.)
Age: Late Jurassic (Gubin and Sinitza
1996)
3. Daril Formation (Graham et al. 1997)
Sauropoda indet. (=Mamenchisaurus sp.)
Ornithischia indet.
Age: Late Jurassic (Graham et al. 1997)
48.2.
SICHUAN
,
PEOPLE
’
S REPUBLIC
OF CHINA
1. Kyangyan Series (Young 1942b)
Theropoda indet. (=?Szechuanosaurus
campi, Sinocoelurus fragilis)
Age: Tithonian (Dong, pers. comm.)
2. Upper Shaximiao Formation (Camp
1935; Young 1939; Dong et al. 1983; He
et al. 1984; Gao 1992; Currie and Zhao
1993a; Pi et al. 1996; Zhang and Chen
1996; Zhang et al. 1998; Ye et al. 2001;
Zhang and Li 2001)
Theropoda
Allosauroidea
Yangchuanosaurus shangyouensis
Yangchuanosaurus magnus
Sinraptor hepingensis
Theropoda indet. (including
Szechuanosaurus campi,
S. yandonensis)
Sauropoda
Eusauropoda
Mamenchisaurus constructus
Mamenchisaurus hochuanensis
Mamenchisaurus jingyanensis
Mamenchisaurus youngi
Sauropoda indet. (including
Omeisaurus changshouensis,
O. fuxiensis)
Ornithischia indet. (=Gongbusaurus
shiyii)

Stegosauria
Stegosauridae
Tuojiangosaurus multispinus
Chialingosaurus kuani
Chungkingosaurus jiangbeiensis
Chungkingosaurus sp.
Stegosauridae indet.
Age: Late Jurassic (Dong et al. 1983);
Oxfordian (Dong, pers.
comm.)
3. Suining Formation (He et al. 1996)
Theropoda
Yangchuansaurus sp.
Sauropoda
Eusauropoda
Mamenchisaurus anyuensis
Age: Late Jurassic (He et al. 1996)
4. Penglaizhen Formation (Zhen et al.
1983; He et al. 1996)
Sauropoda
Eusauropoda
Mamenchisaurus anyuensis
Ornithischian tracks
Age: Late Jurassic (Zhen et al. 1983)
5. Lower Chiating Series (Young 1960;
Zhen et al. 1989)
Euornithopod tracks
Age: Late Jurassic (Young 1960)
48.4.
LIAONING
,
PEOPLE
’
S REPUBLIC
OF CHINA
1. Tuchengzi Formation (Wang et al. 1985;
Zhao et al. 1999; Dong 2001; Lockley
et al. 2001b)
Theropod tracks (including avialan
tracks)
Sauropoda indet. (=Mamenchisauridae
indet., Brachiosauridae
indet.)
Ceratopsia
Neoceratopsia
Chaoyangosaurus youngi
Age: Tithonian (Wang et al. 1985;
Lockley et al. 2001b)
48.5.
XINJIANG UYGUR ZIZHIQU
,
PEOPLE
’
S REPUBLIC OF CHINA
1. Shishugou Formation (Young 1937;
Dong 1989; Currie and Zhao 1993a;
Russell and Zheng 1993; Clark, pers.
comm.)
Theropoda
Allosauroidea
Sinraptor dongi
Undescribed ornithomimosaurian
Tetanurae indet.
Sauropoda
Eusauropoda
Mamenchisaurus sinocanadorum
Sauropoda indet. (including
Tienshanosaurus chitaiensis)
Undescribed stegosaur
Undescribed ankylosaur
Ornithopoda
Unnamed euornithopodan
(=“Gongbusaurus”
wucaiwanensis)
Age: Bathonian-Oxfordian (Eberth et al.
2001a)
2. Kalaza Formation (Young and Sun 1957;
Dong 1977, 1997d; Sun et al. 1992; Yao et
al., in press; =Karaza Formation, Keilozo
Formation, Huoyanshan Formation,
Kalazha Formation)
Theropoda indet. (=cf. Szechuanosaurus
campi)
Sauropoda
Eusauropoda incertae sedis
Hudiesaurus sinojapanorum
Sauropoda indet. (including
Chiayusaurus lacustris)
Age: Tithonian (Eberth et al. 2001a)
3. Unnamed unit (Dong 1973b)
Theropoda indet.
Age: ?Late Jurassic (Dong 1973b)
4. Qigu Formation (Maisch et al. 2003; Yao
et al., in press)
Theropoda
Tetanurae indet.
Coelurosauria indet.
Sauropoda indet.
Age: Oxfordian (Eberth et al. 2001a)
48.6.
SHAANXI
,
PEOPLE
’
S REPUBLIC
OF CHINA
1. Unnamed unit (Kuhn 1958b; Lockley
and Matsukawa 1998)
Euornithopod tracks
Age: Late Jurassic (Kuhn 1958; ?Early
Cretaceous, Lockley and
Matsukawa 1998)
48.8.
GANSU
,
PEOPLE
’
S REPUBLIC
OF CHINA
1. Hantong Formation (Young 1958b;
Dong, pers. comm.)
Sauropoda
Eusauropoda
Mamenchisaurus constructus
Age: Late Jurassic (Young 1958b)
48.9.
SHANDONG
,
PEOPLE
’
S REPUBLIC
OF CHINA
1. Meng-Yin Formation (Wiman 1929;
Mateer and McIntosh 1985)
Sauropoda
Eusauropoda
Euhelopus zdanskyi
Stegosauria
Stegosauridae indet.
Age: early Tithonian (Chen et al. 1982);
Kimmeridgian (Dong, pers.
comm.)
2. Laiyang Series (Young 1960; Zhen et al.
1989)
Theropod tracks
Age: Late Jurassic (Zhen et al. 1989)
48.10.
HEBEI
,
PEOPLE
’
S REPUBLIC
OF CHINA
1. Houcheng Formation (Wang et al. 1985;
Zhao unpubl. data)
Ceratopsia
Xuanhuasaurus nieii
Age: Late Jurassic (Wang et al. 1985;
Zhao unpubl. data)
51.2.
CHANGWAT KALASIN
,
THAILAND
1. Phu Kradung Formation (Buffetaut et al.
1997b, 2001, 2002; Buffetaut and
Suteethorn 1998b)
Theropoda indet.
Sauropoda indet.
Stegosauria indet.
Ornithopoda indet.
Age: Late Jurassic (Buffetaut et al.
1997b)
51.3.
CHANGWAT MUKDAHAN
,
THAILAND
1. Phu Kradung Formation (Buffetaut et al.
1997b; Buffetaut and Suteethorn 1998b)
Theropoda indet.
Sauropoda indet.
Stegosauria indet.
Ornithopoda indet.
Age: Late Jurassic (Buffetaut et al. 1
997b)
53.2.
GIFU PREFECTURE
,
JAPAN
1. Kagidani Formation (Kunimitsu et al.
1990; Matsukawa and Obata 1994;
Azuma and Tomida 1995)
?Theropod tracks
?Euornithopod tracks
Age: Tithonian (Azuma and Tomida
1995)
53.3.
FUKUSHIMA PREFECTURE
,
JAPAN
1. Tochikubo Formation (Takahashi and
Taira 1996, 1997)
Theropod tracks
Euornithopod tracks
Age: Late Jurassic (Takahashi and Taira
1996, 1997)
DINOSAUR DISTRIBUTION 551
South America
The Late Jurassic dinosaur faunas of South America are known
from seven locations (a 75% increase; figs. 24.8 and 24.9). Four
are represented by dinosaur tracks or trackways from Brazil, Chile,
and Argentina, described by G. Leonardi, R. M. Casamiquela,
O. Galli, R. Dingman, and D. Rubilar, among others. Another,
from Colombia, is represented by possible cetiosaurid material
described by W. Langston Jr. and J. W. Durham, while G. C. Diaz
and Z. B. de Gasparini report an undescribed dinosaur fauna from
northern Chile.

54.1.
DEPARTAMENTO DE MAGDALENA
,
COLOMBIA
1. “Giron” formation (Langston and
Durham 1955)
Sauropoda indet. (=?Cetiosauridae
indet.)
Age: ?Late Jurassic (Bonapar te 1979a)
57.4.
ESTADO DO MINAS GERAIS
,
BRAZIL
1. Areado Group (Carvalho and Kattah
1996)
Theropod tracks
Age: Late Jurassic–Early Cretaceous
(Carvalho and Kattah 1996)
60.1.
UPPER TAKUTU
,
GUYANA
1. Takutu Formation (Montalvão et al.
1975)
Theropod tracks
Age: Late Jurassic–Early Cretaceous
(Hammen and Burger 1966)
62.1.
PROVINCIA DE ANTOFAGASTA
,
CHILE
1. Unnamed unit (Diaz and Gasparini
1976)
Undescribed dinosaur
Age: Late Jurassic (Diaz and Gasparini
1976)
62.2.
PROVINCIA DE TARAPACA
,
CHILE
1. Chacarilla Formation (Galli and
Dingman 1962; Moreno et al. 2000)
Theropod tracks
Sauropod tracks
?Stegosaur tracks
Euornithopod tracks
Age: ?Oxfordian (Diaz and Gasparini
1976)
62.3.
PROVINCIA DE COLCHAGUA
,
CHILE
1. Baños del Flaco Formation (Rubilar,
pers. comm.)
Dinosaur tracks
Age: Late Jurassic (Rubilar, pers.
comm.)
63.3.
PROVINCIA DE SANTA CRUZ
,
ARGENTINA
1. La Matilde Formation (Casamiquela
1964b; Coria 1991)
Theropod tracks
Age: Oxfordian (Stipanicic and Bonetti
1970a, 1970b)
552 DINOSAUR DISTRIBUTION AND BIOLOGY
Africa
The Tendaguru fauna of Tanzania is perhaps the best known of
all dinosaur faunas from Africa. This site was originally worked by
E. Fraas from the Königlichen Naturalienkabinett Stuttgart (now
the Staatliches Museum für Naturkunde, Stuttgart), but most
extensively between 1909 and 1912 by W. Janensch, E. Hennig,
and H. Reck under the auspices of the Humboldt-Universität
Museum für Naturkunde (for further information on the Ger-
man expedition to Tendaguru, see Hennig 1912). From 1924 to
1929, Tendaguru was worked by an expedition from the British
Museum (Natural History), headed successively by W. E. Cutler,
F. W. H. Migeod, and J. Parkinson (Parkinson 1930). Although
the Tendaguru fauna is not presently worked (Zils et al. 1995),
revisionary studies of the fauna continue to recognize new taxa,
principally through the work of O. W. M. Rauhut. Other African
work on the Late Jurassic has been conducted by D.-M. Dutuit
and A. Ouazzou in Morocco, L. Ginsburg and co-workers in
Niger, and M. A. Raath in Zimbabwe. Recently, M. B. Goodwin
and collaborators have discovered the first Ethiopian dinosaur
fauna, also from the Late Jurassic. Upper Jurassic dinosaur loca-
tions in Africa are given in figures 24.10 and 24.11.
64.1.
PROVINCE DE MARRAKECH
,
MOROCCO
1. Unnamed unit (Dutuit and Ouazzou
1980)
Sauropod tracks
Age: Late Jurassic or Early Cretaceous
(Dutuit and Ouazzou
1980)
68.1.
DÉPARTEMENT D
’
AGADEZ
,
NIGER
1. Grès d’Assaouas (Ginsburg et al. 1966)
Theropod tracks
Sauropod tracks
Age: Late Jurassic–Hauterivian
(Ginsburg et al. 1966;
Taquet 1976)
2. Argiles de l’Irhazer (Ginsburg et al. 1966;
Congleton 1990)
Theropod tracks
Euornithopod tracks
Age: Late Jurassic–Hauterivian
(Ginsburg et al. 1966;
Taquet 1976)
72.1.
SHEWA PROVINCE
,
ETHIOPIA
1. Mugher Mudstone (Goodwin et al. 1996,
1999)
Theropoda
Carnosauria
cf. Acrocanthosaurus sp.
Theropoda indet.
Ornithischia indet.
Ornithopoda
Euornithopoda indet.
Age: Tithonian–?Early Cretaceous
(Goodwin et al. 1996,
1999)
75.1.
MKOAWA MTWARA
,
TANZANIA
1. Tendaguru Formation (Janensch 1914,
1929b, 1935–36, 1955; Hennig 1924;
Swinton 1950; Rauhut 1998; Bonaparte
et al. 2000; Madsen and Welles 2000)
Theropoda
Ceratosauria
Elaphrosaurus bambergi
Ceratosaurus sp. (=Ceratosaurus
roechlingi, Labrosaurus
stechowi)
Allosauroidea
?Allosaurus tendagurensis
Theropoda indet. (=Megalosaurus
ingens)
Sauropoda
Sauropoda incertae sedis
Tendaguria tanzaniensis
Diplodocoidea
Dicraeosaurus hansemanni
Dicraeosaurus sattleri
Tornieria africana
Barosaurus gracilis
Brachiosauridae
Brachiosaurus brancai (including
B. fraasi)
Titanosauria
Janenschia robusta
Stegosauria
Stegosauridae
Kentrosaurus aethiopicus
Ornithopoda
Iguanodontia
Dryosaurus lettowvorbecki
?Dinosaur eggs
Age: Kimmeridgian (Aitken 1961;
Schudack 1999)
77.1.
MASHONALAND NORTH
,
ZIMBABWE
1. Kadsi Formation (Armstrong et al.
1967; Raath 1967; Raath and McIntosh
1987 )
Sauropoda
Brachiosauridae
Brachiosaurus sp.
?Brachiosauridae indet.
Diplodocidae
Barosaurus sp.
Dicraeosauridae
Dicraeosaurus sp.
Camarasauromorpha
?Camarasaurus sp.
Titanosauria
Janenschia sp.
Age: Late Jurassic (Molnar 1980b; Raath
and McIntosh 1987)
77.5.
MIDLANDS
,
ZIMBABWE
1. Gokwe Formation (Bond and Bromley
1970; Raath and McIntosh 1987)
Theropoda indet.
Sauropoda indet.
Age: Late Jurassic (Raath and McIntosh
1987 )

Early Cretaceous
Dinosaur fossils of Early Cretaceous age are known from nearly
280 locations in North America, Europe, Asia, South America,
Africa, and Australasia.
North America
Lower Cretaceous dinosaurs are presently known from 48 loca-
tions (an increase of nearly 50%). The most important among
them are found primarily in the Western Interior of the United
States, thanks to the early work of B. Brown and P. Kaisen and
later work by P. R. Bjork, B. B. Britt, K. Carpenter, J. A. Dorr Jr.,
J. G. Eaton, A. P. Hunt, J. I. Kirkland, M. G. Lockley, S. G. Lucas,
W. D. Maxwell, J. H. Ostrom, R. P. Ratkevich, and D. B. Wei-
shampel. The first Early Cretaceous dinosaur occurrence from
California was reported by R. P. Hilton and colleagues. Addi-
tional western North American records include important ichno-
faunas from British Columbia and Alberta, Canada, and Alaska,
United States, described by P. J. Currie, R. A. Gangloff, and C. M.
DINOSAUR DISTRIBUTION 553
Sternberg. To the southeast, important Comanchean sites from
Oklahoma, central Texas, and Arkansas have yielded both track
and skeletal material, described in detail by D. D. Gillette, L. L.
Jacobs, W. Langston Jr., J. G. Pittman, D. A. Thomas, and D. A.
Winkler. Finally, an intriguing but not yet well-known fauna
comes from the Arundel Clay of Maryland, Washington, D.C.,
and Virginia, studied by C. W. Gilmore, T. R. Lipka, R. S. Lull,
O. C. Marsh, R. Stanford, and R. E. Weems. These Lower Creta-
ceous locations are plotted in figures 24.1, 24.13, 24.17, and
24.18. (Listed here is an ichnofauna from Estado de Puebla,
Mexico, described by Leonardi (1994), which can be dated only
as ?Cretaceous in age [location 44.1-2].)
1.3.
BRITISH COLUMBIA
,
CANADA
1. Minnes Formation (Currie 1989a)
Theropod tracks
Age: Berriasian-Valanginian (Currie
1989a)
2. Cadomin Formation (Currie 1989a)
Theropod tracks
Euornithopod tracks
Age: Barremian-Aptian (Currie
1989a)
3. Gething Formation (Sternberg 1932b;
Currie and Sarjeant 1979; Carpenter 1984)
Theropod tracks (including avialan
tracks)
Ankylosaur tracks
Euornithopod tracks
Age: Barremian-Aptian (Stott 1975)
1.4.
ALBERTA
,
CANADA
1. Gates Formation (McCrea and Currie
1998; McCrea and Sarjeant 2001)
Theropod tracks (including avialan
tracks)
Ankylosaur tracks
Euornithopod tracks
Age: Albian (McCrea and Currie 1998)
3.1.
UTAH
,
UNITED STATES
1. Kelvin Formation (Jensen 1970;
Zelenitsky et al. 2000)
Dinosaur eggs
Age: Berriasian-Albian (Hintze 1973)
FIGU RE 24.17. Early and Late Cretaceous localities in the southwestern United States (3). Scale =600 km.

2. Lower Cedar Mountain Formation
(Galton and Jensen 1979b; Kirkland
et al. 1993, 1997, 1998a, 1998b; Britt
and Stadtman 1997; Kirkland 1998a;
Carpenter 1999; Lockley et al. 1999;
Tidwell et al. 1999, 2001; Carpenter et al.
2001a; DiCroce and Carpenter 2001;
Scheetz et al. 2001)
Theropoda
Coelurosauria incertae sedis
Nedcolbertia justinhofmanni
Dromaeosauridae
Utahraptor ostrommaysi
Theropoda incertae sedis
cf. Ricardoestesia sp.
Theropod tracks
Sauropoda
Brachiosauridae
Cedarosaurus weiskopfae
?Brachiosauridae indet.
Undescribed ?macronarian
(=“camarasaurid”)
Titanosauria
Venenosaurus dicrocei
Sauropod tracks
Ankylosauria
Nodosauridae
Cedarpelta bilbeyhallorum
Ankylosauridae
Gastonia burgei
Ornithopoda
Iguanodontia
Planicoxa venenica
Undescribed iguanodontians
Iguanodontia indet. (=Iguanodon
ottingeri)
Euornithopod tracks
Dinosaur eggs
Age: Berriasian-Hauterivian (Kirkland
et al. 1993, 1997, 1998a)
3. Middle Cedar Mountain Formation
(Bodily 1969; DeCourten 1991; Kirkland
et al. 1997, 1998a; Carpenter and
Kirkland 1998; Harris 1998c; Carpenter
1999; Lockley et al. 1999; Zelenitsky
et al. 2000; Scheetz et al. 2001)
Theropoda
Carnosauria
Acrocanthosaurus sp.
Undescribed allosaurid
Undescribed coelurosaurian
Dromaeosauridae
Deinonychus sp.
Theropoda incertae sedis
cf. Ricardoestesia sp.
Sauropoda
Titanosauriformes incertae sedis
Pleurocoelus sp.
Ankylosauria
Nodosauridae
Unnamed Sauropelta species
cf. Sauropelta sp. (=Hoplitosaurus sp.)
Undescribed ankylosaurid
(=cf. Shamosaurus sp.)
Ornithopoda
Iguanodontia
Tenontosaurus sp.
Euornithopod tracks
Dinosaur eggs
Age: Aptian-Albian (Kirkland et al. 1998a)
3.2.
ARIZONA
,
UNITED STATES
1. Dakota Formation (unpublished)
Euornithopod tracks
Age: late Aptian–early Cenomanian
(Tschudy et al. 1984b)
2. Shellenberger Canyon Formation (Miller
1964; Galton and Jensen 1979b;
Ratkevich 1997)
Ornithopoda
Iguanodontia
cf. Tenontosaurus sp.
Age: Aptian-Albian (Galton and Jensen
1979b)
554 DINOSAUR DISTRIBUTION AND BIOLOGY
FIGU RE 24.18. Early and Late Cretaceous localities in the central and southeastern United States (3). Scale =600 km.

3. Sand Wells Formation (Miller 1964;
Ratkevich 1997)
?Stegosauria
?Stegosauridae indet.
Age: Early Cretaceous (Miller 1964)
4. Amole Formation (Ratkevich 1997)
Ornithopoda
Iguanodontia indet.
Age: Aptian-Albian (Ratkevich 1997)
5. Unnamed unit (Ratkevich 1997)
Theropoda
Dromaeosauridae
cf. Deinonychus sp.
Ornithopoda
Iguanodontia
cf. Tenontosaurus sp.
Age: ?Early Cretaceous (Ratkevich 1997)
6. Turney Ranch Formation (Ratkevich
1997, 1998)
Sauropoda indet. (including
Sonorasaurus thompsoni)
Age: Albian-Cenomanian (Ratkevich
1997)
3.3.
NEW MEXICO
,
UNITED STATES
1. Mesa Rica Formation (Hunt and Lucas
1998; Lockley et al. 2000; Lucas et al.
2000)
Theropod tracks
Age: late Albian (Lucas et al. 1989;
Lockley et al. 1993)
2. Pajarito Formation (Lucas et al. 1989;
Lockley et al. 1993, 2000; Hunt and
Lucas 1998)
Theropod tracks
Euornithopod tracks
Age: late Albian (Lucas et al. 1989;
Lockley et al. 1993)
3.4.
COLORADO
,
UNITED STATES
1. South Platte Formation (Johnson 1931;
Mehl 1931; Lockley 1985; Currie et al.
1991; Lockley et al. 1992a, 2001a;
McCrea et al. 2001a)
Theropod tracks (including avialan
tracks)
?Ankylosaur tracks
Euornithopod tracks
Age: late Aptian–early Cenomanian
(Lucas et al. 1998b)
2. Plainview Formation (Kurtz et al. 2001;
McCrea et al. 2001a)
Theropod tracks
Ankylosaur tracks
Euornithopod tracks
Age: Albian (Kurtz et al. 2001)
3.5.
TEXAS
,
UNITED STATES
1. Bissett Formation (Langston, pers.
comm.)
Ornithopoda
Iguanodontia
cf. Iguanodon sp.
Age: ?Berriasian–early Albian (Langston,
pers. comm.)
2. Twin Mountain Formation (Winkler and
Murry 1989; Winkler et al. 1997b; Harris
1998c; Jacobs and Winkler 1998)
Theropoda
Carnosauria
Acrocanthosaurus atokensis
Dromaeosauridae indet.
Theropoda indet.
Sauropoda
Sauropoda indet.
Titanosauriformes incertae sedis
Pleurocoelus sp.
Ornithopoda
Unnamed euornithopodan
Iguanodontia
Tenontosaurus dossi
Iguanodontia indet.
Age: Aptian (Winkler and Murry 1989;
Jacobs et al. 1991)
3. Bluff Dale Sandstone (Langston,
pers. comm.)
Sauropoda
Titanosauriformes incertae sedis
Pleurocoelus sp.
Age: late Aptian (Langston 1974,
pers. comm.)
4. Glen Rose Formation (Langston 1974;
Farlow 1987a; Pittman and Lockley
1994; Rogers 2002; Hawthorne et al.
2002; Jacobs and Winkler 1998)
Theropod tracks
Sauropoda
Titanosauriformes
Pleurocoelus cf. nanus
Unnamed titanosauriform
Sauropod tracks
Ornithopoda
Iguanodontia
Tenontosaurus cf. tilletti
Euornithopod tracks
Age: late Aptian–early Albian (Langston
1974)
5. Antlers Formation (Langston 1974;
Winkler and Murry 1989; Jacobs and
Winkler 1998)
Theropoda
Carnosauria
Acrocanthosaurus atokensis
Dromaeosauridae indet.
Theropoda indet.
Sauropoda indet.
Ornithopoda
Euornithopoda
Undescribed euornithopodan
Iguanodontia
Tenontosaurus sp.
Age: late Aptian–middle Albian
(Langston 1974)
6. Paluxy Formation (Langston 1974)
Theropoda
Ornithomimosauria
?Ornithomimus sp.
?Dromaeosauridae indet.
Theropoda indet.
Sauropoda
Titanosauriformes
Pleurocoelus cf. nanus
?Pleurocoelus sp.
Ornithischia indet.
Ankylosauria incertae sedis
?Nodosaurus sp.
Ornithopoda
Iguanodontia
Tenontosaurus cf. tilletti
Age: middle Albian (Young 1986)
7. Edwards Formation (Stanford, pers.
comm.)
Sauropod tracks
Age: middle Albian (Stephenson et al.
1942)
8. Paw Paw Formation (Jacobs et al. 1994;
Coombs 1995b; Lee 1996)
Ankylosauria
Nodosauridae
Pawpawsaurus campbelli
Nodosauridae indet.
Ankylosauria incertae sedis
Texasetes pleurohalio
Age: late Albian (Jacobs et al. 1991)
9. Segovia Formation (Pittman 1989)
Theropod tracks
Age: late Albian–early Cenomanian
(Pittman 1989)
10. Fort Terrett Formation (Pittman 1989)
Theropod tracks
Age: late Albian (Pittman 1989)
11. Hensel Formation (Pittman 1989)
Euornithopod tracks
Age: middle Albian (Pittman 1989)
3.6.
OKLAHOMA
,
UNITED STATES
1. Antlers Formation (Larkin 1910;
Langston 1974; Cifelli et al. 1997a;
Brinkman et al. 1998; Currie and
Carpenter 2000; Wedel et al. 2000)
Theropoda
Carnosauria
Acrocanthosaurus atokensis
Dromaeosauridae
Deinonychus antirrhopus
cf. Deinonychus sp.
Avialae indet.
Sauropoda
Brachiosauridae
Sauroposeidon proteles
Titanosauriformes incertae sedis
Pleurocoelus sp. (=?Astrodon sp.)
Sauropoda indet.
Sauropod tracks
Ornithopoda
Iguanodontia
Tenontosaurus sp.
Age: late Aptian–middle Albian
(Langston 1974)
3.7.
KANSAS
,
UNITED STATES
1. Dakota Sandstone (Eaton 1960;
McAllister 1989)
Ankylosauria
Nodosauridae
Silvisaurus condrayi
?Euornithopod tracks
Age: late Aptian–early Cenomanian
(Tschudy et al. 1984b)
3.10.
CALIFORNIA
,
UNITED STATES
1. Budden Canyon Formation (Hilton et al.
1997)
Ornithopoda
DINOSAUR DISTRIBUTION 555

Euornithopoda indet.
Age: late Aptian (Hilton et al. 1997)
3.11.
MONTANA
,
UNITED STATES
1. Cloverly Formation (Ostrom 1970a;
Maxwell and Horner 1994; Makovicky
and Sues 1998; Makovicky and Grellet-
Tinner 1999)
Theropoda
Ornithomimidae indet.
(=?Ornithomimus velox)
Oviraptorosauria
Microvenator celer
Dromaeosauridae
Deinonychus antirrhopus
Theropoda indet.
Sauropoda
Titanosauriformes indet.
(=?Pleurocoelus sp.)
Ankylosauria
Nodosauridae
Sauropelta edwardsorum
Ornithopoda
Euornithopoda
Zephyrosaurus schaffi
Iguanodontia
Tenontosaurus tilletti
Dinosaur eggs
Age: late Aptian (Chen and Lubin 1997)
2. Kootenai Formation (Horner 1989)
Dinosauria indet.
Age: Aptian (Cobban and Reeside 1952)
3.12.
WYOMING
,
UNITED STATES
1. Mowry or Thermopolis Shale (Marsh
1889a; Williston 1905; Lull 1921;
Coombs 1978a)
Ankylosauria incertae sedis
Nodosaurus textilis
Age: late Aptian (Vuke 1984)
2. Cloverly Formation (Ostrom 1970a)
Theropoda
Ornithomimidae indet.
(=?Ornithomimus velox)
Oviraptorosauria
Microvenator celer
Dromaeosauridae
Deinonychus antirrhopus
Theropoda indet.
Sauropoda
Sauropoda indet.
Titanosauriformes incertae sedis
Pleurocoelus sp.
Ankylosauria
Nodosauridae
Sauropelta edwardsorum
Ornithopoda
Iguanodontia
Tenontosaurus tilletti
Age: late Aptian (Chen and Lubin 1997)
3. Thomas Fork Formation (Dorr 1985)
Dinosauria indet.
Dinosaur eggs
Age: Albian (Dorr 1985)
3.13.
IDAHO
,
UNITED STATES
1. Wayan Formation (Dorr 1985;
Weishampel et al. 2002)
Ankylosauria
?Nodosauridae indet.
Ornithopoda
Ornithopoda indet.
Iguanodontia
Tenontosaurus sp.
Neoceratopsia indet.
Dinosaur eggs
Age: Albian (Dorr 1985)
3.15.
SOUTH DAKOTA
,
UNITED STATES
1. Lakota Formation (Lucas 1901;
Anderson 1939; Galton and Jensen
1979b; Bjork 1985a; Weishampel and
Bjork 1989; Carpenter and Kirkland
1998; Foster et al. 1998; Lockley et al.
2001a)
Theropod tracks (including avialan
tracks)
Sauropoda indet. (=Camarasauridae
indet.)
Ankylosauria incertae sedis
Hoplitosaurus marshi
Ornithopoda
Iguanodontia
Iguanodon lakotaensis
Camptosaurus depressus
Euornithopod tracks
Ornithopoda indet. (=Hypsilophodon
wielandi)
Age: Barremian (Sohn 1979)
3.16.
NEBRASKA
,
UNITED STATES
1. Dakota Sandstone (Barbour 1931; Galton
and Jensen 1979b; Joekel et al. 2001)
Ornithopoda
?Hadrosauridae indet.
Ornithopod tracks
Age: late Aptian–early Cenomanian
(Tschudy et al. 1984b)
3.18.
ARKANSAS
,
UNITED STATES
1. DeQueen Formation (Gillette et al. 1985;
Pittman and Gillette 1989)
Sauropod tracks
Age: Albian (Gillette et al. 1985)
2. Trinity Formation (Quinn 1973)
Theropoda
Ornithomimidae indet.
Age: Early Cretaceous (Quinn 1973)
3.25.
MARYLAND
,
UNITED STATES
1. Arundel Clay (Lull 1911a, 1911b; Gilmore
1921; Ostrom 1970a; Galton and Jensen
1979b; Weishampel and Young 1996;
Chinnery et al. 1998; Lipka 1998)
Theropoda
Allosauroidea indet.
Ornithomimosauria indet.
(=Coelosaurus affinis)
Dromaeosauridae
Deinonychus sp.
Theropoda indet. (=Allosaurus medius,
Coelurus grandis, Creosaurus
potens)
Sauropoda
Titanosauriformes
Pleurocoelus altus
Pleurocoelus nanus
Sauropoda indet. (=Astrodon johnstoni)
Ankylosauria
Nodosauridae
Priconodon crassus
Ornithopoda
Iguanodontia
?Tenontosaurus sp.
Ceratopsia
Neoceratopsia indet.
Age: middle-late Aptian (Doyle 1992;
Lipka, pers. comm.)
2. Patuxent Formation (Stanford 1998;
Stanford and Stanford 1998)
Theropod tracks
Sauropod tracks
Euornithopod tracks
Thyreophoran tracks
Age: early Aptian (Doyle 1992; Lipka,
pers. comm.)
3.26.
VIRGINIA
,
UNITED STATES
1. Patuxent Formation (Weems and
Bachman 1997)
Theropod tracks
Euornithopod tracks
Age: Aptian (Weems and Bachman
1997)
3.33.
ALASKA
,
UNITED STATES
1. Chandler Formation (Parrish et al.
1987; Gangloff 1998; McCrea et al.
2001a)
Theropoda
Tyrannosauridae indet.
Theropod tracks
Ankylosaur tracks
Euornithopod tracks
Age: Albian-Cenomanian (Parrish et al.
1987 )
3.34.
WASHINGTON
,
D
.
C
.,
UNITED STATES
1. Arundel Clay (Gilmore 1920, 1921;
Weishampel and Young 1996)
Theropoda indet.
Sauropoda
Titanosauriformes incertae sedis
Pleurocoelus sp. (=Astrodon johnsoni)
Age: late Aptian–early Albian (Robbins
1991 )
556 DINOSAUR DISTRIBUTION AND BIOLOGY
Europe
The Early Cretaceous is well known in Europe (89 locations,
amounting to nearly a 90% increase). The Wealden and Wealden
equivalents have produced the greatest number of dinosaur
faunas (figs. 24.1, 24.5, and 24.19). Important locations include

those of England, Belgium, Germany, France, Spain, and Portu-
gal. The English dinosaur faunas are known through the early
work of W. Buckland, J. W. Hulke, G. A. Mantell, R. Owen, H. G.
Seeley, and recently through the research of P. M. Barrett, W. T.
Blows, A. J. Charig, P. C. Ensom, S. Hutt, D. M. Martill, D. B. Nor-
man, and J. L. Wright. Much more spectacular though more
limited in scope, the Early Cretaceous dinosaurs of Belgium,
inextricably linking the Bernissart iguanodons and the name of
L. Dollo, remain one of the best investigated of all dinosaurian
occurrences (viz. Casier 1978; Norman 1987c). Germany has pro-
vided additional perspectives on Early Cretaceous mass assem-
blages and other faunal aspects through the work of H. Holder,
E. Koken, D. B. Norman, and K. Oekentorp. Elsewhere in Europe,
dinosaurian faunas appear to have a more differentiated com-
position. In France, these have been studied principally by H.-E.
Sauvage, A. F. de Lapparent, V. Stchepinsky, E. Buffetaut, J. Le
Loeuff, and V. Martin, and in Spain and Portugal by, among
many others, L. M. Aguirrezabala, A. Buscalioni, J. I. Canudo,
M. L. Casanovas, C. A. Meyer, J. J. Moratalla, B. Perez-Moreno,
N. H. Platt, J. I. Ruiz-Omeñaca, B. Sanchiz, J. V. Santafé, J. L.
Sanz, X. Pereda-Suberbiola, and L. I. Viera. Finally, three dino-
saur occurrences, less well known, deserve mention. One, from
western Romania, was originally worked by T. Jurcsák and E. Popa,
and later by M. J. Benton and co-workers. Another, in Croatia, is
known through the work of F. M. Dalla Vecchia. Finally, C. Dal
Sasso and M. Signore (1998) announced a new theropod skele-
ton from Italy that also has exceptionally well-preserved soft
tissue.
DINOSAUR DISTRIBUTION 557
FIGU RE 24.19. Early and Late Cretaceous localities in Sweden (6), England (10), Belgium (11), The Netherlands (12), France (14), Spain (15), Portugal
(16), Germany (17), Switzerland (18), Italy (19), and Denmark (21). Position of England not to scale. Scale =600 km.

10.4.
NORTH YORKSHIRE
,
ENGLAND
1. Speeton Clay (Norman and Barrett, 2003)
Ornithopoda
Iguanodontia
cf. Camptosaurus hoggi (=Iguanodon
cf. fittoni)
Age: Berriasian (Rawson et al. 1978)
10.7.
DORSET
,
ENGLAND
1. Lower Purbeck Beds (=lower Lulworth
Formation; Galton 1980e; Clements
1993; Ensom 1995; Norman and Barrett
2003)
Ankylosauria
Ankylosauria indet.
Ankylosaur tracks
Age: Berriasian (Allen and Wimbledon
1991 )
2. Middle Purbeck Beds (=upper Lulworth
Formation and lower Durlston
Formation; Owen 1861b, 1874; Delair
1959; Galton 1978, 1980e; Clements
1993; Ensom 1995, 1997; Wright et al.
1998; Wright 1999; Milner 2003;
Norman and Barrett 2003)
Theropoda indet. (=Megalosaurus
dunkeri)
Theropod tracks
Sauropod tracks
Ankylosauria
Nodosauridae indet.
?Nodosaurid tracks
Ornithopoda
Heterodontosauridae
Echinodon becklesii
Iguanodontia
Camptosaurus hoggi
Euornithopod tracks
?Dinosaur eggs
Age: Berriasian (Allen and Wimbledon
1991 )
3. Upper Purbeck Beds (Ensom 1995)
Theropoda tracks
Ankylosaur tracks
Ornithopoda tracks
Age: Berriasian (Allen and Wimbledon
1991 )
4. “Wealden Beds” (Delair 1966; Ensom
1995)
Theropoda
Theropoda indet. (=Megalosaurus sp.)
Theropod tracks
?Sauropoda indet.
Ornithopoda
Iguanodontia
Iguanodon sp.
Euornithopod tracks
Dinosaur tracks
Age: late Barremian–early Aptian
(Rawson et al. 1978)
5. Lower Greensand (Blows 1987)
Ankylosauria incertae sedis
Polacanthus foxii
Age: late Aptian (Rawson et al. 1978)
10.9.
CAMBRIDGESHIRE
,
ENGLAND
1. Cambridge Greensand (Seeley 1869,
1876, 1879; Lydekker 1888a; Le Loeuff
1993; Pereda-Suberbiola and Barrett
1999; Upchurch, pers. comm.)
Theropoda
Avialae
Hesperornithiformes
Enaliornis barretti
Enaliornis sedgwicki
Sauropoda indet. (=Acanthopholis
platypus partim,
Macrurosaurus semnus)
Ornithischia indet. (=Acanthopholis
eucercus)
Ankylosauria incertae sedis
Anoplosaurus curtonotus
Nodosauridae indet.
Ankylosauria indet. (including
Acanthopholis macrocercus
partim, A. platypus partim,
A. stereocercus partim,
Anoplosaurus major partim,
Eucercosaurus tanyspondylus,
Syngonosaurus macrocercus)
Ornithopoda
Ornithopoda indet. (=Acanthopholis
macrocercus partim,
A. stereocercus partim,
Anoplosaurus major partim)
Hadrosauridae indet. (=Trachodon
cantabrigiensis)
Age: late Albian (reworked into
Cenomanian; Rawson et al.
1978)
2. Lower Greensand (Keeping 1883;
Barrett, unpubl. data)
Sauropoda indet. (=?Pelorosaurus sp.)
Ornithopoda
Iguanodontia indet.
Age: late Aptian (Rawson et al. 1978)
10.10.
BEDFORDSHIRE
,
ENGLAND
1. Potton Sands (reworked; Seeley 1874;
Galton 1981a; Benton and Spencer
1995; Barrett and Upchurch, unpubl.
data)
Theropoda indet. (=?Megalosaurus sp.)
Sauropoda
Titanosauriformes indet.
Stegosauria
Stegosauridae
Craterosaurus pottonensis
Ornithopoda
Iguanodontia
Iguanodon cf. bernissartensis
Age: Early Cretaceous (reworked into late
Aptian–early Albian; Rawson
et al. 1978)
10.11.
OXFORDSHIRE
,
ENGLAND
1. Lower Greensand (Benton and Spencer
1995)
Dinosauria indet.
Age: late Aptian (Rawson et al. 1978)
10.14.
WILTSHIRE
,
ENGLAND
1. Upper Purbeck Beds (Huddleston 1876;
Delair 1973)
Theropoda indet. (=Megalosaurus sp.)
Age: Berriasian (Allen and Wimbledon
1991 )
10.17.
ISLE OF WIGHT
,
ENGLAND
1. Wessex Formation (=Wealden Marls;
Hulke 1874b, 1881; Lydekker 1889d,
1890c; Van Straelen 1928; Galton 1971c,
1975; Charig 1980; Delair and Sarjeant
1985; Blows 1987, 1995; Norman 1990b;
Howse and Milner 1993; Le Loeuff 1993;
Insole and Hutt 1994; Benton and
Spencer 1995; Barrett 1996b; Hutt et al.
1996, 2001; Mar till and Hutt 1996;
Wright et al. 1998; Naish 2000; Blows
2001b; Martill and Naish 2001; Naish and
Martill 2002; Upchurch, pers. comm.)
Theropoda
Allosauroidea
Neovenator salerii
Spinosauridae
cf. Baryonyx sp.
Tyrannosauroidea
Eotyrannus lengi
Coelurosauria
Aristosuchus pusillus
Oviraptorosauria indet.
(=Thecocoelurus daviesi)
Theropoda indet. (including
Calamospondylus foxi,
Megalosaurus dunkeri,
Megalosaurus sp.)
?Dromaeosauridae indet.
(=Ornithodesmus cluniculus)
Sauropoda
Diplodocidae indet.
Titanosauriformes
Ornithopsis hulkei
Undescribed titanosauriform
Sauropoda incertae sedis
Oplosaurus armatus
Sauropoda indet. (=Chondrosteosaurus
gigas, Bothriospondylus
magnus, Eucamerotus foxi,
Titanosaurus valdensis,
Cetiosaurus brevis,
Cetiosaurus sp., Pleurocoelus
valdensis)
Sauropod tracks
?Stegosauria indet.
Ankylosauria incertae sedis
Polacanthus foxii (including
Polacanthus becklesi)
Polacanthus sp.
Nodosaurid indet. (including
Polacanthoides ponderosus)
?Ankylosaurian tracks
Ornithopoda
Euornithopoda
Euornithopod tracks
Hypsilophodon foxii
Hypsilophodon sp.
Iguanodontia
Valdosaurus canaliculatus
(?including Camptosaurus
valdensis)
Iguanodon bernissartensis
(including I. seeleyi)
Iguanodon atherfieldensis
(including Sphenospondylus
gracilis, Vectisaurus valdensis)
Iguanodon sp.
558 DINOSAUR DISTRIBUTION AND BIOLOGY

Pachycephalosauria
Yaverlandia bitholus
?Dinosaur eggs (questionable; Carpenter
and Alf 1994)
Age: Barremian (Rawson et al. 1978)
2. Vectis Formation (=Wealden Shale;
Norman 1986; Blows 1987; Wright et al.
1998; Hutt et al. 2001)
Theropod
Theropoda indet. (=Calamospondylus
oweni)
Theropod tracks
Ankylosauria incertae sedis
Polacanthus foxii
Ornithopoda
Euornithopoda
Hypsilophodon foxii
Euornithopoda indet.
Euornithopod tracks
Iguanodontia
Iguanodon atherfieldensis
Age: Barremian–early Aptian (Allen and
Wimbledon 1991)
3. Lower Greensand (Casey 1961; Blows
1987; Benton and Spencer 1995)
Ankylosauria incertae sedis
Polacanthus foxii
Ornithopoda
Iguanodontia
Iguanodon sp.
Age: late Aptian (Rawson et al. 1978)
10.18.
WEST SUSSEX
,
ENGLAND
1. Hastings Beds (=lower Wealden Group;
Lydekker 1890a; Harrison and Walker
1973; Galton 1975, 1985c; Norman
1987b, 1990b; Barrett and Upchurch
1995b; Barrett 1996b; Benton and
Spencer 1995; Upchurch, pers. comm.)
Theropoda
Theropoda indet. (=Megalosaurus
dunkeri)
?Avialae indet. (=Wyleyia valdensis)
Theropoda indet. (=Megalosaurus oweni)
Sauropoda
Titanosauriformes incertae sedis
Pelorosaurus conybearei
Sauropoda indet. (=Pleurocoelus
valdensis, Cetiosaurus brevis)
Stegosauria indet. (=Regnosaurus
northamptoni)
Ankylosauria incertae sedis
Hylaeosaurus armatus
Ornithopoda
Euornithopoda
Euornithopoda indet.
Iguanodontia
Valdosaurus canaliculatus
Iguanodon anglicus (including
Cetiosaurus brevis,
Streptospondylus major)
Iguanodon?bernissartensis
Iguanodon sp.
Age: late Berriasian–Valanginian
(Rawson et al. 1978)
2. Weald Clay (=upper Wealden Group;
Norman 1987b; Benton and Spencer
1995; Blows 1996)
Ankylosauria incertae sedis
Polacanthus rudgwickensis
Ornithopoda
Iguanodontia
Iguanodon atherfieldensis
Iguanodon bernissartensis
Iguanodon sp.
Age: Barremian (Rawson et al. 1978)
10.19.
EAST SUSSEX
,
ENGLAND
1. Hastings Beds (=lower Wealden Group;
Lydekker 1889f, 1892, 1893c; Delair
and Sarjeant 1985; Norman 1987b;
Woodhams and Hines 1989; Barrett and
Upchurch 1995b; Barrett 1996b; Benton
and Spencer 1995; Upchurch 1995, pers.
comm.; Naish 1999b, 1999c, 2003; Blows
2001b)
Theropoda
Tetanurae
Becklespinax altispinax
Allosauroidea indet.
Theropoda indet. (including
Megalosaurus dunkeri,
Megalosaurus sp.)
Theropod tracks
Sauropoda indet. (=Cetiosaurus brevis,
Cetiosaurus sp., Ornithopsis
hulkei, Pleurocoelus valdensis,
Pelorosaurus becklesii)
Thyreophora
Thyreophora indet.
?Stegosauria indet.
Ankylosauria incertae sedis
Hylaeosaurus armatus
Nodosauridae indet.
Ornithopoda
Euornithopoda
Euornithopod tracks
Iguanodontia
Iguanodon dawsoni
Iguanodon fittoni (including
I. hollingtoniensis)
Iguanodon sp.
Age: late Berriasian–Valanginian
(Rawson et al. 1978)
2. Weald Clay (=upper Wealden Group;
Norman 1987b)
Ornithopoda
Iguanodontia
Iguanodon atherfieldensis
Iguanodon sp.
Age: Hauterivian-Barremian (Rawson
et al. 1978)
3. Purbeck Limestone Formation (Benton
and Spencer 1995)
Theropoda indet. (=Megalosaurus sp.)
Age: Berriasian (Allen and Wimbledon
1991 )
10.20.
SURREY
,
ENGLAND
1. Weald Clay (=upper Wealden Group;
Rivett 1953, 1956; Delair and Sarjeant
1985; Charig and Milner 1986; Benton
and Spencer 1995)
Theropoda
Spinosauridae
Baryonyx walkeri
Sauropoda
?Lithostrotia indet.
Ornithopoda
Iguanodontia
Iguanodon atherfieldensis
Iguanodon bernissartensis
Iguanodon sp.
Euornithopod tracks
Age: Barremian (Rawson et al. 1978)
10.21.
KENT
,
ENGLAND
1. Hastings Beds (=lower Wealden Group;
Seeley 1882; Lydekker 1889c; Norman
1987b; Benton and Spencer 1995)
?Dinosauria indet. (=Thecospondylus
horneri)
Theropoda
Avetheropoda incertae sedis
Valdoraptor oweni (=Megalosaurus
oweni)
Theropoda indet. (=Megalosaurus sp.)
Sauropoda indet. (=Ornithopsis sp.)
Ornithopoda
Iguanodontia
Iguanodon sp.
Age: late Berriasian–Valanginian
(Rawson et al. 1978)
2. Lower Greensand (Owen 1884; Casey
1961; Pereda-Suberbiola and Barrett
1999)
Sauropoda indet. (=Dinodocus mackesoni)
Ankylosauria indet. (=Acanthopholis
horridus, Camptosaurus sp.)
Ornithopoda
Iguanodontia
Iguanodon atherfieldensis
Age: middle Aptian-early Albian
(Rawson et al. 1978)
11.2.
PROVINCE DE LA HAINAUT
,
BELGIUM
1. Wealden (Dollo 1882, 1883a, 1883b,
1884, 1909, 1923; Dames 1884; Bertrand
1903; Norman 1986)
Theropoda indet. (=Megalosaurus
dunkeri)
Ornithopoda
Iguanodontia
Iguanodon bernissartensis
Iguanodon atherfieldensis
Dinosaur coprolites
Age: Barremian-Aptian (Taquet 1976;
Norman 1980)
14.2.
DÉPARTEMENT DE LA HAUTE
-
MARNE
,
FRANCE
1. Calcaire à spatangue Formation
(Lapparent and Stchepinsky 1968;
Martin and Buffetaut 1992)
Ornithopoda
Iguanodontia
Iguanodon atherfieldensis
Age: Hauterivian (Lapparent and
Stchepinsky 1968)
2. Argiles à plicatules Formation
(Lapparent and Stchepinsky 1968;
Martin and Buffetaut 1992)
Ornithopoda
Iguanodontia
Iguanodon bernissartensis
Iguanodon atherfieldensis
DINOSAUR DISTRIBUTION 559

Age: early Aptian (Lapparent and
Stchepinsky 1968)
3. Grès et sables piquetés Formation
(Martin and Buffetaut 1992)
Ornithopoda
Iguanodontia
Iguanodon bernissartensis
Age: late Barremian (Martin and
Buffetaut 1992)
14.6.
DÉPARTEMENT DE L
’
AIN
,
FRANCE
1. Unnamed unit (Mojon 2001)
Theropoda indet.
Age: early Berriasian (Mojon 2001)
14.7.
DÉPARTEMENT DU GARD
,
FRANCE
1. Unnamed unit (Buffetaut et al. 1991;
Pérez-Moreno et al. 1993)
Theropoda
Carnosauria
Unnamed allosauroid
Age: Valanginian (Buffetaut et al. 1991)
14.21.
DÉPARTEMENT DE LA SEINE
-
MARITIME
,
FRANCE
1. Unnamed unit (Buffetaut 1984)
Sauropoda indet.
Age: Albian (Buffetaut 1984)
14.22.
DÉPARTEMENT DU PAS
-
DE
-
CALAIS
,
FRANCE
1. Unnamed unit (Lapparent 1967)
Theropoda indet. (=Megalosaurus
insignis)
Age: Valanginian (Lapparent 1967;
Buffetaut et al. 1991)
2. Unnamed unit (Sauvage 1876a)
Theropoda incertae sedis
Erectopus superbus
Age: Albian (=Gault; Huene 1932)
14.26.
DÉPARTEMENT DE LA MEUSE
,
FRANCE
1. Argiles ostréenne (Lapparent and
Stchepinsky 1968; Martin and Buffetaut
1992)
Ornithopoda
Iguanodontia
Iguanodon bernissartensis
Age: early Barremian (Lapparent and
Stchepinsky 1968)
2. Grès et sables piquetés (Lapparent and
Stchepinsky 1968; Damotte et al. 1978;
Martin and Buffetaut 1992)
Sauropoda indet.
Ornithopoda
Iguanodontia
Iguanodon bernissartensis
Age: late Barremian (Lapparent and
Stchepinsky 1968)
3. Unnamed unit (Barrois 1875; Sauvage
1882; Huene 1926a, 1932; Lapparent
1967; Buffetaut et al. 1991)
Theropoda
Tetanurae
Erectopus sauvagei
Erectopus superbus (=Megalosaurus
superbus partim)
Theropoda indet.
(=Megalosaurus sp.)
Age: Albian (Lapparent 1967)
4. Argile du Gault (Martin et al. 1993)
Sauropoda indet.
Age: middle Albian (Magniez 1980)
14.28.
DÉPARTEMENT DE L
’
AUBE
,
FRANCE
1. Sables verts de l’Albien (Buffetaut 1995b;
Knoll et al. 2000)
Sauropoda indet.
Ankylosauria indet.
Age: early Albian (Buffetaut 1995b;
Knoll et al. 2000)
14.29.
DÉPARTEMENT DE L
’
OISE
,
FRANCE
1. Unnamed unit (Lapparent 1946, 1967)
Sauropoda indet.
Age: Albian (Lapparent 1967)
14.30.
DÉPARTEMENT DES ARDENNES
,
FRANCE
1. Unnamed unit (Barrois 1875; Lapparent
1967; Buffetaut et al. 1991; Buffetaut
1995b)
Theropoda indet. (=Megalosaurus sp.)
Age: Albian (Lapparent 1967)
14.31.
DÉPARTEMENT DU VAUCLUSE
,
FRANCE
1. Grès vert (Gervais 1852; Le Loeuff 1993)
Sauropoda indet. (=Aepysaurus
elephantinus)
Age: Albian (Buffetaut et al. 1991)
14.32.
DÉPARTEMENT DES ALPES DE
HAUTE
-
PROVENCE
,
FRANCE
1. “Bevons” beds (Accarie et al. 1995)
Theropoda
Ceratosauria
Genusaurus sisteronis
Age: Albian (Accarie et al. 1995)
15.3.
PROVINCIA DE VALENCIA
,
SPAIN
1. Unnamed unit (Royo y Gómez 1925;
Bataller 1960)
Tetanurae indet. (=Megalosaurus cf.
dunkeri, Megalosaurus sp.)
Sauropoda indet. (=Cetiosaurus sp.)
Age: Valanginian-Barremian (i.e.,
=Wealden; Bataller 1960)
2. El Collado Formation (Casanovas-
Cladellas 1992; Casanovas-Cladellas et al.
1993b, 1995c; Cuenca Bescós et al. 1997)
Theropoda indet.
Sauropoda indet. (=Camarasauridae
indet.)
Stegosauria
Stegosauridae
Dacentrurus armatus
Age: Hauterivian-Aptian (Sanz 1996)
15.4.
PROVINCIA DE TERUEL
,
SPAIN
1. Urbion Group (Casanovas et al. 1995)
Theropod tracks
Sauropod tracks
Age: late Berriasian–Hauterivian
(Casanovas et al. 1995)
15.5.
PROVINCIA DE VIZCAYA
,
SPAIN
1. Monte Grande Formation (Moratalla et
al. 1994a)
Sauropod tracks
Age: Albian (Moratalla et al. 1994a)
15.6.
PROVINCIA DE BURGOS
,
SPAIN
1. Rupelo Formation (Platt and Meyer 1991)
Sauropod tracks
Euornithopod tracks
Age: Berriasian (Platt 1989)
2. Urbion Group (Sanz 1983; Platt and
Meyer 1991; Sanz et al. 1992;
Bengoechea et al. 1993; Moratalla et al.
1994b; Pereda-Suberbiola et al. 1999)
Theropoda
Maniraptora indet.
Theropod tracks
Sauropoda
Sauropoda indet.
Ankylosauria incertae sedis
Polacanthus sp.
Nodosauridae indet.
(=cf. Hylaeosaurus sp.)
Ankylosaur tracks
Ornithopoda
Iguanodontia
Iguanodon atherfieldensis
Euornithopod tracks
Age: Barremian-Aptian (Pereda-
Suberbiola et al. 1999)
3. Tera-Beds (Maisch 1997)
Theropoda indet.
Ornithopoda
Euornithopoda
Euornithopoda indet.
Iguanodontia
Iguanodon cf. fittoni
Age: Berriasian-Valanginian (Maisch
1997)
4. Piedrahita de Muñó Formation (Platt
and Meyer 1991)
Theropod tracks
Age: ?Valanginian-Barremian (Platt 1989)
15.7.
PROVINCIA DE LA RIOJA
,
SPAIN
1. Urbion Group (Casanovas et al. 1995)
Theropod tracks
Sauropod tracks
Ornithopod tracks
Age: late Berriasian–Hauterivian
(Casanovas et al. 1995)
2. Unnamed unit (Viera 1991)
Ornithopoda
Iguanodontia
Iguanodon atherfieldensis
Age: Valanginian (Viera 1991)
3. Castellar Formation (Estes and
Sanchíz1982; Sanz 1982; Buscalioni and
Sanz 1984, 1987; Casanovas-Cladellas
et al. 1984, 1994; Sanz et al. 1984a, 1984b,
1984c, 1987; Kohring 1990; Cuenca et al.
1993; Zinke and Rauhut 1994; Canudo
et al. 1996c, 1997; AmoSanjuan et al.
1999; Ruiz-Omeñaca et al. 1998a, 1998b;
Cuenca-Bescós et al. 1999a, 1999b; Royo-
Torres et al. 1999; Canudo et al. 2001)
560 DINOSAUR DISTRIBUTION AND BIOLOGY

Theropoda
Coelurosauria indet.
?Troodontidae indet.
(=?Paronychodon sp.)
Dromaeosauridae indet.
Theropoda indet.
Theropod tracks
Sauropoda
Eusauropoda incertae sedis
Aragosaurus ischiaticus
?Titanosauriformes indet.
(=“Pleurocoelus” sp.)
Sauropoda indet. (=Camarasaurus sp.)
Sauropod tracks
Ornithischia indet. (=?Echinodon sp.)
Stegosauria indet.
Ornithopoda
Euornithopoda
cf. Hypsilophodon sp.
Iguanodontia indet. (=Iguanodon
cf. atherfieldensis)
Euornithopod tracks
Dinosaur eggs
Age: late Hauterivian–early Barremian
(Cuenca-Bescós et al. 1999b)
4. Camarillas Formation (Buscalioni and
Sanz 1984, 1987; Sanz et al. 1984a,
1984b, 1984c, 1987; Canudo et al. 1996c,
1997, 1999; Ruiz-Omeñaca et al. 1998a,
1998b)
Theropoda
Coelurosauria indet.
Dromeosauridae indet.
Theropoda indet.
Sauropoda
Titanosauriformes indet. (including
cf. Pleurocoelus sp.)
Ornithischia indet. (=Echinodon sp.)
Ankylosauria
Nodosauridae indet.
Ornithopoda
Euornithopoda
Hypsilophodon foxii
Euornithopoda indet. (including
cf. Hypsilophodon sp.)
Iguanodontia
cf. Valdosaurus sp.
Iguanodon bernissartensis
Iguanodon cf. atherfieldensis
Dinosaur eggs
Age: early Barremian (Canudo et al.
1996a, 1996b)
5. Blesa Formation (Cuenca-Bescós et al.
1997, 1999a, 1999b; Ruiz-Omeñaca et al.
1997; Amo 1998)
Theropoda indet.
Sauropoda indet. (=Camarasauridae
indet.)
Ornithopoda
Euornithopoda
Iguanodontia
Iguanodon sp.
Iguanodontia indet.
Euornithopoda indet.
Dinosaur coprolites
Dinosaur eggs
Age: early Barremian (Ruiz-Omeñaca
et al. 1997)
6. Artoles Formation (Canudo et al. 1997;
Ruiz-Omeñaca and Cuenca- Bescós
1995; Ruiz-Omeñaca et al. 1996, 1998a,
1998b; Cuenca-Bescós et al. 1996, 1997)
Theropoda
Spinosauridae indet.
(=Baryonychidae indet.)
Coelurosauria indet.
Dromaeosauridae indet.
Theropoda indet.
Sauropoda
Titanosauriformes indet.
Sauropoda indet.
Ornithopoda
Euornithopoda
Euornithopoda indet.
Iguanodontia
Iguanodon sp.
Iguanodontia indet.
Age: late Barremian (Ruiz-Omeñaca
et al. 1996)
7. Alacón Formation (Auqué Sanz et al.
1986; Canudo et al. 1999)
Sauropoda tracks
Euornithopoda tracks
Age: late Barremian (Canudo et al. 1999)
8. Calizas y margas de Xert Formation
(Royo-Torres and Canudo 1999; Canudo,
pers. comm.)
Sauropoda
Titanosauriformes indet.
Age: early Aptian (Royo-Torres and
Canudo 1999)
9. Forcall Formation (Ruiz-Omeñaca et al.
1997)
Theropoda indet.?
Age: early Aptian (Ruiz-Omeñaca et al.
1997)
10. Enciso Group (Casanovas-Cladellas
and Santafé-Llopis 1971, 1974; Viera
and Torres 1979, 1992, 1996; Viera and
Aguirrezabala 1982, 1984; Casanovas-
Cladellas et al. 1985a, 1985b, 1990a,
1990b, 1991a, 1992d, 1993a, 1993b,
1994, 1995; Aguirrezabala et al. 1985;
Sanz et al. 1985; Martin Escorza 1986;
Moratalla et al. 1988a, 1988b, 1988c,
1992a, 1992b, 1993, 1994a, 1996,
2000; Casanovas et al. 1989a, 1989b;
Moratalla and Sanz 1992; Pérez Lorente
1993; Torres and Viera 1994; Moratalla
2000a, 2000b, 2000c, 2000d, 2000e,
2000f, 2000g, 2000h)
Theropod tracks (including ?avialan
tracks)
Sauropod tracks
Ornithopoda
Euornithopoda
Hypsilophodon foxii
Euornithopod tracks
Iguanodontian tracks
Age: Aptian (Wright et al. 1998)
11. Escucha Formation (Vilanova 1873;
Royo y Gómez1926; Cuenca-Bescós
et al. 1999a, 1999b)
Ornithopoda
Iguanodontia
Iguanodon sp.
Age: Aptian (Cuenca-Bescós et al.
1999a, 1999b)
12. Unnamed unit (Calderon 1876; Bataller
1960)
Ornithopoda
Iguanodontia
Iguanodontia indet. (=Iguanodon sp.)
?Hadrosauridae indet.
(=Trachodon sp.)
Age: Early Cretaceous (Bataller 1960)
13. Unnamed unit (Lapparent et al. 1969)
Ornithopoda indet. (=Iguanodon sp.)
Sauropoda indet.
Age: Hauterivian-Berremian
(=Wealden, Lapparent et al.
1969)
15.8.
PROVINCIA DE LLEIDA
,
SPAIN
1. La Pedrera de Rubies Lithographic
Limestones Formation (Gómez Pallerola
1979, 1986; Barale et al. 1984; Lacasa-
Ruiz 1985, 1989, 1991; Sanz et al. 1988,
1997a)
Theropoda
Avialae
Enantiornithes
Noguerornis gonzalezi
Enantiornithes indet.
Avialae indet. (including Ilerdopteryx
viai)
Age: late Berriasian–early Barremian
(Brenner et al. 1974)
15.9.
PROVINCIA DE CUENCA
,
SPAIN
1. Unnamed unit (Sanz 1986)
Ornithopoda
Iguanodontia
Iguanodon bernissartensis
Age: Valanginian-Hauterivian (Sanz
1986)
2. Unnamed unit (Francés and Sanz 1989)
Theropoda indet.
Sauropoda indet.
Ornithopoda
Iguanodontia
Iguanodon sp.
Age: Hauterivian-Barremian (Francés
and Sanz 1989)
3. Unnamed unit (Lapparent et al. 1969)
Theropoda indet.
Sauropoda indet.
Age: Valanginian-Barremian
(i.e., =Wealden; Lapparent
et al. 1969)
4. Calizas de La Huergina Formation
(Sanz et al. 1988, 1995, 1996; Sanz and
Buscalioni 1992; Sanz and Bonaparte
1992; Pérez-Moreno et al. 1994; Krebs
1995; Pérez-Moreno and Sanz 1995;
Rauhut and Zinke 1995; Rauhut 2002)
Theropoda
Theropoda incertae sedis
cf. Ricardoestesia sp.
Ornithomimosauria
Pelecanimimus polyodon
?Troodontidae indet. (including
cf. Euronychodon sp.,
cf. Paranychodon sp.)
DINOSAUR DISTRIBUTION 561

?Dromaeosauridae indet.
Avialae
Enantiornithes
Concornis lacustris
Eoalulavis hoyasi
Iberomesornis romerali
Theropoda indet.
Sauropoda indet. (including
Brachiosauridae indet.)
Ornithopoda
Euornithopoda indet.
Iguanodontia
Iguanodon sp.
Age: late Barremian (Frengenal-Martínez
and Meléndez 1995; Rauhut
and Zinke 1995)
15.10.
PROVINCIA DE CASTELLÓN
,
SPAIN
1. Arcillas de Morella Formation (Royo y
Gómez 1918, 1921, 1926; Bataller 1960;
Santafé-Llopis et al. 1979, 1981, 1982;
Santafé et al. 1981; Sanz et al. 1983
[including Capas Rojas Formation])
Theropoda indet.
Sauropoda indet. (including
Brachiosauridae indet.)
Ankylosauria
?Nodosauridae indet.
Ornithopoda
Euornithopoda
Hypsilophodon sp.
Iguanodontia
Iguanodon bernissartensis
Age: early Aptian (Canerot 1974; Canerot
et al. 1982)
16.2.
DISTRITO DO SANTARÉM
,
PORTUGAL
1. Conglomerados de Caranguejeira
(Sauvage 1897–98; Lapparent and
Zbyszewski 1957)
Theropoda indet. (=Morosaurus marchei)
Sauropoda indet. (=Pelorosaurus
humerocristatus)
Age: Aptian-Cenomanian (Manupella
et al. 2000)
16.4.
DISTRITO DE LISBOA
,
PORTUGAL
1. Unnamed unit (Galton 1980a; Mateus,
pers. comm.)
Ankylosauria incertae sedis
Dracopelta zbyszewskii
Age: Early Cretaceous (Mateus,
pers. comm.)
16.5.
DISTRITO DO SETÚBAL
,
PORTUGAL
1. Unnamed unit (Antunes 1976; Madeira
and Dias 1983)
Theropod tracks
Sauropod tracks
Euornithopod tracks
Age: Hauterivian (Antunes 1976)
2. Grès à Dinosauriens Formation
(Lapparent and Zbyszewski 1957; Galton
1981b, 1994; Mateus, pers. comm.)
Theropoda indet. (including
Megalosaurus superbus)
Sauropoda
Titanosauriformes indet. (=“Astrodon”
valdensis)
Ornithopoda
Iguanodontia
Iguanodon sp.
Age: early Barremian (Rey 1972)
17.1.
NIEDERSACHSEN
,
GERMANY
1. Obernkirchen Sandstein (=Wealden;
Meyer 1857; Koken 1887; Ballerstedt
1922; Nopcsa 1923b; Kuhn 1958b; Sues
and Galton 1982; McCrea et al. 2001a)
Theropoda indet. (=Megalosaurus
dunkeri)
Theropod tracks
?Ornithischian tracks
Ankylosaur tracks
Ornithopoda
Euornithopoda
Euornithopod tracks
Iguanodontia
Iguanodon sp.
Pachycephalosauria
Stenopelix valdensis
Age: Berriasian (Schmidt 1969)
17.8.
NORDRHEIN
-
WESTPHALEN
,
GERMANY
1. Unnamed unit (Sachs 1997)
?Ankylosauria
?Nodosauridae indet.
Age: Berriasian (Sachs 1997)
2. Wealden (Huckriede 1982; Hölder and
Norman 1986; Norman 1987b)
Theropoda indet. (=Megalosaurus sp.)
Ornithopoda
Euornithopoda
?Euornithopoda indet.
(=Hypsilophodon sp.)
Iguanodontia
Iguanodon atherfieldensis
Iguanodon bernissartensis
Age: Aptian (Huckriede 1982)
18.6.
KANTON UNTERWALDEN
,
SWITZERLAND
1. Upper Schrattenkalk Formation (Meyer,
pers. comm.)
Euornithopod tracks
Age: late Aptian (Meyer, pers. comm.)
19.4.
REGIONE FRIULI
–
VENEZIA GIULIA
,
ITALY
1. Calcare di Cellina Formation (Dalla
Vecchia and Venturini 1995, 1996;
Venturini 1995; Dalla Vecchia 1997a,
1998b, 1999a)
Theropod tracks
Sauropod tracks
Age: late Hauterivian–early Barremian
(Dalla Vecchia and Venturini
1995)
19.7.
REGIONE CAMPANIA
,
ITALY
1. “Calcari selciferi ed ittiolitiferi di
Pietraroia” (Dal Sasso and Signore 1998)
Theropoda
Coelurosauria incertae sedis
Scipionyx samniticus
Age: early Albian (Catenacci and
Manfredini 1963; Bravi
1996)
19.8.
REGIONE PUGLIA
,
ITALY
1. San Giovanni Rotondo Formation
(Gianolla et al. 2000)
Theropod tracks
?Ornithopod tracks
Age: Hauterivian-Barremian (Gianolla
et al. 2000)
21.1.
BORNHOLM AMT
,
DENMARK
1. Unnamed unit (Christiansen and Bonde
2003)
Theropoda
Dromaeosauridae indet.
(=Dromaeosauroides
barnhomensis)
Age: Valanginian-Barremian (=Wealden
equivalent; Christiansen and
Bonde 2003)
22.1.
SVALBARD
,
NORWAY
1. Festningen Sandstone (=Helvetiafjellet
Formation; Lapparent 1962; Edwards
et al. 1978)
Theropod tracks
Euornithopod tracks
Age: Barremian (Lapparent 1962)
29.1.
JUDET
¸
UL BIHOR
,
ROMANIA
1. Bauxite of Cornet (Jurcsák and Popa
1979, 1983; Jurcsák 1982; Kessler 1984;
Jurcsák and Kessler 1985; Kessler and
Jurcsák 1986; Benton et al. 1997; Tallodi-
Pos¸mos¸ anu and Popa 1997; Pos¸mos¸anu
2000)
Theropoda
Neornithes
Palaeocursornis corneti
Eurolimnornis corneti
?Avialae indet. (=Archaeopteryx sp.)
Theropoda indet. (=Aristosuchus sp.)
Ornithopoda
Euornithopoda
Hypsilophodon sp.
Euornithopoda indet.
(=cf. Thescelosaurus sp.)
Iguanodontia
Valdosaurus canaliculatus
?Iguanodontia indet.
(=Dryosaurus sp.)
Iguanodon sp. (=I. mantelli)
Age: Berriasian-Hauterivian (Patrulius
et al. 1983)
29.2.
JUDET
¸
UL CONSTANT
¸
A
,
ROMANIA
1. Unnamed unit (Simionescu 1913)
Theropoda indet. (=Megalosaurus
cf. superbus)
Age: Valanginian-Barremian
(Simionescu 1913)
28.1.
ISTARSKA ZUPANIJA
,
CROATIA
1. Unnamed unit (Lockley et al. 1994b)
?Sauropod track
Age: Berriasian (Tisljar et al. 1983)
2. Unnamed unit (Dalla Vecchia et al. 2000)
Theropod tracks
Sauropod tracks
Age: Hauterivian (Dalla Vecchia et al.
2000)
562 DINOSAUR DISTRIBUTION AND BIOLOGY

3. Unnamed unit (Boscarolli et al. 1993;
Dalla Vecchia 1994, 1997a, 1997b,
1998a, 1998b, 1999b, 2001a, 2001b)
Theropoda
Theropoda indet.
Sauropoda
?Camarasauridae indet.
Titanosauriformes indet.
Sauropoda indet. (=Histriasaurus
boscarollii)
Age: late Hauterivian–early Barremian
(Boscarolli et al. 1993; Dini
et al. 1998)
4. Unnamed unit (Velic and Tislja 1987; Dalla
Vecchia et al. 1993; Dalla Vecchia 1998b)
Theropod tracks
Age: late Barremian (Velic and Tislja 1987)
5. Unnamed unit (Gogala and Pavlovec
1978; Dalla Vecchia and Tarlao 1995)
?Theropod tracks
Age: ?late Barremian (Velic and Tislja
1987; Dalla Vecchia and
Tarlao 1995)
6. Unnamed unit (Bachofen-Echt 1925a,
1925b; Dalla Vecchia and Tarlao 1995,
2000; Dalla Vecchia 1997a, 1998b)
Theropod tracks
Ornithopod tracks
Age: late Albian (Velic and Tisljar 1987;
Tunis and Venturini 2000)
7. Unnamed unit (Dalla Vecchia et al.
1993; Dalla Vecchia 1994, 1996a,
1997a, 1998b; Dalla Vecchia and
Tarlao 2000)
Sauropod tracks
Theropod tracks
Age: late Albian (Dalla Vecchia et al.
1993; Tunis and Venturini
2000)
DINOSAUR DISTRIBUTION 563
Asia
Asian records of Early Cretaceous dinosaurs (107 locations,
producing a 160% increase) compare well with those of the Late
Cretaceous on this continent as well as elsewhere (figs. 24.7,
24.16, 24.20, 24.21, and 24.22). Studies of Early Cretaceous
dinosaurs in northern China and Mongolia include those by
B. Brown, P. R. Currie, Dong Z.-M., W. Granger, K. Mikhailov,
H. F. Osborn, E. Schlaikjer, A. K. Rozhdestvensky, D. A. Russell,
C.-C. Young, and Zhao Z.-C. But by far the most important dis-
covery in China, if not the world, was a Lagerstatt in Liaoning
Province. It is considered by many to be as important as the
discovery of Archaeopteryx in the last century. Both avialan and
nonavialan dinosaurs, with and without feathers, are presently
under investigation by Ji Q., Ji S.-A., P. R. Currie, and others. The
dinosaurs of South Korea, known mostly from ichnofaunas,
have been studied by H.-M. Kim, S.-K. Lim, and M. G. Lockley,
among others. In Japan, scattered but intriguing assemblages
are known through the research of H. Hasegawa, Y. Azuma,
Y. Tomida, and M. Matsukawa. J. H. Hoffet, E. Buffetaut, R. In-
gavat, V. Suteethorn, P. Taquet, and R. Allain have conducted
important work on the Early Cretaceous dinosaurs of Thailand
and Laos. Finally, Asian Russia, Kazakhstan, Tadzhikistan, Uzbek-
istan, and Kirghizia have been investigated by A. N. Riabinin,
A. K. Rozhdestvensky, V. P. Novikov, and L. A. Nessov. (A series
of ornithopod trackways is reported from Nei Mongol Zizhiqu,
People’s Republic of China [Gao et al. 1981], but they are dated
only as Cretaceous in age [location 48.11-15].)
37.2.
YAKUTSKAYA AVTONOMNAYA OBLAST
,
RUSSIA
1. Sangarskaya Svita (Rozhdestvensky
1973b)
Ankylosauria
Ankylosauridae indet.
Age: Berriasian-Hauterivian
(=Neocomian;
Rozhdestvensky 1973b)
37.3.
ULYANOVSKAYA AV TONOMNAYA
OBLAST
,
RUSSIA
1. Unnamed unit (Efimov 1997)
Sauropoda indet. (=Brachiosauridae
indet.)
Age: Hauterivian (Efimov 1997)
37.4.
GORNO
-
ALTAYASKAYA AVTONOMNAYA
OBLAST
,
RUSSIA
1. Shestakovskaya Svita (=Ilekskaya
Svita; Rozhdestvensky 1955b, 1960a;
Novikov et al. 1998; Alifanov et al. 1999;
Leshchinskiy et al. 2001; Averianov
et al. 2002)
Theropoda
?Tyrannosauridae indet.
Troodontidae indet.
Dromaeosauridae indet.
Avialae indet.
Sauropoda
Titanosauria indet.
Ceratopsia
Psittacosauridae
Psittacosaurus mongoliensis
Psittacosaurus cf. xinjiangensis
Psittacosaurus sibericus
Age: Aptian-Albian (Averianov et al. 2002)
37.5.
BURIAT
-
MONGOLSKAYA
AVTONOMNAYA OBLAST
,
RUSSIA
1. Turginskaya Svita (Riabinin 1914)
Theropoda
Tetanurae incertae sedis
?Chilantaisaurus sibiricus
Age: Berriasian-Hauterivian
(=Neocomian; Osmólska,
pers. comm.)
2. Murtoi Formation (Riabinin 1937;
Rozhdestvensky 1970a; Nessov and
Starkov 1992; Starkov 1999)
Theropoda indet. (two species?)
Sauropoda indet. (including cf.
Chiayusaurus sp.)
Ankylosauria indet.
Age: late Barremian–middle Aptian
(Nessov and Starkov 1992)
3. Unnamed unit (Rozhdestvensky 1968b,
1970b; Kurzanov, pers. comm.)
Theropoda indet. (=?Therizinosaurus sp.)
Sauropoda indet.
Age: Early Cretaceous (Kurzanov, pers.
comm.; Osmólska, pers.
comm.)
38.2.
K
’
ALAK
’
IK
’
UT
’
AISI
,
GEORGIA
1. Unnamed unit (Gabuniya 1951, 1958;
Kurzanov, pers. comm.)
Theropod tracks
Euornithopod tracks
Age: Berriasian-Hauterivian
(=Neocomian;
Rozhdestvensky 1973b)
39.2.
AQTOBE OBLYSY
,
KAZAKHSTAN
1. Neocomian Sands (Riabinin 1931b)
Theropoda indet. (=Embasaurus
minax)
Age: Berriasian-Hauterivian
(=Neocomian; Riabinin
1931b)
40.2.
SURKHONDARYO WILOYATI
,
UZBEKISTAN
1. Kiezeltashskaya Formation (Lockley,
pers. comm.)
Theropod tracks
Sauropod tracks
Age: Hauterivian-Barremian (Lockley,
pers. comm.)
40.3.
QORAQALPOGHISTON RESPUBLIKASI
,
UZBEKISTAN
1. Lower Khodzhakul Formation
(Nessov et al. 1994; Kurochkin
2000)
Theropoda
Avialae
Ornithurae
Horezmavis eocretaceous
Ornithopoda
Hadrosauridae indet.
Age: late Albian (Nessov et al. 1994)
42.1.
OSH OBLAST
,
KIRGHIZIA
1. Khodzhaosmansk Formation (Nessov
and Kaznachkin 1986)
Dinosaur eggs
Age: Aptian-?Albian (Nessov and
Kaznachkin 1986)

43.2.
SOUTHWESTERN TADZHIKISTAN
(
NO OBLAST
-
LEVEL DIVISION EXISTS
FOR THIS REGION
)
1. Luchak Svita (Lockley, pers. comm.)
Theropod tracks
Sauropod tracks
Age: Albian (Lockley, pers. comm.)
43.3.
VILOYATI KHATLON
,
TADZHIKISTAN
1. Shirabad Svita (McCrea et al. 2001a)
?Theropod tracks
?Sauropod tracks
?Ankylosaur tracks
Age: Albian (McCrea et al. 2001a)
47.1.
GOV
’
ALTAI
,
MONGOLIA
1. Unnamed unit (Nessov and Kaznachkin
1986)
Dinosaur eggs
Age: Berriasian-Hauterivian
(=Neocomian; Nessov and
Kaznachkin 1986)
47.2.
KHOVD
,
MONGOLIA
1. Tsagaantsav Svita (Osmólska,
pers. comm.)
Sauropoda indet.
Ceratopsia
Psittacosauridae indet.
Age: Valanginian (Osmólska,
pers. comm.)
47.3.
BAYANKHONGOR
,
MONGOLIA
1. Öndörukhaa Formation (Kurochkin
1985, 1991)
Theropoda
Avialae
Ornithurae
Ambiortus dementjevi (?including
Holbotia ponomarenkoi)
Age: Hauterivian-Aptian (Krassilov 1982)
2. Döshuul Formation (Mikhailov et al.
1994)
Dinosaur eggs
Age: Aptian (Mikhailov et al. 1994)
3. Unnamed unit (Kalandadze and
Kurzanov 1974)
Ankylosauria
Ankylosauridae indet.
Ceratopsia
Psittacosauridae
Psittacosaurus mongoliensis
Age: Aptian-Albian (Shuvalov 1974)
4. Unnamed unit (Kurochkin 2000)
Theropoda
Avialae
Enantiornithes indet.
Age: Early Cretaceous (Kurochkin 2000)
47.4.
ÖMNÖGOV
’,
MONGOLIA
1. Sainshand Svita (Kurzanov, pers. comm.;
Osmólska, pers. comm.)
Dinosaur eggs
Age: Albian-Cenomanian (Kurzanov,
pers. comm.; Osmólska,
pers. comm.)
2. Unnamed unit (Watabe et al. 2001)
Theropoda
Avialae indet.
Age: Early Cretaceous (Watabe et al.
2001)
47.5.
ÖVÖRKHANGAI
,
MONGOLIA
1. Döshuul Formation (Mikhailov et al.
1994)
Dinosaur eggs
Age: Aptian (Mikhailov et al. 1994)
2. Hühteeg Svita (=Ondai Sair and Oshih
formations; Osborn 1923a, 1924c;
Rozhdestvensky 1952a; Martinsson
and Shuvalov 1973; Coombs 1982;
Tumanova 1985; Nessov and
Kaznachkin 1986; Norman 1996;
Barrett et al. 2002b)
Theropoda indet. (=Prodeinodon
mongoliensis)
Sauropoda
Sauropoda indet. (=Asiatosaurus
mongoliensis)
Ankylosauria
Ankylosauridae
Shamosaurus scutatus
Ornithopoda
Iguanodontia
Iguanodon bernissartensis
(=Iguanodon orientalis)
Iguanodontia indet.
Ceratopsia
Psittacosauridae
Psittacosaurus mongoliensis
(including Protiguanodon
mongoliensis, Psittacosaurus
protiguanodonensis)
Age: Aptian-Albian (Shuvalov 1974)
47.6.
DUNDGOV
’,
MONGOLIA
1. Khooldzin Svita (Shuvalov 1974;
Osmólska, pers. comm.)
564 DINOSAUR DISTRIBUTION AND BIOLOGY
FIGU RE 24.20. Late Cretaceous localities in Mongolia (47), People’s Republic of China (48), South Korea (50), and Japan (53). Scale =600 km.

Theropoda
Dromaeosauridae indet.
Ceratopsia
Psittacosauridae
Psittacosaurus mongoliensis
Age: Aptian-Albian (Osmólska,
pers. comm.)
2. Shinekhudag Svita (=Khooldzin Svita;
Barsbold and Perle 1984; Norman
1996)
Theropoda
Ornithomimosauria
Harpymimus okladnikovi
Ornithopoda
Iguanodontia
Iguanodon bernissartensis
(=Iguanodon orientalis)
Ceratopsia
Psittacosauridae
Psittacosaurus mongoliensis
Age: late Albian (Hicks et al. 1999)
47.7.
DORNOGOV
’,
MONGOLIA
1. Bayan Bologai Formation (Gilmore
1933b; Coombs 1987)
?Ankylosauria
?Ankylosauridae indet.
Ornithopoda
?Hadrosauridae indet.
Age: ?early Berriasian (Morris 1936)
2. Hühteeg Svita (=Upper Züünbayan
Svita; Barsbold and Perle 1983; Norman
1998; Osmólska, pers. comm.)
Theropoda
Troodontidae indet.
Theropoda indet.
Ankylosauria
Ankylosauridae
Shamosaurus scutatus
Ornithopoda
Iguanodontia
Altirhinus kurzanovi
Ceratopsia
Psittacosauridae
Psittacosaurus mongoliensis
Age: late Albian (Hicks et al. 1999)
48.1.
XIZANG ZIZHIQU
,
PEOPLE
’
S REPUBLIC
OF CHINA
1. Lura Formation (?=Loe-ein Formation;
Zhao 1983; Dong 1990b; Sun et al. 1992;
Zhao unpubl. data)
Theropoda
Tetanurae indet.
?Coelurosauria indet.
Sauropoda indet. (=cf. Asiatosaurus
kwangshiensis)
Stegosauria
Stegosauridae
Monkonosaurus lawulacus
Age: Early Cretaceous (Sun et al. 1992;
Yao et al., in press)
2. Unnamed unit (Hao et al. 1986)
Undescribed theropods (two species?)
Undescribed sauropods
Age: Early Cretaceous (Hao et al. 1986)
48.2.
SICHUAN
,
PEOPLE
’
S REPUBLIC
OF CHINA
1. Jianguan Formation (Zhen et al. 1987;
Lockley and Matsukawa 1998)
Theropod tracks
Euornithopod tracks
Age: Aptian-Albian (Hao and Guan
198 4)
48.3.
YUNNAN
,
PEOPLE
’
S REPUBLIC
OF CHINA
1. Upper Jingxing Formation (Yeh 1975;
Zhao 1983)
Theropoda
Tyrannosauroidea indet.
(=Tyrannosaurus
lanpingensis)
Age: Berriasian-Hauterivian (Chen 1983)
48.4.
LIAONING
,
PEOPLE
’
S REPUBLIC
OF CHINA
1. Jiufotang Formation (Sereno et al. 1988;
Zhou et al. 1992; Dong 1993c; Wang
et al. 1999a, 1999b; Xu et al. 2000c;
Zhang et al. 2001; Zhou and Zhang
2001, 2002a, 2002b, 2003; Ji et al. 2003)
Theropoda
Dromaeosauridae
Microraptor zhaoianus
Avialae
Avialae incertae sedis
Sapeornis chaoyangensis
Shenzhouraptor sinensis
Jeholornis prima
Enantiornithes
Boluochia zhengi
Cathayornis yandica
Eocathayornis walkeri
Sinornis santensis
Cathayornis caudatus
Longchengornis sanyannensis
DINOSAUR DISTRIBUTION 565
FIGU RE 24.21. Middle Jurassic to Late Cretaceous localities in Moskva Oblast, Russia (37), Georgia (38), Kazakhstan (39),
Uzbekistan (40), Turkmenist an (41), Kyrgyzia (42), and Tadzhikistan (43). Scale =600 km.

Cuspirostrisornis hinfengi
Largirostrornis sexdentornis
Songlingornis linghensis
Longipteryx chaoyangensis
Ornithurae
Chaoyangia beishanensis
Yanornis martini
Yixianornis grabaui
Sauropoda indet.
Ceratopsia
Psittacosauridae
Psittacosaurus meileyingensis
Psittacosaurus mongoliensis
Age: Early Cretaceous (Sereno et al.
1988); Late Jurassic (Wang
et al. 1999b)
2. Yixian Formation (includes
Chaomidianzi Formation) (Hou et al.
1995a; Ji and Ji 1996, 1997; Hou 1997;
Chen et al. 1998; Ji et al. 1998, 1999,
2001; Xu and Wang 1998; Chiappe et al.
1999a; Hou and Chen 1999; Hou et al.
1999; Wang et al. 1999a; Xu et al. 1999a,
1999b, 2000a, 2000c, 2001a, 2001b,
2002a, 2002b, 2002c, 2003; Zhang and
Zhou 2000; Zhou and Wang 2000; Zhou
et al. 2000; Currie and Chen 2001; Wang
and Xu 2001a, 2001b; Xu and Wu 2001;
Ji et al. 2003; You et al. 2003c; Hou et al.
2004; Hwang et al. 2002, 2004; Xu and
Wang 2003)
Theropoda
Compsognathidae
Sinosauropteryx prima
Huaxiagnathus orientalis
Therizinosauroidea
Beipiaosaurus inexpectus
ornithomimosauria
Shenzhousaurus orientalis
Maniraptora incertae sedis
Yixianosaurus longimanus
Oviraptorosauria
Caudipteryx zoui
Caudipteryx dongi
Incisivosaurus gauthieri
Troodontidae
Sinovenator changii
Dromaeosauridae
Sinornithosaurus milllenii
Microraptor gui
Microraptor zhaoianus
Undescribed dromaeosaurid
Avialae
Protarchaeopteryx robusta
Avialae incertae sedis
Jibeinia luanhera
Confuciusornithidae
Confuciusornis sanctus (including
Confuciusornis chuonzhous,
C. suniae)
Confuciusornis dui
Changchengornis hengdaoziensis
Enantiornithes
Protopteryx fengningensis
Eoenantiornis buhleri
Liaoxiornis delicatus
Longirostravis hani
Ornithurae
Liaoningornis longidigitus
Theropoda indet.
Sauropoda indet.
Ankylosauria incertae sedis
Liaoningosaurus paradoxus
Ornithopoda
Euornithopoda
Jeholosaurus shangyuanensis
Iguanodontia
Jinzhousaurus yangi
Ceratopsia
Psittacosauridae
Hongshanosaurus houi
Psittacosaurus sp.
Neoceratopsia
Liaoceratops yanzigouensis
566 DINOSAUR DISTRIBUTION AND BIOLOGY
FIGU RE 24.22. Late Jurassic to Late Cretaceous localities in Yakutskaya Avtonomnaya Oblast (37.2), Gorno-Atlayaskaya Avtonomnaya Oblast
(37.4), Buriatmongolskaya Avtonomnaya Oblast (37.5), Yevreyaskaya Avtonomnaya Oblast (37.7), Chukotka Avtonomnaya Oblast (37.8),
Amurskaya Avtonomnaya Oblast (37.10), Sachalinskaya Oblast (37.11), Russia, north-central and eastern Asia. Scale =600 km.

Dinosaur eggs
Age: Barremian (Xu and Wang 1998;
Barrett 2000b; Smith et al.
2001a; Wang et al. 2001b;
Swisher et al. 2002)
3. Sunjiawan Formation (Dong 2002; You
et al. 2003a) (not plotted)
Ankylosauria incertae sedis
Crichtonsaurus bohlini
Ornithopoda
Iguanodontia
Shuangmiaosaurus gilmorei
Age: Cenomanian-Turonian (Dong
2002)
48.5.
XINJIANG UYGUR ZIZHIQU
,
PEOPLE
’
S REPUBLIC OF CHINA
1. Tugulu Group (Dong 1973a, 1992, 1993;
Sereno and Chao 1988; Shen and Mateer
1992; Brinkman et al. 2001)
Theropoda
Tetanurae incertae sedis
Kelmayisaurus petrolicus
Coelurosauria incertae sedis
Tugulusaurus faciles
Theropoda indet. (=Phaedrolosaurus
ilikensis)
Sauropoda indet. (=Asiatosaurus
mongoliensis)
Stegosauria
Stegosauridae
Wuerhosaurus homheni
Ornithopoda
Hadrosauridae indet.
Ceratopsia
Psittacosauridae
Psittacosaurus xinjiangensis
Age: ?Valanginian-Albian (Shen and
Mateer 1992)
2. Kezilesu Group (Yao et al., in press)
Ceratopsia
Psittacosaurus sp.
Age: Early Cretaceous (Yao et al.,
in press)
48.6.
SHAANXI
,
PEOPLE
’
S REPUBLIC
OF CHINA
1. Donghe Group (Xue et al. 1996)
Theropod tracks
Age: Early Cretaceous (Xue et al. 1996)
48.7.
JILIN
,
PEOPLE
’
S REPUBLIC OF CHINA
1. Tongfosi Formation (Matsukawa et al.
1995; Lockley and Matsukawa 1998)
Theropod tracks
Euornithopod tracks
Age: Hauterivian (Matsukawa et al.
1995)
2. Quantou Formation (Wood et al. 2001)
Undescribed theropods
Undescribed ornithischians
Dinosaur eggs
Age: ?Aptian (Wood et al. 2001)
48.8.
GANSU
,
PEOPLE
’
S REPUBLIC
OF CHINA
1. Xinminbao Group (Bohlin 1953; Dong
1993a, 1997a, 1997b, 1997c; Dong and
Azuma 1997; Dong and Yu 1997; Lu
1997; Xu 1997; Tang et al. 2001b; You et
al. 2003b; Barrett et al. 2002b; You et al.
2001, 2003, in press; You and Dodson
2003)
Theropoda
Allosauroidea indet.
Therizinosauroidea
?Nanshiungosaurus bohlini
Undescribed ornithomimid
Troodontidae
Sinornithoides sp.
Dromaeosauridae indet.
Theropoda indet.
Sauropoda
Titanosauria
Gobititan shenzhouensis
Titanosauriformes indet.
(=?Chiayusaurus sp.,
?Mongolosaurus sp.)
Sauropoda indet. (=Chiayusaurus
lacustris, Nemegtosauridae
indet., Mamenchisauridae
indet.)
Ankylosauria
Ankylosauridae indet.
Ankylosauria indet. (=Stegosaurides
excavatus)
Ornithopoda
Euornithopoda
Euornithopoda indet. (=Siluosaurus
zhangqiani)
Iguanodontia
Probactrosaurus mazongshanensis
Equijubus normani
Ceratopsidae
Psittacosauridae
Psittacosaurus mazongshanensis
Neoceratopsia
Archaeoceratops oshimai
Neoceratopsia indet.
(=Microceratops gobiensis,
Microceratops sulcidens)
Age: Barremian-Albian (Tang et al. 2001b)
2. Xiagou Formation (Hou and Liu 1984)
(not plotted)
Theropoda
Aves
Ornithurae
Gansus yumenensis
Age: Early Cretaceous (Hou and Liu
198 4)
48.9.
SHANDONG
,
PEOPLE
’
S REPUBLIC
OF CHINA
1. Qingshan Formation (Wiman 1929;
Young 1958a; Chao 1962; Dong 1992,
1993c
Sauropoda indet.
Ceratopsia
Psittacosauridae
Psittacosaurus sinensis
Psittacosaurus youngi
Age: Aptian-Albian (Mateer, pers.
comm.)
48.10.
HEBEI
,
PEOPLE
’
S REPUBLIC
OF CHINA
1. Tujingzi Formation (Dong 1987; Russell
and Zhao 1996)
Ceratopsia
Psittacosauridae
Psittacosaurus sp.
Age: early Albian (Russell and Zhao
1996)
2. Xiguayuan Formation (You and Azuma
1995; Lockley and Matsukawa 1998)
Theropod tracks
Euornithopod tracks
Age: Early Cretaceous (You and Azuma
1995)
3. Yixian Formation (Yao et al., in press)
Ceratopsia
Psittacosaurus sp.
Age: Barremian (Xu and Wang 1998;
Barrett 2000a; Smith et al.
2001a; Swisher et al. 2002)
48.11.
NEI MONGOL ZIZHIQU
,
PEOPLE
’
S REPUBLIC OF CHINA
1. Luohandong Formation (Russell and
Zhao 1996)
Ceratopsia
Psittacosauridae indet.
Age: ?Valanginian-?Albian (Russell and
Zhao 1996)
2. Qagannur Formation (Dong 1992,
1993c)
Sauropoda indet.
Stegosauria indet.
Age: ?Valanginian-?Albian (Russell and
Zhao 1996)
3. Zhidan Group (Dong 1993c)
Ceratopsia
Psittacosauridae
Psittacosaurus sp.
Age: ?Valanginian-?Albian (Russell and
Zhao 1996)
4. Jing Chuan Formation (White et al.
2001; Wright et al. 2001)
Theropod tracks (including avialan
tracks)
Sauropod tracks
Age: Barremian-Aptian (White et al.
2001)
5. On Gong Formation (Gilmore 1933b;
Barrett et al. 2002b; =“Tairum Nor”
formation, Reeds 1932; Gilmore 1933b;
=“Shirigu” formation, Reeds 1932)
Dinosauria indet.
Theropoda
Ornithomimidae indet.
Theropoda indet.
Sauropoda indet. (=Mongolosaurus
haplodon)
Age: Aptian-Albian (i.e., approximately
equivalent with the Ondai
Sair Formation; Dong 1993c)
6. Dashuigou Formation (Chow and
Rozhdestvensky 1960; Hu 1964; Sun
et al. 1992; Dong 1993c; Clure 1998b)
Theropoda
?Tyrannosauroidea
“Chilantaisaurus” maortuensis
Ornithopoda
Iguanodontia
Probactrosaurus alashanicus
Probactrosaurus gobiensis
Age: Albian (Dong, pers. comm.)
DINOSAUR DISTRIBUTION 567

7. Lisangou Formation (Young 1931;
Cheng 1983)
Ceratopsia
Psittacosauridae
Psittacosaurus guyangensis
Psittacosaurus osborni
8. Xinpongnaobao Formation (Young
1931 )
Ceratopsia
Psittacosauridae
Psittacosaurus osborni
Age: ?Aptian-Albian (Dong, pers.
comm.)
9. Ulansuhai Formation (Hu 1964; Sun
et al. 1992; Kobayashi et al. 1999, 2001;
Vickaryous et al. 2001b; Kobayashi and
Lü 2003))
Theropoda
Tetanurae incertae sedis
Chilantaisaurus tashuikouensis
Ornithomimosauria
Sinornithomimus dongi
Ankylosauria
Ankylosauridae
Gobisaurus domoculus
Ornithopoda
Iguanodontia indet.
Age: Aptian-?Albian (Dong, pers.
comm., contra Dong 1980);
?early Late Cretaceous
(Kobayashi and Lü 2003)
10. Bayin-Gobi Formation (Dong 1993b;
Russell and Dong 1993a; Russell and
Zhao 1996)
Theropoda
Therizinosauroidea
Alxasaurus elesitaiensis
Theropoda indet.
Sauropoda indet.
Ceratopsia
Psittacosauridae
Psittacosaurus sp.
Age: Albian (Russell and Dong 1993a)
11. Ejinhoro Formation (Dong 1993a,
1993b; Russell and Dong 1993b; Russell
and Zhao 1996; Currie and Dong
2001a)
Theropoda
Troodontidae
Sinornithoides youngi
Sauropoda indet. (=Diplodocidae indet.,
cf. Chiayusaurus sp.)
Stegosauria
Stegosauridae
Wuerhosaurus ordosensis
Stegosauridae indet.
Ceratopsia
Psittacosauridae
Psittacosaurus neimongoliensis
Psittacosaurus ordosensis
Psittacosauridae indet.
Age: Early Cretaceous (Dong 1993a)
12. Unnamed unit (Bohlin 1953)
Theropoda indet. (=Prodeinodon sp.)
Ankylosauria indet. (=Sauroplites scutiger)
Ceratopsia
Psittacosauridae
?Psittacosaurus mongoliensis
Age: Early Cretaceous (not younger than
mid-Aptian; Eberth et al.
1993)
13. Unnamed unit (Li pers. comm.)
Theropod tracks
Age: Early Cretaceous (Zhen et al. 1989)
14. Yijinholuo Formation (Hou 1994)
Theropoda
Avialae
Enantiornithes
Otogornis genghisi
Age: Early Cretaceous (Hou 1994)
48.12.
HENAN
,
PEOPLE
’
S REPUBLIC
OF CHINA
1. Sangping Formation (Xu et al. 2000b)
Ornithopoda
Iguanodontia
Nanyangosaurus zhugeii
Dinosaur eggs
Age: ?Albian (Xu et al. 2000b)
48.13.
ZHEJIANG
,
PEOPLE
’
S REPUBLIC
OF CHINA
1. Jinhua Formation (Tang et al. 2001a)
Sauropoda
Titanosauridae
Jiangshanosaurus lixianensis
Age: late Early Cretaceous (Tang et al.
2001a)
48.14.
NINGXIA HUIZU ZIZHIQU
,
PEOPLE
’
S REPUBLIC OF CHINA
1. Unnamed unit (Dong 1987; Russell and
Zhao 1996)
Ceratopsia
Psittacosauridae
Psittacosaurus sp.
Age: ?Valanginian-?Albian (Russell and
Zhao 1996)
2. Unnamed unit (Young 1979)
Dinosaur eggs
Age: ?Early Cretaceous (Young 1979)
48.15.
SHANXI
,
PEOPLE
’
S REPUBLIC
OF CHINA
1. Unnamed unit (Young 1958b)
Theropoda
Allosauroidea indet.
Sauropoda indet.
Age: Early Cretaceous (Mateer, pers.
comm.)
48.16.
GUANGXI ZHUANGZU ZIZHIQU
,
PEOPLE
’
S REPUBLIC OF CHINA
1. Napan Formation (Hou et al. 1975; Dong
1980, 1993c)
Theropoda indet. (=Prodeinodon
kwangshiensis)
Sauropoda indet. (=Asiatosaurus
kwangshiensis)
Ornithopoda indet.
Age: ?Aptian-Albian (?equivalent to
Oshih Formation; Early
Cretaceous, Hou et al. 1975)
48.17.
BEIJING MUNICIPAL AREA
,
PEOPLE
’
S REPUBLIC OF CHINA
1. Lushangfen Formation (Tian 1994)
?Sauropoda indet.
Age: Early Cretaceous (Tian 1994)
49.1.
P
’
YONGAN PUKDO
,
NORTH KOREA
1. Sinuiju Series (Lee et al. 2001)
Theropoda
?Avialae indet. (=Proornis coreae)
Age: Early Cretaceous (Lee et al. 2001)
50.1.
KYONGSANG PUKDO
,
SOUTH KOREA
1. Chilgog Formation (Kim 1983)
Sauropoda indet.
Age: Berriasian-Hauterivian (Choi 1986)
2. Guyedong Formation (=Dogyedong
Formation; Kim 1983, 1986; Lee et al.
2001)
Theropoda indet. (=?Deinonychus sp.)
Sauropoda indet. (=Ultrasaurus
tabriensis)
Ornithopoda
?Euornithopoda indet.
Age: Aptian-Albian (Choi 1985)
3. Jeomgog Formation (Lee et al. 2001)
Theropod tracks (including avialan
tracks)
Age: Aptian-Albian (Lee et al. 2001)
4. Geoncheonri Formation (Lee et al. 2001)
Theropod tracks
Euornithopod tracks
Age: Albian (Lee et al. 2001)
5. Sagog Formation (Lee et al. 2001)
Theropod tracks
Sauropod tracks
Euornithopod tracks
Age: Albian (Lee et al. 2001)
50.2.
KYONGSANG NAMDO
,
SOUTH KOREA
1. Iljig Formation (Kim 1983)
?Sauropoda indet.
Age: Berriasian-Hauterivian (Choi 1986)
2. Jinju Formation (Lee et al. 2001; Lim
et al. 2001)
Theropod tracks
Sauropoda
Titanosauriformes indet.
(=Brachiosauridae indet.)
Euornithopod tracks
Age: Barremian (Lee et al. 2001)
3. Hasandong Formation (Kim 1983; Lee
et al. 2000, 2001; Paik 2000; Park et al.
2000; Dong et al. 2001; Barrett et al.
2002b)
Theropoda indet.
Theropod tracks
Sauropoda
Sauropoda indet. (=Chiayusaurus
asianensis, Pukyongosaurus
millenniumi)
Camarasauridae indet.
Titanosauridae indet.
Sauropoda indet.
Sauropod tracks
Euornithopod tracks
Dinosaur eggs
Age: Aptian-Albian (Choi 1986)
4. Nagdong Formation (Kim 1983)
Sauropoda indet.
Age: Aptian-Albian (Choi 1986)
568 DINOSAUR DISTRIBUTION AND BIOLOGY

5. Haman Formation (Kim 1983; Lockley
et al. 1992c; Lim et al. 1995, 2000; Lee
et al. 2001)
Theropod tracks (including avialan
tracks)
Sauropod tracks
Euornithopod tracks
Age: Aptian-Albian (Choi 1986)
6. Jindong Formation (Kim 1983; Lim et al.
1989,1994; Lockley et al. 1992c; Lee et al.
2000, 2001; Paik et al. 2001)
Theropod tracks (including avialan
tracks)
Sauropod tracks
Euornithopod tracks
Dinosaur eggs
Age: Albian (Choi 1985, 1986; Lee et al.
2001)
7. Unnamed unit (Zelenitsky, pers.
comm.)
Dinosaur eggs
Age: Early Cretaceous (Zelenitsky,
pers. comm.)
50.3.
CH
ˇ
ULLA NAMDO
,
SOUTH KOREA
1. Unnamed unit (Lee et al. 2001)
Dinosaur eggs
Age: Early Cretaceous (Lee et al. 2001)
51.1.
CHANGWAT KHON KAEN
,
THAILAND
1. Sao Khua Formation (Ingavat and
Taquet 1978; Ingavat et al. 1978;
Buffetaut 1982; Buffetaut and Ingavat
1983, 1984, 1986a; Buffetaut et al. 1996a,
1997b; Buffetaut and Suteethorn 1998b;
Martin et al. 1999)
Theropoda
Carnosauria
Siamotyrannus isanensis
Ornithomimosauria indet.
Theropoda indet. (including
Siamosaurus suteethorni)
Sauropoda
Titanosauria
Phuwiangosaurus sirindhornae
Sauropoda indet.
Age: Hauterivian-Valanginian
(Buffetaut and Suteethorn
1998b)
51.2.
CHANGWAT KALASIN
,
THAILAND
1. Phra Wihan Formation (Buffetaut et al.
1997b, 2002; Le Loeuff et al. 1999, 2002)
Theropod tracks
Sauropod tracks
Euornithopod tracks
Age: Berriasian (Buffetaut and
Suteethorn 1998b)
2. Sao Khua Formation (Buffetaut 1982;
Suteethorn et al. 1995a, 1995b;
Buffetaut et al. 1997b, 2002)
Sauropoda
Titanosauria
Phuwiangosaurus sirindhornae
Sauropoda indet.
Age: Hauterivian-Valanginian
(Buffetaut and Suteethorn
1998b)
51.4.
CHANGWAT UDON THANI
,
THAILAND
1. Sao Khua Formation (Buffetaut 1982;
Buffetaut and Ingavat 1986b; Buffetaut
and Suteethorn 1998b; Buffetaut et al.
2002)
Sauropoda indet.
Age: Hauterivian-Valanginian (Buffetaut
and Suteethorn 1998b)
51.5.
CHANGWAT NAKHON RATCHASIMA
,
THAILAND
1. Khok Kruat Formation (Buffetaut 1982;
Buffetaut and Ingavat 1986a; Buffetaut
and Suteethorn 1998b)
Theropoda indet.
Age: ?Aptian-Albian (Buffetaut and
Suteethorn 1998b)
51.6.
CHANGWAT UBON RATCHATHANI
,
THAILAND
1. Khok Kruat Formation (Buffetaut et al.
1997b, 2002; Buffetaut and Suteethorn
1998a, 1998b)
Sauropoda
Nemegtosauridae indet.
Ornithopoda
Iguanodontia indet.
Age: ?Aptian-Albian (Buffetaut and
Suteethorn 1998b)
51.7.
CHANGWAT CHAIYAPHUM
,
THAILAND
1. Khok Kruat Formation (Buffetaut and
Suteethorn 1992, 1998b)
Ceratopsia
Ceratopsia indet. (=Psittacosaurus
sattayaraki)
Age: ?Aptian-Albian (Buffetaut and
Suteethorn 1998b)
51.8.
CHANGWAT LOEI
,
THAILAND
1. Phu Phan Formation (Buffetaut et al.
1985a, 2002; Buffetaut and Suteethorn
1998b)
Theropod tracks
Sauropoda indet.
Age: Barremian-Aptian (Buffetaut and
Suteethorn 1998b)
52.1.
SAVANNAKHET KHOUENG
,
LAOS
1. “Grès supérieurs” formation (Hoffet
1942, 1943a, 1943b; Buffetaut 1991;
Allain et al. 1997, 1999; Buffetaut and
Suteethorn 1998b; equivalent to the
Khok Kruat Formation of Thailand)
Theropod tracks
Sauropoda
Titanosauria
Tangvayosaurus hoffeti
Lithostrotia
Titanosaurus falloti
Sauropod tracks
Ornithopoda
Iguanodontia indet.
(=Mandschurosaurus
laosensis)
Euornithopod tracks
Age: Aptian-Albian (Buffetaut and
Suteethorn 1998b)
53.2.
GIFU PREFECTURE
,
JAPAN
1. Amagodani Formation (Hasegawa et al.
1995)
Ornithopoda
Iguanodontia indet.
Age: ?Berriasian (=early Neocomian;
Hasegawa et al. 1995)
2. Okurodani Formation (Matsukawa and
Obata 1994; Azuma and Tomida 1995;
Hasegawa et al. 1995; Evans et al. 1998)
Theropoda
Dromaeosauridae indet.
Avialae indet.
Theropoda indet.
Ornithopoda
Euornithopoda
Euornithopoda indet.
Iguanodontia indet.
Ornithopoda indet.
Age: Berriasian-Valanginian (Matsukawa
and Obata 1994)
3. Unnamed unit (Kunimitsu et al. 1990)
Euornithopod tracks
Age: Valanginian (Azuma and Tomida
1997)
53.4.
KUMAMOTO PREFECTURE
,
JAPAN
1. Mihune Formation (Mateer,
pers. comm.)
Theropoda indet.
Age: ?Albian (Mateer, pers. comm.)
53.5.
MIE PREFECTURE
,
JAPAN
1. Matsuo Group (Tanimoto 1998, 1999;
Tomida et al. 2001)
Sauropoda
Lithostrotia
Undescribed lithostrotian
Lithostrotia indet.
Ornithopoda tracks
Age: Berriasian-Hauterivian (Sugiyama
et al. 1993)
53.6.
TOKUSHIMA PREFECTURE
,
JAPAN
1. Tazukawa Formation (Morozumi et al.
1995; Azuma and Tomida 1997)
Ornithopoda
Iguanodontia indet.
Age: late Hauterivian–Barremian
(Azuma and Tomida 1997)
53.7.
ISHIKAWA PREFECTURE
,
JAPAN
1. Kuwajima Formation (Matsukawa and
Obata 1994; Azuma and Tomida 1995;
Hasegawa et al. 1995; Barrett and
Manabe 2000; Manabe et al. 2000;
Barrett et al. 2002b; Manabe, pers.
comm.)
Theropoda
?Oviraptorosauria indet.
Theropoda indet.
Theropod tracks
Sauropoda
Titanosauriformes indet.
Ornithischia indet.
Ornithopoda.
Euornithopoda indet.
Iguanodontia indet.
DINOSAUR DISTRIBUTION 569

Age: Berriasian-Hauterivian (Manabe
et al. 2000; Barrett et al.
2002b)
2. Akaiwa Formation (Kondo et al. 1990;
Matsukawa and Obata 1994; Azuma
and Tomida 1995)
?Theropod tracks
?Euornithopod tracks
Age: Hauterivian-Aptian (Azuma and
Tomida 1995)
53.8.
FUKUI PREFECTURE
,
JAPAN
1. Izuki Formation (Matsukawa and Obata
1994; Azuma and Tomida 1995; Manabe
1999; Azuma et al. 2002)
Theropoda
Tyrannosauridae indet.
Avialan tracks
?Theropod tracks
Ornithischian tracks
Euornithopod tracks
Age: Berriasian (Azuma and Tomida
1995)
2. Akaiwa Formation (Kondo et al. 1990;
Matsukawa and Obata 1994)
?Theropod tracks
?Euornithopod tracks
Age: Hauterivian-Aptian (Azuma and
Tomida 1995)
3. Kitadani Formation (Azuma 1989, 1990,
1991, 2003; Azuma et al. 1990a, 1990b,
1991; Dong et al. 1990; Matsukawa and
Obata 1994; Azuma and Currie 1995,
2000; Azuma and Tomida 1995;
Kobayashi and Azuma 1999; Kobayashi
and Azuma 2003)
Theropoda
Carnosauria
Fukuiraptor kitadaniensis
Dromaeosauridae indet.
Theropoda indet.
Theropod tracks (including avialan
tracks)
Sauropoda
Titanosauriformes indet.
Sauropoda indet.
Sauropod tracks
Ornithischia tracks
Ornithopoda
Iguanodontia
Fukuisaurus tetoriensis
Iguanodontia indet.
Euornithopod tracks
Ceratopsia
Psittacosauridae
Psittacosaurus sp.
Dinosaur eggs
Age: Aptian-Albian (Azuma and Tomida
1995)
53.9.
GUMMA PREFECTURE
,
JAPAN
1. Sebayashi Formation (Hasegawa et al.
1999; Matsukawa and Obata 1985, 1989;
Manabe et al. 1989; Azuma and Tomida
1997)
Theropoda
Ornithomimidae indet.
Theropoda indet.
?Theropod tracks
Euornithopod tracks
Age: late Barremian–Aptian (Matsukawa
and Obata 1994)
53.10.
IWATE PREFECTURE
,
JAPAN
1. Tanohata Formation (Hasegawa et al.
1991; Azuma and Tomida 1997)
Sauropoda indet. (including
?Mamenchisaurus sp.)
Age: late Aptian (Matsukawa and Obata
1994)
53.11.
FUKUOKA PREFECTURE
,
JAPAN
1. Sengoku Formation (Okazaki 1990, 1992a)
Theropoda indet. (=Wakinosaurus satoi)
Age: late Hauterivian (Matsukawa and
Obata 1991)
2. Unnamed unit (Okazaki 1992b)
Theropoda indet.
Sauropoda indet.
?Ceratopsian indet.
Age: Berriasian-Hauterivian
(=Neocomian; Azuma and
Tomida 1997)
53.12.
TOYAMA PREFECTURE
,
JAPAN
1. Nagatogawa Formation (Goto 1992;
Azuma and Tomida 1997; Matsukawa
et al. 1997)
Theropod or euornithopod tracks
Age: Early Cretaceous (Matsukawa et al.
1997); Valanginian (Azuma
and Tomida 1997)
53.13.
YAMAGUCHI PREFECTURE
,
JAPAN
1. Kiyosu-e Formation (Okazaki 1994;
Azuma and Tomida 1997)
Euornithopod tracks
Age: earliest Cretaceous (Azuma and
Tomida 1997)
570 DINOSAUR DISTRIBUTION AND BIOLOGY
South America
Seventeen South American dinosaur locations are currently
known from Lower Cretaceous strata (an increase of 40%; figs.
24.8 and 24.9). Most include skeletal remains from Brazil and
Argentina, described by J. F. Bonaparte, J. O. Calvo, R. A. Coria,
A. W. A. Kellner, D. M. Martill, D. Rivarola, and L. Salgado, among
others. The remaining Early Cretaceous dinosaur occurrences
consist of Brazilian ichnofaunas investigated by G. Leonardi.
54.2.
DEPARTAMENTO DE SANTANDER
,
COLOMBIA
1. Unnamed unit (Buffetaut 2000)
Theropod tracks
Age: ?Early Cretaceous (Buffetaut 2000)
55.1.
PROVINCE OF LOJA
,
ECUADOR
1. Cazaderos Formation (Leonardi 1994)
Theropod tracks
Age: Albian-Cenomanian (Leonardi
1994)
57.3.
ESTADO DO PARANÁ
,
BRAZIL
1. Caiua Formation (Leonardi 1989)
Theropod tracks
Age: Early Cretaceous (Leonardi 1989)
57.5.
ESTADO DO GOIÁS
,
BRAZIL
1. Corda Formation (Leonardi 1980b)
Sauropod tracks
Euornithopod tracks
Age: pre-Aptian (Leonardi 1989)
57.6.
ESTADO DO BAHIA
,
BRAZIL
1. Bahia Series (Mawson and Woodward
1907 )
Theropoda indet.
Ornithopoda
Iguanodontia indet.
Age: Aptian (Taquet 1977)
57.7.
ESTADO DO CEARÁ
,
BRAZIL
1. Antenor Navarro Formation (Leonardi
1989)
Theropod tracks
Euornithopod tracks
Age: pre-Aptian (Leonardi 1989)
2. Santana Formation (Leonardi and
Borgomanero 1981; Campos 1985;
Campos and Kellner 1991; Frey and
Martill 1995; Kellner 1996a, 1996b,
1999; Kellner and Campos 1996; Martill
et al. 1996a)
Dinosauria indet.
Theropoda
Spinosauridae
Irritator challengeri
Angaturama limai
Unnamed compsognathid
?Tyrannosauroidea
Santanaraptor placidus
?Oviraptorosauria indet.
Avialae indet.
Theropoda indet.
?Ornithischia indet.
Age: ?Albian (Frey and Martill 1995)

57.8.
ESTADO DO PARAÍBA
,
BRAZIL
1. Antenor Navarro Formation (Leonardi
1981b)
Theropod tracks
Sauropod tracks
Euornithopod tracks
Age: pre-Aptian (Leonardi 1989)
2. Sousa Formation (Leonardi 1979b)
Theropod tracks
Sauropod tracks
?Ankylosaur tracks
Euornithopod tracks
Age: pre-Aptian (Leonardi 1989)
3. Piranhas Formation (Leonardi 1981b)
Theropod tracks
Euornithopod tracks
Age: pre-Aptian (Leonardi 1989)
62.3.
PROVINCIA DE COLCHAGUA
,
CHILE
1. Banos del Flaco Formation (Casamiquela
and Fasola 1968)
Euornithopod tracks
Age: Hauterivian (Diaz and Gasparini
1976)
63.4.
PROVINCIA DE CHUBUT
,
ARGENTINA
1. Cerro Barcino Formation (Corro 1966,
1975; Bonaparte and Gasparini 1978;
Salgado 1993; Bonaparte 1996a, 1996b;
Rich et al. 1999b; Vickers-Rich et al.
1999a; Rauhut et al. 2003)
Theropoda
Abelisauridae indet.
?Carcharodontosauridae indet.
Theropoda indet. (Megalosaurus
inexpectatus)
Sauropoda
Undescribed titanosauriform
Titanosauria
Chubutisaurus insignis
Titanosauria indet.
Age: Albian (Bonaparte 1996b)
63.5.
PROVINCIA DE SAN LUIS
,
ARGENTINA
1. La Cantera Formation (Lull 1942; not
Paganzo Beds III, Bonaparte, pers. comm.)
Theropod tracks
Age: Berriasian-Hauterivian
(=Neocomian; Leonardi
1989)
2. El Jume Formation. (Rivarola 2000)
Dinosaur tracks
Age: Early Cretaceous (Rivarola 2000)
63.6.
PROVINCIA DE NEUQUÉN
,
ARGENTINA
1. La Amarga Formation (Salgado and
Bonaparte 1991; Bonapar te 1996a, 1996b)
Theropoda
Ceratosauria
Ligabueino andesi
Sauropoda
Diplodocoidea
Amargasaurus cazaui
Stegosauria
?Stegosauridae indet.
Age: Barremian (Leanza and Hugo
2001); Berriasian-
Valanginian (=late
Neocomian; Salgado and
Bonaparte 1991; Bonapar te
1996b)
2. Lohan Cura Formation (Bonaparte
1999b)
Sauropoda
Titanosauria
Agustinia ligabuei
Age: late Aptian–early Albian (Leanza
and Hugo 2001); Aptian
(Bonaparte 1999b)
3. Rayoso Formation (Bonaparte 1996b)
Theropoda
Ceratosauria indet.
Sauropoda
Diplodocoidea
Rayososaurus agrioensis
Age: Aptian (Bonaparte 1996b)
DINOSAUR DISTRIBUTION 571
Africa
The African record of the Lower Cretaceous (24 locations, a
60% increase) has yielded a variety of dinosaur fossils, primarily
through the work of C. Deperét and A. F. de Lapparent, R. Lavo-
cat, M. Monbaron, J. Savornin, P. Taquet, and in the “Conti-
nental intercalaire” and Elrhaz Formation of Morocco, Algeria,
Tunisia, Libya, Mali, and Niger, as well as the Kalahari Deposits
of South Africa. Additional Late Jurassic to Early Cretaceous
African material is known from South Africa, originally studied
by R. Broom and later by W. A. Coombs Jr., P. M. Galton, R. E.
Molnar, P. V. Rich, and T. H. V. Rich. Other recent research on
Afrcan faunas include studies by S. Bouaziz, M. Brunet, J. Con-
gleton, J. J. Flynn, C. A. Forster, E. M. P. Gomani, L. L. Jacobs,
T. H. V. Rich, and P. C. Sereno. Figures 24.10, 24.11, and 24.23
indicate the positions of these locations.
64.6.
PROVINCE DE KSAR
-
ES
-
SOUK
,
MOROCCO
1. Tegana Formation (Lavocat 1954;
Monbaron 1978; Jacobs et al. 1996)
Theropoda
Allosauroidea
Carcharodontosaurus saharicus
Theropoda indet. (=Elaphrosaurus sp.)
Sauropoda
Diplodocoidea
Rebbachisaurus garasbae
Age: Albian (Lapparent 1960a)
65.3.
WILAYA ADRAR
,
ALGERIA
1. “Continental intercalaire” (Depéret
and Savornin 1925; Lapparent 1960a;
Bassoulet and Iliou 1967; Broin et al.
1971; Taquet 1977)
Theropoda
Allosauroidea
Carcharodontosaurus saharicus
Undescribed theropods (two species)
Theropoda indet. (=Elaphrosaurus
iguidiensis)
Sauropoda
Sauropoda indet.
Undescribed sauropod
Diplodocoidea
Nigerosaurus taqueti (including
Rebbachisaurus tamesnensis)
Age: Albian–early Cenomanian
(Lapparent 1960a; Petters
1981; Mateer et al. 1992)
65.4.
WILAYA TAMENGHEST
,
ALGERIA
1. “Continental intercalaire” (Lapparent
1960a)
Theropoda
Allosauroidea
Carcharodontosaurus saharicus
Sauropoda
Diplodocoidea
Nigerosaurus taqueti (including
Rebbachisaurus tamesnensis)
Age: Albian–early Cenomanian
(Lapparent 1960a; Petters
1981; Mateer et al. 1992)
65.5.
WILAYA WARGLA
,
ALGERIA
1. “Continental intercalaire” (Lapparent
1960a)
Theropoda
Allosauroidea
Carcharodontosaurus saharicus
Theropoda indet. (=Elaphrosaurus
iguidiensis)
Sauropoda
Diplodocoidea
Nigerosaurus taqueti (including
Rebbachisaurus tamesnensis)
Brachiosauridae
Brachiosaurus nougaredi
Age: Albian–early Cenomanian
(Lapparent 1960a; Petters
1981; Mateer et al. 1992)
66.1.
GOUVERNORAT DE MEDENINE
,
TUNISIA
1. “Continental intercalaire” (Lapparent
1960a; Schlüter and Schwarzhans 1978)
Theropoda
Allosauroidea
Carcharodontosaurus saharicus
cf. Carcharodontosaurus sp.
Theropoda indet. (=Elaphrosaurus
gautieri)
Sauropoda
Sauropoda indet.

Diplodocoidea
Nigerosaurus taqueti (including
Rebbachisaurus tamesnensis)
Ornithopoda
Iguanodontia indet.
Age: Albian–early Cenomanian
(Lapparent 1960a; Petters
1981; Mateer et al. 1992)
2. Chenini Formation (Bouaziz et al. 1988;
Jacobs et al. 1996; Benton et al. 2000a)
Theropoda
Allosauroidea
Carcharodontosaurus saharicus
?Spinosauridae
?Spinosaurus sp.
Sauropoda indet.
Ornithopoda
Iguanodontia indet.
Age: late Aptian–early Albian (Le Loeuff
et al. 2000)
66.2.
GOUVERNORAT DE TATAOUINE
,
TUNISIA
1. Douiret Formation (Le Loeuff et al. 2000)
Ornithopoda
Iguanodontia indet.
Age: early Aptian (Le Loeuff et al. 2000)
2. Chenini Formation (Le Loeuff et al.
2000; Buffetaut and Ouaja 2002)
Theropoda
Spinosauridae indet.
Theropoda indet.
Sauropoda indet.
Ornithopoda
Iguanodontia indet.
Age: late Aptian–early Albian (Le Loeuff
et al. 2000)
3. Oum ed Diab Formation (Le Loeuff et al.
2000)
Ornithopoda
Iguanodontia indet.
Age: late Albian (Le Loeuff et al. 2000)
67.1.
REGION DE GAO
,
MALI
1. “Continental intercalaire” (Lapparent
1960a; O’Leary 1999)
Sauropoda indet.
Ankylosauria
Nodosauridae indet.
Age: Albian–early Cenomanian
(Lapparent 1960a; Petters
1981; Mateer et al. 1992)
68.1.
DÉPARTEMENT D
’
AGADEZ
,
NIGER
1. Elrhaz Formation (Lapparent 1960a;
Taquet 1976; Galton and Taquet 1982;
Sereno et al. 1998, 1999; Taquet and
Russell 1998, 1999; Wilson et al. 1999b;
Sereno and Wilson 2001; Wilson, pers.
comm.)
Theropoda
Spinosauridae
Suchomimus tenerensis
Cristatusaurus lapparenti
?Spinosauridae indet.
Tetanurae indet.
Theropoda indet. (=Elaphrosaurus
iguidiensis)
Sauropoda
Diplodocoidea
Nigersaurus taqueti
Undescribed titanosaurian
Sauropoda indet.
Ornithopoda
Iguanodontia
Valdosaurus nigeriensis
Ouranosaurus nigeriensis
Lurdusaurus arenatus
Age: late Aptian (Taquet 1976; Moody
and Sutcliffe 1991)
2. “Continental intercalaire” (Lapparent
1960a)
Theropoda
Avetheropoda incertae sedis
Bahariasaurus ingens
Allosauroidea
Carcharodontosaurus saharicus
Theropoda indet. (=Inosaurus
tedreftensis, Elaphrosaurus
iguidiensis, E. gautieri)
Sauropoda
Sauropoda indet. (=Astrodon sp.,
Rebbachisaurus tamesnensis,
Aegyptosaurus baharijensis)
Ankylosauria
Nodosauridae indet.
572 DINOSAUR DISTRIBUTION AND BIOLOGY
FIGU RE 24.23. Late Jurassic and Early Cretaceous localities in Israel (31), Syria (32), Jordan (33), Sultanate of Oman (34),
Republic of Yemen (35), Egypt (69), and Sudan (71). Scale =600 km.

Age: Albian–early Cenomanian
(Lapparent 1960a; Petters
1981; Mateer et al. 1992)
3. Tiourarén Formation (Sereno et al. 1994;
Jacobs et al. 1996; Sadleir, pers. comm.;
Wilson, pers. comm.)
Theropoda
Tetanurae
Afrovenator abakensis
Sauropoda
Macronaria
Jobaria tiguidensis
Age: Hauterivian-Barremian (Sereno
et al. 1994)
68.2.
DÉPARTEMENT DE LA TAHOUA
,
NIGER
1. Farak Formation (Greigert et al. 1954;
Lapparent 1960a; Taquet 1976; Jacobs
et al. 1996)
Theropoda
Avetheropoda incertae sedis
Bahariasaurus ingens
Sauropoda
Macronaria
Jobaria tiguidensis
Lithostrotia indet. (=Aegyptosaurus
baharijensis)
Age: ?late Albian–early Cenomanian
(Taquet 1976; Petters 1981;
Lefranc and Guiraud 1990)
70.1.
MUHAFAZAT GHARYAN
,
LIBYA
1. “Continental intercalaire” (Lapparent
1960a)
Theropoda
Allosauroidea
Carcharodontosaurus saharicus
Theropoda indet. (=Elaphrosaurus
iguidiensis)
Age: Albian–early Cenomanian
(Lapparent 1960a; Petters
1981; Mateer et al. 1992)
73.1.
PROVINCE DE NORD
,
CAMEROUN
1. Koum Formation (Flynn et al. 1987;
Congleton 1990; Colin et al. 1992;
Congleton et al. 1992; Jacobs et al. 1989,
1996)
Theropoda
Spinosauridae
Spinosaurus sp.
?Spinosauridae indet.
Theropoda indet.
Theropod tracks
Sauropoda indet.
Sauropod tracks
?Thyreophora indet.
Ornithopoda
Iguanodontia
Ouranosaurus sp.
Euornithopod tracks
Age: Barremian-Aptian (Colin et al.
1992)
76.1.
TETE PROVINCE
,
MOZAMBIQUE
1. Lupata Series (Dixey and Smith
1929)
Sauropoda indet.
Age: Aptian (Dixey and Smith 1929;
Pentel’kov and Voronovsky
1979)
78.1.
NORTHERN PROVINCE
,
ZAMBIA
1. Dinosaur Beds (Dixey 1937; Colin and
Jacobs 1990; Jacobs et al. 1996)
Sauropoda indet.
Age: Aptian (Colin and Jacobs 1990;
Pentel’kov and Voronovsky
1979)
79.1.
NORTHERN PROVINCE
,
MALAWI
1. Dinosaur Beds (Dixey 1928; Haughton
1928; Colin and Jacobs 1990; Jacobs et
al. 1990, 1993, 1994, 1996; Gomani
1998, 1999; Gomani et al. 1999)
Undescribed theropod
Sauropoda
Lithostrotia
Malawisaurus dixeyi
Undescribed sauropods (two species)
Age: Aptian (Pentel’kov and Voronovsky
1979; Colin and Jacobs 1990;
Eby et al. 1995)
81.3.
CAPE PROVINCE
,
SOUTH AFRICA
1. Kalahari Deposits (Haughton 1915;
Cooper 1985)
Ornithopoda
Iguanodontia indet. (=Kangnasaurus
coetzeei)
Age: ?Late Jurassic–Valanginian (Dingle
1983)
2. Enon Formation (Mateer 1987)
Theropoda indet.
Age: ?Late Jurassic–Valanginian
(McLachlan and
McMillan 1976; Dingle
et al. 1983)
3. Upper Kirkwood Formation (Broom
1904; Galton and Coombs 1981; Rich
et al. 1983; Forster 1996c; Jacobs et al.
1996; de Klerk et al. 1998, 2000; Forster,
pers. comm.)
Theropoda
Coelurosauria incertae sedis
Nqwebasaurus thwazi
Theropoda indet.
Sauropoda
Sauropoda indet. (=Algoasaurus
bauri, aff. Camarasauridae
indet., Diplodocidae indet.,
aff. Pleurocoelus sp., aff.
Astrodon sp.)
?Ornithischian indet.
Stegosauria
Stegosauridae
Paranthodon africanus
Ornithopoda
Undescribed iguanodontian
?Iguanodontia indet.
Age: ?Late Jurassic–early Valanginian
(McLachlan and McMillan
1976; Dingle1983; Rich et al.
1983)
4. Sundays River Formation (Rich et al.
1983; Mateer 1987)
Theropoda indet.
Age: ?Hauterivian (McLachlan and
McMillan 1976; Dingle
1983)
DINOSAUR DISTRIBUTION 573
Australasia
Since the late 1970s, Australasia has claimed nearly all major di-
nosaur groups known to exist during this time interval elsewhere
in the world. This information on Australasian dinosaurs (fig.
24.12) is the result of the efforts of A. Bartholomai, J. A. Long,
R. E. Molnar, T. H. Rich, and R. A. Thulborn, and P. Vickers-Rich.
84.1.
QUEENSLAND
,
AUSTRALIA
1. Bungil Formation (Molnar 1980a)
Ankylosauria
Ankylosauridae
Minmi paravertebra
Age: Aptian (Exon and Vine 1970)
2. Unnamed unit (possibly Toolebuc or
Allaru Formation; Molnar 1982; Long
1998)
Sauropoda indet.
Age: Albian (Molnar 1980b)
3. Toolebuc Formation (Molnar 1980b,
1984, 1986; Kurochkin and Molnar
1997; Rich and Vickers-Rich 1997;
Long 1998)
Theropoda
Avialae
Enantiornithes
Nanantius eos
Nanantius sp.
Ornithischian indet.
Ankylosauria
Ankylosauridae
Minmi sp.
?Ankylosauria indet.
Ornithopoda
Iguanodontia
Muttaburrasaurus sp.
Ornithopoda indet.
Age: Albian (Molnar 1980b)

4. Mackunda Formation (Bartholomai and
Molnar 1981)
Ornithopoda
Iguanodontia
Muttaburrasaurus langdoni
Age: Albian (Vine and Day 1965)
5. Allaru Formation (Longman 1933;
Molnar 1980b, 1996; Rich and Vickers-
Rich 1997)
Sauropoda
Titanosauria
Austrosaurus mckillopi
Ankylosauria
Ankylosauridae
Minmi sp.
?Ankylosauria indet.
Ornithopoda
Iguanodontia
?Muttaburrasaurus sp.
Ornithopoda indet.
Age: Albian (Molnar 1982)
84.2.
WESTERN AUSTRALIA
,
AUSTRALIA
1. Broome Sandstone (Colbert and
Merrilees 1967; Long 1990; Thulborn
et al. 1994; Rich and Vickers-Rich 1997)
Theropod tracks
Sauropod tracks
?Stegosaur tracks
Euornithopod tracks
Age: Berriasian-Hauterivian
(=Neocomian; Colbert
and Merrilees 1967)
2. Birdsong Sandstone (Long 1998)
Theropoda indet.
Age: Hauterivian-Barremian (Long
1998)
84.3.
SOUTH AUSTRALIA
,
AUSTRALIA
1. Maree Formation (Molnar and Pledge
1980; Long 1998)
Theropoda
Coelurosauria incertae sedis
Kakuru kujani
Theropoda indet.
Age: Aptian (Carr et al. 1979)
2. ?Cadn-Owie Formation (Molnar 1980b)
Ornithopoda indet.
Age: ?Aptian (Molnar 1980b)
84.4.
NORTHERN TERRITORY
,
AUSTRALIA
1. Unnamed unit (Molnar, pers. comm.)
Ornithopoda
?Iguanodontia indet.
Age: Early Cretaceous (Molnar,
pers. comm.)
84.5.
NEW SOUTH WALES
,
AUSTRALIA
1. Griman Creek Formation (Huene 1932;
Molnar 1980b, 1996, 1999; Molnar and
Galton 1986; Rich and Vickers-Rich
1997)
Theropoda
Tetanurae incertae sedis
Rapator ornitholestoides
Theropoda indet. (=Walgettosuchus
woodwardi, Avialae indet.)
Sauropoda indet. (=Brachiosauridae
indet. )
Ornithopoda
Euornithopoda
Fulgurotherium australe partim
Euornithopoda indet.
Iguanodontia incertae sedis
Muttaburrasaurus sp.
Euornithopod tracks
Age: Albian (Molnar 1980b)
2. Coreena Formation (Molnar 1980b)
Theropoda indet.
Age: Albian (Molnar 1980b)
84.6.
VICTORIA
,
AUSTRALIA
1. Wonthaggi Formation (Woodward
1906; Molnar et al. 1981, 1985; Rich
and Rich 1989; Rich and Vickers-Rich
1994, 1997, 2003; Currie et al. 1996;
Vickers-Rich and Rich 1997; Chure
1998b)
Theropoda
?Allosauroidea indet.
Ornithomimosauria indet.
?Dromaeosauridae indet.
Theropoda indet.
Ankylosauria indet.
Ornithopoda
Euornithopoda
Fulgurotherium australe
Qantassaurus intrepidus
Fulgurotherium sp.
Euornithopoda indet.
?Ceratopsia
?Neoceratopsia indet.
(=Serendipaceratops
arthurclarkei)
Age: Valanginian-Aptian (Rich and Rich
1989)
2. Eumeralla Formation (Rich and Rich
1989; Rich and Vickers-Rich 1994, 1997;
Currie et al. 1996; Vickers-Rich and Rich
1997; Long 1998)
Theropoda
Ornithomimosauria indet. (=Timimus
hermani)
?Oviraptorosauria indet.
?Dromaeosauridae indet.
Theropoda indet.
Ornithischian tracks
Ornithopoda
Euornithopoda
Leaellynasaura amicagraphica
Atlascoposaurus loadsi
Euornithopoda indet.
Dinosaur tracks
Age: early Albian (Rich and Rich 1989;
Vickers-Rich and Rich
1997)
574 DINOSAUR DISTRIBUTION AND BIOLOGY
Late Cretaceous
Late Cretaceous dinosaurs are known from nearly four hundred
locations, particularly from the Campanian and Maastrichtian
and principally from Canada and the United States in North
America and from Mongolia and the People’s Republic of China
in eastern Asia. Additional Late Cretaceous material is known
from Europe, South America, Africa, Australasia, and Antarctica.
North America
The greatest diversity and abundance of dinosaurs are known
from the Late Cretaceous of North America (148 locations, an
increase of nearly 50%; figs. 24.1, 24.2, 24.3, 24.14, 24.17, 24.18,
and 24.24). Histories of the great Late Cretaceous dinosaur dis-
coveries of the late 1800s and early 1900s exist in the writings of
Sternberg (1909, 1917), Russell (1967), and Colbert (1968). The
likes of B. Brown, E. D. Cope, W. E. Cutler, G. M. Dawson, C. W.
Gilmore, J. B. Hatcher, L. M. Lambe, J. Leidy, O. C. Marsh, W. A.
Parks, and C. H. Sternberg and his sons (George, Charles M., and
Levi), and established the dinosaurian wealth from North Amer-
ica. At a pace unabated to this day, Late Cretaceous dinosaurs
from the Western Interior of Canada, particularly southeastern
Alberta, continue to be revealed through the important research
of D. B. Brinkman, P. J. Currie, D. A. Eberth, R. C. Fox, W. Lang-
ston Jr., D. A. Russell, L. S. Russell, J. E. Storer, D. Tanke, and
T. To ka ry k.
At the same time, a great array of dinosaur paleontologists—
among them J. D. Archibald, B. H. Breithaupt, K. Carpenter,
W. A. Clemens, P. Dodson J. G. Eaton, D. E. Fastovsky, A. R. Fio-
rillo, D. Gabriel, R. A. Gangloff, J. R. Horner, A. P. Hunt, J. K.

Kirkland, T. M. Lehman, S. G. Lucas, M. C. McKenna, J. K. Rigby
Jr., P. M. Sheehan, R. E. Sloan, R. M. Sullivan, L. Van Valen, D. B.
Weishampel, T. E. Williamson, and D. G. Wolfe—carry on the
tradition first established by Cope and Marsh in the northern
realm of the Western Interior and Alaska of the United States.
Elsewhere in the United States, Upper Cretaceous exposures
of the San Juan Basin of New Mexico, of the Big Bend area of
Texas, of the states of the Gulf Coast, and along the eastern sea-
board of the United States have also yielded important dinosaur
material. These areas were initially studied by B. Brown, C. W.
Gilmore, J. Leidy, and C. H. Sternberg, and more recently by J. G.
Armstrong-Ziegler, W. A. Clemens, E. H. Colbert, R. K. Denton,
W. B. Gallagher, J. J. Head, J. P. Lamb, W. Langston Jr., Y.-M.
Lee, T. M. Lehman, S. G. Lucas, N. J. Mateer, D. Parris, and D. R.
Schwimmer, among others.
Several small but important Late Cretaceous dinosaur faunas
from Mexico and Honduras have been documented. They have
been studied by R. E. Molnar, W. I. Morris, G. E. Murray, and
others, and most recently by C. Gonzáles-Leon, R. Hernández-
Riviera, and R. Rodriguez de la Rosa.
DINOSAUR DISTRIBUTION 575
1.1.
NORTHWEST TERRITORIES
,
CANADA
1. Summit Creek Formation (Russell
1984c)
Ceratopsia
?Ceratopsidae indet.
Age: late Maastrichtian (Yorath and
Cook 1981)
2. Kanguk Formation (Russell 1988; Rich
et al. 1997)
Theropoda
Avialae
Hesperornis sp.
Theropoda indet.
Ornithopoda
Hadrosauridae indet.
Age: Maastrichtian (Russell 1988; Rich
et al. 1997)
3. Unnamed unit (Russell 1967)
Theropoda
Avialae
Hesperornithiformes
Hesperornis regalis
Age: Maastrichtian (Russell 1967)
1.2.
YUKON TERRITORY
,
CANADA
1. Bonnet Plume Formation (Russell 1984c)
Ornithopoda
FIGU RE 24.24. Late Cretaceous localities in central and western Canada (1) and the northwestern United States (3). Scale =600 km.

Hadrosauridae indet.
Age: Maastrichtian (Rouse and
Srivastava 1972)
2. Tantalus Formation (Storer,
pers. comm.)
Theropod tracks
Ankylosaur tracks
Ornithopod tracks
Age: Late Cretaceous (Storer,
pers. comm.)
1.3.
BRITISH COLUMBIA
,
CANADA
1. Dunvegan Formation (Storer 1975;
McCrea et al. 2001a, 2001b; Scott et al.
2001)
Theropod tracks (including avialan
tracks)
Ankylosaur tracks
Ornithopod tracks
Age: Cenomanian (Storer 1975)
1.4.
ALBERTA
,
CANADA
1. Dunvegan Formation (Currie 1989a;
McCrea et al. 2001a; Scott et al. 2001)
Theropod tracks
Ankylosaur tracks
Ornithopod tracks
Age: Cenomanian (Storer 1975)
2. Cardium Formation (Currie 1989a)
Theropod tracks
Age: Turonian (Currie 1989a)
3. Kaskapau Formation (Fox 1984)
Theropoda
Avialae
Ichthyornithiformes
Ichthyornis sp.
Age: early Turonian (Fox 1984)
4. Belly River Group (Ryan and Currie
1998; Ryan and Russell 2001)
Theropoda
Ornithomimidae indet.
Ornithopoda
Hadrosauridae indet.
Ceratopsia
Neoceratopsia
Montanoceratops sp.
Age: Campanian (Braman et al. 1995)
5. Milk River Formation (Russell 1935,
1964; Currie 1989a; Baszio 1997; Ryan
and Russell 2001)
Theropoda
Coelurosauria incertae sedis
Ricardoestesia gilmorei
Unnamed Ricardoestesia
species
Tyrannosauridae indet.
Ornithomimidae indet.
?Troodontidae indet. (=Paronychodon
lacustris)
Dromaeosauridae
cf. Saurornitholestes langstoni
Ankylosauria
Nodosauridae indet.
(=cf. Palaeoscincus)
Ankylosauridae indet.
Ornithopoda
Hadrosaurinae indet. (including
cf. Kritosaurus sp.)
Euornithopod tracks
Pachycephalosauria
Pachycephalosauridae indet.
Ceratopsia
Neoceratopsia indet.
Ceratopsidae indet.
(=cf. Brachyceratops sp.)
Age: early Campanian (Fox 1978)
6. Foremost Formation (Fox 1974; Ryan
and Russell 2001)
Theropoda
Coelurosauria incertae sedis
Ricardoestesia sp.
Tyrannosauroidea
Aublysodon sp.
Tyrannosauridae indet.
?Troodontidae indet.
(=Paronychodon sp.)
Dromaeosauridae
Dromaeosaurus sp.
Saurornitholestes sp.
Avialae
Hesperornithiformes
Hesperornis cf. regalis
Ankylosauria
Nodosauridae indet.
Ankylosauridae indet.
Ornithopoda
Euornithopoda indet.
Hadrosauridae indet.
Pachycephalosauria
Pachycephalosauridae
Stegoceras sp.
Ceratopsia
Ceratopsidae indet.
Age: middle Campanian (L. S. Russell
1970 )
7. Oldman Formation (equivalent to the
middle part of Two Medicine Formation;
Horner and Currie 1994; Zelenitsky et al.
1996; Eberth 1997; Eberth et al. 2001b;
Ryan and Russell 2001)
Theropoda
Coelurosauria incertae sedis
cf. Ricardoestesia sp.
Tyrannosauroidea
Aublysodon sp.
Daspletosaurus torosus
Daspletosaurus sp.
cf. Gorgosaurus sp.
Tyrannosauridae indet.
Ornithomimidae indet.
Troodontidae
cf. Troodon sp.
?Troodontidae indet.
(=cf. Paronychodon sp.)
Dromaeosauridae
cf. Dromaeosaurus sp.
Saurornitholestes langstoni
Avialae
Apatornithidae
Apatornis sp.
Ankylosauria
Ankylosauridae
Undescribed Euoplocephalus
species
Ankylosauridae indet.
Nodosauridae indet.
Ornithopoda
Euornithopoda
Orodromeus makelai
Euornithopoda indet.
Hadrosauridae
Brachylophosaurus canadensis
Hypacrosaurus stebingeri
Undescribed lambeosaurine
Hadrosauridae indet.
Pachycephalosauria
Pachycephalosauridae
?Stegoceras validum
Pachycephalosauridae indet.
Ceratopsia
Undescribed centrosaurine
Undescribed chasmosaurine
Dinosaur eggs
Age: late Campanian (Eberth and
Hamblin 1993)
8. Dinosaur Park Formation (Brown 1933b;
Russell 1964; Dodson 1971, 1983, 1984;
Currie 1985, 1986, 1987b, 1987c, 1989a;
Brinkman 1986, 1990; Ryan and Currie
1998; Ryan et al. 1998b; Chin et al. 1999,
2003; Holmes et al. 1999; Eberth et al.
2001b; Holmes et al. 2001; Coy, pers.
comm.)
Theropoda
Coelurosauria incertae sedis
Ricardoestesia gilmorei
Ricardoestesia isosceles
Tyrannosauroidea
Gorgosaurus libratus
Daspletosaurus torosus
Tyrannosauroidae indet.
(including Aublysodon
mirandus)
Therizinosauroidea
cf. Erlikosaurus sp.
Ornithomimosauria
Struthiomimus altus
Ornithomimus edmontonensis
(including Struthiomimus
samueli)
Ornithomimidae indet.
Oviraptorosauria
Chirostenotes pergracilis
(=Macrophalangia
canadensis, Caenagnathus
collinsi)
Chirostenotes elegans (including
Caenagnathus sternbergi,
Ornithomimus elegans)
?Chirostenotes sp.
Avimimus sp.
Troodontidae
Troodon formosus
(=Stenonychosaurus inequalis,
Pectinodon bakkeri)
?Troodon sp.
?Troodontidae indet.
(=?Paronychodon lacustris,
cf. Paronychodon sp.)
Dromaeosauridae
Dromaeosaurus albertensis
Dromaeosaurus sp.
Saurornitholestes langstoni
Dromaeosauridae indet.
576 DINOSAUR DISTRIBUTION AND BIOLOGY

Avialae
Baptornithidae
Baptornis sp.
Undescribed ichthyornithiform
Ankylosauria
Nodosauridae
Edmontonia rugosidens
Edmontonia longiceps
Panoplosaurus mirus
?Panoplosaurus sp.
Nodosauridae indet.
(=Palaeoscincus costatus)
Ankylosauridae
Euoplocephalus tutus
Ornithopoda
Euornithopod tracks
Euornithopoda
Orodromeus cf. makelai
Hadrosauridae
Brachylophosaurus canadensis
Gryposaurus notabilis
Gryposaurus incurvimanus
Prosaurolophus maximus
?Maiasaura sp.
Corythosaurus casuarius
Lambeosaurus lambei
Lambeosaurus magnicristatus
Parasaurolophus walkeri
Unnamed lambeosaurine
Hadrosauridae indet. (including
Cionodon stenopsis, Pteropelyx
grallipes, Trachodon altidens,
T. marginatus, T. mirabilis,
T. selwyni)
Pachycephalosauria
Homalocephaloidea
Ornatotholus browni
Pachycephalosauridae
Stegoceras validum
Gravitholus albertae
Pachycephalosaurus sp.
Ceratopsia
Neoceratopsia
Leptoceratops sp.
Ceratopsidae
Anchiceratops ornatus
Chasmosaurus belli (including
C. brevirostris, C. canadensis,
C. kaiseni)
Chasmosaurus russelli
Chasmosaurus irvinensis
Chasmosaurus sp.
Monoclonius crassus (including
M. lowei)
Monoclonius sp.
Centrosaurus apertus (including
C. longirostris, Monoclonius
cutleri, M. dawsoni,
M. flexus)
Centrosaurus sp.
Styracosaurus albertensis (including
Monoclonius nasicornus,
S. parksi)
Ceratopsidae indet. (=?Ceratops sp.)
Dinosaur eggs
Dinosaur coprolites
Age: late Campanian (Dodson 1971;
Fox 1978); middle-late
Campanian (Thomas et al.
1990)
9. Allison Formation (Russell 1949; Currie
1989a, pers. comm.)
Theropoda indet.
Theropod tracks
Ornithopoda
Euornithopoda
Euornithopoda indet. (=Laosaurus
minimus)
Hadrosauridae indet.
Ceratopsia
Neoceratopsia
Montanoceratops sp.
Dinosauria eggs
Age: late Campanian (Russell 1949)
10. Bearpaw Shale Formation (Ryan and
Russell 2001)
Theropoda
Ornithomimidae indet.
Ankylosauria
Nodosauridae
Edmontonia sp.
Ornithopoda
Hadrosauridae
Brachylophosaurus sp.
Prosaurolophus sp.
Hadrosauridae indet.
Pachycephalosauria
Pachycephalosauridae
Stegoceras sp.
Ceratopsia
Ceratopsidae indet.
Age: late Campanian–early
Maastrichtian (Caldwell
1982)
11. Horseshoe Canyon Formation
(Sternberg 1926a; Russell and Chamney
1967; D. A. Russell 1969, 1970b, 1972;
Wall and Galton 1979; Currie 1989a;
Ryan et al. 1998a; Sues 1997; Eberth
et al. 2001b; Makovicky 2001; Ryan
and Russell 2001; Varricchio 2001a)
Theropoda
Coelurosauria incertae sedis
Ricardoestesia gilmorei
cf. Ricardoestesia isosceles
Tyrannosauroidea
Aublysodon sp.
Albertosaurus sarcophagus
Daspletosaurus sp.
Tyrannosauridae indet.
Ornithomimosauria
Ornithomimus edmontonensis
(including Struthiomimus
brevitertius)
Struthiomimus altus
Ornithomimidae indet.
Oviraptorosauria
Chirostenotes pergracilis
Caenagnathus sp.
Troodontidae
cf. Troodon sp.
?Troodontidae indet.
(=Paronychodon sp.)
Dromaeosauridae
Undescribed Dromaeosaurus
species
cf. Dromaeosaurus sp.
Saurornitholestes sp.
Undescribed dromaeosaurid
?Dromaeosauridae indet.
Avialae indet.
Theropod tracks
Ankylosauria
Nodosauridae
Edmontonia longiceps
Panoplosaurus sp.
Ankylosauridae
Euoplocephalus tutus
Ornithopoda
Euornithopoda
Parksosaurus warreni
Thescelosaurus neglectus
Hadrosauridae
Edmontosaurus regalis
Saurolophus osborni
Hypacrosaurus altispinus
Euornithopod tracks
Pachycephalosauria
Pachycephalosauridae
Stegoceras edmontonense
Stegoceras validum (=Prenocephale
brevis)
Ceratopsia
Neoceratopsia
Montanoceratops cerorhynchus
Ceratopsidae
Anchiceratops ornatus (including
A. longirostris)
Arrhinoceratops brachyops
Pachyrhinosaurus canadensis
Ceratopsidae indet.
Ceratopsid tracks
Age: early Maastrichtian (Fox 1978)
12. Wapiti Formation (Tanke 1988; Currie
1989a; Ryan and Russell 2001)
Theropoda
Tyrannosauroidea
Albertosaurus sp.
Tyrannosauridae indet.
Ornithomimidae indet.
Troodontidae
Troodon sp.
Dromaeosauridae
Saurornitholestes sp.
Theropod tracks
Ankylosauria
?Ankylosauridae indet.
Ankylosaur tracks
Ornithopoda
Hadrosauridae indet.
Euornithopod tracks
Ceratopsia
Ceratopsidae
Pachyrhinosaurus sp.
Age: Campanian-Maastrichtian
(Eberth 1997)
13. St. Mary River Formation (Langston
1960b, 1975, 1976; Currie et al. 1991;
Carpenter and Alf 1994)
Theropoda
Tyrannosauroidea
Albertosaurus sp.
Tyrannosauridae indet.
Ornithomimidae indet.
DINOSAUR DISTRIBUTION 577

?Troodontidae
?Troodon sp.
Dromaeosauridae
cf. Saurornitholestes sp.
Theropod indet.
Theropod tracks
Ankylosauria
Nodosauridae
Edmontonia cf. longiceps
Ornithopoda
Euornithopoda
Euornithopod tracks
Hadrosauridae
Edmontosaurus cf. regalis
Hadrosauridae indet.
Ceratopsia
Ceratopsidae
Pachyrhinosaurus canadensis
Anchiceratops sp. (=?A. ornatus)
Dinosauria eggs
Age: early Maastrichtian (Langston
1976)
14. Scollard Formation (Sternberg 1949;
L. S. Russell 1964, 1987; Russell and
Chamney 1967; Currie 1989a; Ryan
and Currie 1998; Eberth et al. 2001b;
Ryan and Russell 2001)
Theropoda
Coelurosauria incertae sedis
cf. Ricardoestesia gilmorei
cf. Ricardoestesia isosceles
Tyrannosauroidea
Tyrannosaurus rex
Tyrannosauridae indet.
Ornithomimosauria
Undescribed ornithomimid
Ornithomimidae indet.
Oviraptorosauria
Caenagnathidae indet.
Troodontidae
cf. Troodon sp.
?Troodontidae indet.
(=cf. Paronychodon sp.)
Dromaeosauridae
Dromaeosaurus albertensis
cf. Saurornitholestes langstoni
Dromaeosauridae indet.
Ankylosauria
Ankylosauridae
Ankylosaurus magniventris
Ornithopoda
Euornithopoda
Thescelosaurus neglectus (including
T. edmontonensis)
Parksosaurus warreni
Iguanodontia
?Iguanodontia indet.
Hadrosauridae
Edmontosaurus regalis
Edmontosaurus annectens
Hadrosauridae indet.
Pachycephalosauria
Pachycephalosauridae
cf. Pachycephalosaurus sp.
Ceratopsia
Neoceratopsia
Leptoceratops gracilis
?Leptoceratops sp.
Ceratopsidae
Triceratops horridus
(=?T.albertensis)
cf. Torosaurus sp.
Ceratopsidae indet.
?Ceratopsid tracks
Age: late Maastrichtian (Russell 1964;
Fox 1978)
15. Willow Creek Formation (Abler 1984;
Carpenter and Alf 1994; Horner,
pers. comm.)
Theropoda
Tyrannosauroidea
Tyrannosaurus rex
Tyrannosauridae indet.
Ornithopoda
Hadrosauridae indet.
Ceratopsia
Neoceratopsia
Montanoceratops sp.
Dinosaur eggs
Age: late Maastrichtian (Schmidt
1978)
1.5.
SASKATCHEWAN
,
CANADA
1. ?Ashville Formation (Tokaryk et al. 1997;
Cumbaa and Tokaryk 1999)
Theropoda
Avialae
?Enantiornithes indet.
Hesperornithiformes
Pasquiaornis hardiei
Pasquiaornis tankei
Ichthyornithiformes
Ichthyornis sp.
Ornithischia indet.
Ornithopoda indet.
Age: late Cenomanian (Tokaryk et al.
1997)
2. Pierre Shale (Tokaryk 1999)
Theropoda
Avialae
Hesperornithoformes
Hesperornis sp.
Age: early Campanian (Tokaryk 1999)
3. Dinosaur Park Formation (Eberth et al.
1990; Tokaryk and Harington 1992;
Tokaryk 1999)
Theropoda
Tyrannosauridae indet.
Ornithomimidae indet.
Oviraptorosauria
cf. Chirostenotes sp.
(=cf. Macrophalangia sp.)
Troodontidae
Troodon formosus
Dromaeosauridae
Saurornitholestes langstoni
Dromaeosaurus sp. (two species?)
Avialae
Hesperornithiformes
Baptornis sp.
Theropoda indet.
Ornithopoda
Hadrosauridae
Hadrosaurinae indet.
Lambeosaurinae indet.
Ceratopsia
Ceratopsidae
Centrosaurus sp.
Ceratopsidae indet.
Dinosaur eggs
Age: late Campanian (Dodson 1971;
Fox 1978)
4. Whitemud Formation (Broughton et al.
1978; Broughton 1981; Schmitz and
Binda 1991)
?Dinosaur coprolites (pseudofossils
according to Coy,
pers. comm.)
Age: late Maastrichtian (Irish and
Havard 1968)
5. Frenchman Formation (Russell 1964;
Tokaryk 1985, 1986a, 1986b, 1997a,
1997b, pers. comm.; Tokaryk and James
1989; Galton 1997b; Chin et al. 1998a,
1998b; McIver 2002)
Theropoda
Coelurosauria incertae sedis
Ricardoestesia gilmorei
Tyrannosauroidea
Tyrannosaurus rex
?Tyrannosaurus sp.
Ornithomimidae indet.
(=Ornithomimus sp.)
Oviraptorosauria
?Caenagnathus sp.
cf. Chirostenotes sp.
(=cf. Macrophalangia sp.)
Troodontidae
Troodon sp. (including
Stenonychosaurus sp.)
Dromaeosauridae
Dromaeosaurus sp.
Saurornitholestes sp.
Ankylosauria
Ankylosauridae
cf. Ankylosaurus sp.
Ornithopoda
Euornithopoda
Thescelosaurus neglectus
Hadrosauridae
Edmontosaurus saskatchewanensis
Hadrosauridae indet.
Pachycephalosauria indet.
Ceratopsia
Ceratopsidae
Triceratops horridus
(=T. cf. prorsus)
Triceratops sp.
Torosaurus latus
Chasmosaurinae indet.
Ceratopsidae indet.
Dinosaur coprolites
Age: late Maastrichtian (Russell 1964)
1.6.
MANITOBA
,
CANADA
1. Vermillion River Formation (Bardack
1965; Martin and Stewart 1982)
Theropoda
Avialae
Hesperornithiformes
Hesperornis regalis
Undescribed hesperornithiform
Ichthyornithiformes indet.
Age: early Campanian (Bardack 1965)
578 DINOSAUR DISTRIBUTION AND BIOLOGY

3.1.
UTAH
,
UNITED STATES
1. Upper Cedar Mountain Formation
(Jensen 1970; Cifelli et al. 1997a, 1999b;
Kirkland et al. 1997, 1998a; Bray 1998;
Carpenter and Kirkland 1998; Chinnery
et al. 1998; Kirkland 1998b; Carpenter
et al. 1999; Lockley et al. 1999)
Theropoda
Tyrannosauroidea incertae sedis
cf. Alectrosaurus sp.
Tyrannosauridae indet.
Undescribed ?ornithomimid
?Troodontidae indet. (including
cf. Paronychodon sp.)
Dromaeosauridae indet.
Avialae
?Hesperornithiformes indet.
Avialae indet.
Theropoda indet. (including
cf. Ricardoestesia sp.)
Sauropoda
Titanosauriformes indet. (including
cf. Pleurocoelus sp.)
Sauropod tracks
Ankylosauria
Nodosauridae
Animantarx ramaljonesi
Ankylosaur tracks
Ornithopoda
Euornithopoda
cf. Zephyrosaurus sp.
Euornithopoda indet.
Iguanodontia
Eolambia caroljonesa
Undescribed iguanodontian
Euornithopod tracks
Pachycephalosauria
Pachycephalosauridae indet.
Ceratopsia
Neoceratopsia indet.
Dinosaur eggs
Age: early Cenomanian (Kirkland et al.
1998a)
2. Dakota Formation (Eaton 1991; Eaton
et al. 1997, 1999a; Carpenter 1999)
Theropoda
Tyrannosauridae indet.
Troodontidae
cf. Troodon sp.
?Troodontidae indet.
(=cf. Paronychodon sp.)
Dromaeosauridae indet.
Theropoda indet. (including
cf. Ricardoestesia sp.)
Sauropoda
Diplodocidae
?Barosaurus lentus
Ankylosauria
Nodosauridae indet.
Ankylosauridae indet.
Ornithopoda
Euornithopoda
Euornithopoda indet.
Iguanodontia indet.
Hadrosauridae indet.
Dinosaur eggs
Age: late Cenomanian (Eaton 1993)
3. Tropic Shale (Gillette et al. 2001)
Theropoda
Therizinosauroidea indet.
Age: Turonian (Gillette et al. 2001)
4. Masuk Formation (Eaton 1990)
Theropoda
Dromaeosauridae indet.
Theropoda indet.
Ankylosauria
Ankylosauridae indet.
Ornithopoda
Hadrosauridae indet.
Pachycephalosauria
Pachycephalosauridae indet.
Age: early Campanian (Eaton 1990)
5. Wahweap Formation (Carpenter and Alf
1994; Eaton et al. 1999a; Parrish 1999)
Theropoda
Tyrannosauridae indet.
Troodontidae
Troodon sp.
Dromaeosauridae indet. (two species?)
Theropoda indet. (including
cf. Aublysodon sp.)
Ankylosauria
Nodosauridae indet.
Ankylosauridae indet.
Ornithopoda
Euornithopoda
cf. Thescelosaurus sp.
Hadrosauridae indet.
Ceratopsia
?Ceratopsidae indet.
Dinosaur eggs
Age: early Campanian (Parrish 1999)
6. Mesaverde Formation (Peterson 1924;
Brown 1938; Lockley 1986a)
Theropod tracks
Ornithopoda
Euornithopod tracks
?Iguanodontia indet.
Ceratopsian tracks
Age: late Campanian (Moelnaar 1983)
7. Neslen Formation (Anderson et al. 1995,
1999)
Ornithopoda
Hadrosauridae indet.
Age: late Campanian (Anderson et al.
1995, 1999)
8. Castlegate Sandstone (Lull and Wright
1942)
Ornithopoda
Hadrosauridae indet.
Age: late Campanian (Fouch et al. 1983)
9. Straight Cliffs Formation (Eaton 1991,
1993; Eaton et al. 1999a, 1999b; Parrish
1999)
Theropoda
Tyrannosauridae indet.
?Troodontidae indet. (including
cf. Paronychodon sp.)
Dromaeosauridae indet.
Theropoda indet. (including
cf. Aublysodon sp.)
?Ornithischia indet.
Ankylosauria
Nodosauridae indet.
Ankylosauridae indet.
Ornithopoda
Euornithopoda
cf. Thescelosaurus sp.
Hadrosauridae indet.
Age: middle Turonian–Santonian
(Parrish 1999)
10. Blackhawk Formation (Carpenter
1992a; McCrea et al. 2001a)
Ankylosaur tracks
Euornithopod tracks
Age: early Campanian (Carpenter
1992a)
11. North Horn Formation (Gilmore
1946b; Jensen 1966; Lawson 1976;
Difley and Ekdale 1998; Lockley 1999;
Cifelli et al. 1999b)
Theropoda
Tyrannosauroidea
Tyrannosaurus rex
Tyrannosauridae indet.
Troodontidae indet.
Avialan tracks
Sauropoda
Lithostrotia
Alamosaurus sanjuanensis
Ornithopoda
Hadrosauridae indet.
Ceratopsia
Ceratopsidae
Torosaurus latus (=T. utahensis)
Ceratopsidae indet.
Dinosaur eggs
Age: late Maastrichtian (Hintze
1973)
12. Kaiparowits Formation (H. E.
Gregory 1950, 1951; DeCourten
1978; Weishampel and Jensen
1979; DeCourten and Russell 1985;
Carpenter and Alf 1994; Eaton et al.
1999a; Parrish 1999; Stadtman et al.
1999; Sullivan and Williamson 1999;
Titus et al. 2001)
Theropoda
Tyrannosauroidea
?Albertosaurus sp.
Tyrannosauridae indet.
Ornithomimosauria
Ornithomimus velox
Troodontidae
Troodon sp.
?Troodontidae indet. (including
cf. Paronychodon sp.)
Dromaeosauridae indet.
Avialae
Enantiornithiformes
Unnamed Avisaurus species
Theropoda indet. (including
cf. Ricardoestesia sp.)
Ankylosauria
Nodosauridae indet.
Ankylosauridae
Euoplocephalus sp.
Ankylosauridae indet.
Ornithopoda
Hadrosauridae
Parasaurolophus cyrtocristatus
Hadrosaurinae indet.
Hadrosauridae indet.
DINOSAUR DISTRIBUTION 579

Pachycephalosauria
Pachycephalosauridae
Stegoceras sp.
Ceratopsia
Ceratopsidae
cf. Triceratops sp.
Ceratopsidae indet.
Dinosaur eggs
Age: late Campanian (Parrish 1999)
3.2.
ARIZONA
,
UNITED STATES
1. Fort Crittenden Formation (Miller
1964; McCord and Tegowski 1996;
Thayer and Ratkevich 1996; Ratkevich
1997; McCord, pers. comm.)
Theropoda
Tyrannosauridae indet.
Theropoda indet.
Sauropoda
Lithostrotia
Alamosaurus sp.
Ornithopoda
Hadrosauridae
Hadrosauridae indet.
Ceratopsia
Centrosaurinae indet.
Age: Campanian (Thayer and Ratkevich
1996)
3.3.
NEW MEXICO
,
UNITED STATES
1. Dakota Sandstone (Heckert and Lucas
1998)
Euornithopod tracks
Age: Cenomanian (Heckert and Lucas
1998)
2. Tres Hermanos Formation (Lucas et al.
1988; Hunt and Lucas 1993a)
Dinosauria indet.
Age: Turonian (Hunt and Lucas 1993a)
3. Moreno Hill Formation (Wolfe et al.
1997; Wolfe and Kirkland 1998; Lucas
et al. 2000; Kirkland and Wolfe 2001)
Theropoda
Tyrannosauridae indet.
Therizinosauroidea
Nothronychus mckinleyi
Dromaeosauridae indet.
Ankylosauria
Nodosauridae indet.
Ornithopoda
Hadrosauridae indet.
Ceratopsia
Neoceratopsia
Zuniceratops christopheri
Age: middle Turonian (Wolfe et al. 1997)
4. Crevasse Canyon Formation (Lucas et al.
1988; Hunt and Lucas 1993a)
Dinosauria indet.
Ornithopoda
Iguanodontia indet.
Age: early Santonian (Hunt and Lucas
1993a)
5. Point Lookout Sandstone (Lucas et al.
1988, 2000; Williamson et al. 1989;
Hunt and Lucas 1993a)
Theropoda
Tyrannosauridae indet.
Dromaeosauridae indet.
Ornithopoda
Hadrosauridae indet.
Age: Santonian (Hunt and Lucas 1993a)
6. Menefee Formation (Hunt and Lucas
1993a; Lucas et al. 2000)
Theropoda
Tyrannosauridae indet.
Dromaeosauridae
cf. Saurornitholestes sp.
Ankylosauria indet.
Ornithopoda
Hadrosauridae indet.
Ceratopsidae
Centrosaurinae indet.
Age: late Santonian–early Campanian
(Hunt and Lucas 1993a)
7. Fruitland Formation (Gilmore 1916,
1919a, 1935; Osborn 1923b; Ostrom
1961a; Armstrong Ziegler 1978, 1980;
Lucas 1981; Rowe et al. 1981; Currie
1987a; Lucas et al. 1987, 2000; Hunt
1991; Hall 1993; Hunt and Lucas 1993a;
Lehman 1998; Sullivan and Williamson
1999)
Theropoda
Tyrannosauroidea
?Gorgosaurus libratus
?Albertosaurus sp.
Unnamed tyrannosauroid
Tyrannosauridae indet.
Ornithomimosauria
cf. Ornithomimus sp.
Ornithomimidae indet.
?Troodontidae indet. (=Paronychodon
lacustris)
Dromaeosauridae indet.
Theropoda indet.
Ankylosauria
Nodosauridae indet.
Ankylosauria indet.
Ornithopoda
Euornithopoda
?Thescelosaurus sp.
Hadrosauridae
Parasaurolophus cyrtocristatus
?Corythosaurus sp.
Hadrosaurinae indet.
(=?Kritosaurus navajovius)
Euornithopod tracks
Pachycephalosauria
Pachycephalosauridae indet.
Ceratopsia
Ceratopsidae
Pentaceratops sternbergii (including
P. fenestratus)
?Chasmosaurus sp.
Ceratopsidae indet.
(=?Monoclonius sp.)
Dinosaur eggs
Age: late Campanian (Hunt and Lucas
1993a)
8. Ringbone Formation (Lucas et al. 1990;
Hunt and Lucas 1993a; Anderson et al.
1998)
Theropoda
Tyrannosauridae indet.
Ornithopoda
Hadrosauridae indet.
Age: late Campanian (Lucas et al. 1990;
Hunt and Lucas 1993a)
9. Pictured Cliffs Sandstone (Williamson
and Lucas 1992; Hunt and Lucas 1993a)
Theropoda indet.
Ornithopoda
Hadrosauridae indet.
Age: late Campanian (Hunt and Lucas
1993a)
10. Lower Kirtland Formation (Gilmore
1916, 1919a, 1935; Osborn 1923b;
Wiman 1930, 1931; Lucas 1981; Mateer
1981; Currie 1987a; Lucas et al. 1987,
2000; Hunt 1991; Hunt and Lucas 1992,
1993a; Sullivan 1997, 1998b, 1999;
Sullivan and Williamson 1997, 1999;
Carr and Williamson 1999; Williamson
and Sealey 1999; Sullivan and Lucas
2000)
Theropoda
Tyrannosauroidea
Albertosaurus sp.
Unnamed Daspletosaurus species
Tyrannosauroidea indet.
(including Aublysodon
cf. mirandus)
Ornithomimosauria
cf. Struthiomimus sp.
Ornithomimosauria indet.
(=Ornithomimus antiquus)
Dromaeosauridae
Saurornitholestes langstoni
Dromaeosauridae indet.
Sauropoda
Lithostrotia
Alamosaurus sanjuanensis
Ankylosauria
Nodosauridae indet. (including
Edmontonia australis)
Ankylosauridae
Nodocephalosaurus kirtlandensis
Ornithopoda
Hadrosauridae
Anasazisaurus horneri
Naashoibitosaurus ostromi
Parasaurolophus tubicen
cf. Parasaurolophus sp.
Hadrosaurinae indet.
(=Kritosaurus navajovius)
Hadrosauridae indet.
Pachycephalosauria
Undescribed pachycephalosaurid
Ceratopsia
Ceratopsidae
Pentaceratops sternbergii (including
P. fenestratus)
Pentaceratops sp.
Age: late Campanian (Sullivan and
Lucas 2000); early
Maastrichtian (Hunt and
Lucas 1992, 1993a)
11. Upper Kirtland Formation (Gilmore
1916, 1919a, 1935; Osborn 1923b;
Wiman 1930, 1931; Lucas 1981; Mateer
1981; Currie 1987a; Lucas et al. 1987,
2000; Hunt and Lucas 1992, 1993a;
Williamson and Sullivan 1996; Sullivan
and Williamson 1997, 1999)
580 DINOSAUR DISTRIBUTION AND BIOLOGY

Theropoda
Tyrannosauroidea
?Albertosaurus sp.
cf. Tyrannosaurus sp.
Tyrannosauridae indet.
Ornithomimidae indet.
Troodontidae indet.
Dromaeosauridae indet.
Sauropoda
Lithostrotia
Alamosaurus sanjuanensis
Ankylosauria
Nodosauridae indet. (including
Glyptodontopelta mimus)
Ankylosauridae indet.
Ankylosauria indet.
Ornithopoda
Hadrosauridae
Hadrosauridae indet.
Ceratopsia
Ceratopsidae
cf. Torosaurus latus (=T. utahensis)
Pentaceratops sp.
Ceratopsidae indet.
Age: late Maastrichtian (Hunt and
Lucas 1993a)
12. McRae Formation (Lozinsky et al. 1984;
Hunt and Lucas 1993a; Lucas et al.
1998b, 2000; =Red Beds of New
Mexico, Lull 1933)
Theropoda
Tyrannosauroidea
Tyrannosaurus rex
Theropoda indet.
Sauropoda
Lithostrotia
Alamosaurus sp.
Ankylosauria indet.
Ceratopsia
Ceratopsidae
Torosaurus latus
Ceratopsidae indet.
Age: late Maastrichtian (Hunt and
Lucas 1993a)
13. Raton Formation (Lockley and Hunt
1994; Lockley et al. 2000)
Theropod tracks
Age: Maastrichtian (Lockley et al. 2000)
3.4.
COLORADO
,
UNITED STATES
1. Iles Formation (Mateer, pers. comm.)
Dinosauria indet.
Age: late Campanian (Mateer,
pers. comm.)
2. Williams Fork Formation (Archibald
1987a)
Theropoda indet.
Ornithischia indet.
Age: Maastrichtian (Kiteley 1983)
3. Mesaverde Formation (Peterson 1924;
Brown 1938; Lockley et al. 1982;
Carpenter 1992a)
Theropod tracks
Euornithopod tracks
Age: early Maastrichtian (Carpenter
1992a)
4. Laramie Formation (Carpenter 1979;
Carpenter and Breithaupt 1986; Lockley
1986a; Lockley and Hunt 1995b;
Wright and Lockley 2001; Carpenter,
pers. comm.)
Theropoda
Tyrannosauroidea
cf. Tyrannosaurus rex
?Troodontidae indet.
(=Paronychodon sp.)
?Dromaeosauridae indet.
?Avialae indet. (=Ornithomimus
minutus)
Theropoda/?Troodontidae indet.
(=Tripriodon caperatus)
Theropod tracks
Ankylosauria
Nodosauridae
?Edmontonia sp.
Ornithopoda
Euornithopod tracks
Euornithopoda
Thescelosaurus neglectus
Thescelosaurus sp.
Hadrosauridae
Edmontosaurus regalis
Edmontosaurus annectens
Ceratopsia
Ceratopsidae
Triceratops horridus
Triceratops sp.
Torosaurus latus
Ceratopsid tracks
Age: late Maastrichtian (Carpenter
1979)
5. Arapahoe Formation (Wright
and Lockley 2001; Carpenter,
pers. comm.)
Theropod tracks (may be from
Laramie Formation)
Ornithopod tracks (may be from
Laramie Formation)
Ceratopsia indet.
Age: late Maastrichtian (Cobban and
Reeside 1952; Carpenter,
pers. comm.)
6. Denver Formation (Carpenter,
pers. comm.)
Theropoda
Tyrannosauroidea
Tyrannosaurus rex
Tyrannosauroidea indet.
(=Aublysodon mirandus)
Ornithomimosauria
Ornithomimus velox
Ornithomimidae indet.
Theropoda indet.
Ankylosauria
Nodosauridae indet.
Ornithopoda
Hadrosauridae
Edmontosaurus sp.
Hadrosauridae indet. (=Cionodon
arctatus)
Pachycephalosauria
Pachycephalosauridae
Pachycephalosaurus sp.
Ceratopsia
Ceratopsidae
Triceratops sp.
Ceratopsidae indet. (=Polyonax
mortuarius, Triceratops galeus)
Age: late Maastrichtian (Cobban and
Reeside 1952; Carpenter,
pers. comm.)
7. Cimarron Ridge Formation (Lee 1912;
Reeside 1924; Lehman 1987)
Dinosauria indet.
Age: late Maastrichtian (Lehman 1987)
8. Raton Formation (Lockley and Hunt
1994; Lockley et al. 2000)
Theropod tracks (including avialan
tracks)
Ornithopoda
Hadrosaurid tracks
?Ceratopsian tracks
Age: Maastrichtian (Lockley et al. 2000)
3.5.
TEXAS
,
UNITED STATES
1. Woodbine Formation (Lee 1997a, 1997b;
Head 1998; Jacobs and Winkler 1998)
Theropoda
Theropoda incertae sedis
cf. Ricardoestesia sp.
Theropoda indet.
Theropod tracks
Ankylosauria
Nodosauridae indet.
Ornithopoda
Iguanodontia
Protohadros byrdi
Hadrosauridae indet.
Euornithopod tracks
Age: middle Cenomanian (Birkelund
et al. 1984)
2. Pen Formation (Lehman 1985b)
Ornithopoda
Hadrosauridae
?Gryposaurus sp. (=Kritosaurus sp.)
Age: Cenomanian-early Campanian
(Lehman 1985a)
3. San Carlos Formation (Lehman 1985b)
Theropoda
Tyrannosauridae indet.
Ankylosauria
Nodosauridae indet.
Ornithopoda
Hadrosauridae
?Gryposaurus sp. (=Kritosaurus sp.)
Pachycephalosauria
Pachycephalosauridae
Stegoceras sp.
Ceratopsia
Ceratopsidae indet.
Age: early-late Campanian (Lehman
1985a)
4. Aguja Formation (Lull and Wright 1942;
Davies 1983; Lehman 1985b, 1989;
Carpenter 1990b; Rowe et al. 1992;
Baghai-Riding and DiBenedetto 2001;
Sankey 2001; Wagner and Lehman 2001)
Theropoda
Coelurosauria incertae sedis
Ricardoestesia cf. gilmorei
Ricardoestesia isosceles
cf. Ricardoestesia sp.
Tyrannosauridae indet.
Ornithomimidae indet.
DINOSAUR DISTRIBUTION 581

Troodontidae
cf. Troodon sp.
Dromaeosauridae
Saurornitholestes cf. langstoni
cf. Saurornitholestes sp.
cf. Dromaeosaurus sp.
Theropoda indet.
Ornithischia indet.
Ankylosauria
Nodosauridae
Edmontonia cf. rugosidens
Unnamed Panoplosaurus species
Unnamed Euoplocephalus species
Nodosauridae indet.
Ankylosauridae indet.
Ornithopoda
Iguanodontia
Hadrosauridae
Undescribed hadrosaurine
?Gryposaurus sp. (=Kritosaurus sp.)
Hadrosaurinae indet. (=Kritosaurus
cf. navajovius)
Lambeosaurinae indet.
Hadrosauridae indet.
Ornithopoda indet.
Pachycephalosauria
Pachycephalosauridae
Stegoceras sp.
Pachycephalosauridae indet.
Ceratopsia
Ceratopsidae
Chasmosaurus mariscalensis
Ceratopsidae indet.
Dinosaur coprolites
Age: early-late Campanian (Lehman
1985a)
5. Javelina Formation (Lawson 1976;
Lehman 1985b; Fiorillo 1998d; Davies,
pers. comm.)
Theropoda
Tyrannosauroidea
Tyrannosaurus rex
Tyrannosauridae indet.
Ornithomimidae indet.
Sauropoda
Lithostrotia
Alamosaurus sanjuanensis
Ankylosauria
Nodosauridae indet.
Ornithopoda
Hadrosauridae
cf. Edmontosaurus sp.
Ceratopsia
Ceratopsidae
cf. Torosaurus latus
(=cf. T. utahensis)
Ceratopsidae indet.
Age: late Maastrichtian (Lehman 1985a)
6. El Picacho Formation (Lehman 1985b,
1996)
Theropoda
Tyrannosauridae indet.
Ornithomimidae indet.
Sauropoda
Lithostrotia
Alamosaurus sanjuanensis
Ankylosauria
Nodosauridae indet.
Ornithopoda
Hadrosauridae
?Gryposaurus sp. (=Kritosaurus sp.)
Ceratopsia
Ceratopsidae indet.
Age: late Maastrichtian (Lehman 1985a)
7. Black Peaks Formation (Lehman and
Coulson 2002)
Sauropoda
Lithostrotia
Alamosaurus sanjuanensis
Age: late Maastrichtian (Lehman and
Coulson 2002)
8. Austin Chalk (Marsh 1877b; Clarke,
pers. comm.) (not plotted)
Theropoda
Avialae
Unnamed ichthyornithiform
Age: Late Cretaceous (Marsh 1877b)
3.7.
KANSAS
,
UNITED STATES
1. Greenhorn Formation (Martin and
Stewart 1982; Martin 1983)
Theropoda
Avialae
Hesperornithiformes indet.
Ichthyornithiformes
Ichthyornis sp.
Age: Cenomanian-Turonian (Martin and
Stewart 1982; Martin 1983)
2. Niobrara Chalk Formation (Marsh
1872a, 1872b, 1872c, 1872d, 1873a,
1873b, 1876, 1877b, 1890a; Mehl 1936;
Carpenter et al. 1995; Clarke, pers.
comm.)
Theropoda
Avialae
Ornithurae
Apatornis celer
Hesperornithiformes
Baptornis adventus
Hesperornis regalis
Hesperornis crassipes
Hesperornis gracilis
Parahesperornis alexi
Ichthyornithiformes
Ichthyornis dispar (including
I. agilis, I. anceps, I. validus,
I. victor)
Unnamed ichthyornithiforms
(=Ichthyornis lentus, I. tener)
Ankylosauria incertae sedis
Niobrarasaurus coleii
Ankylosauria indet. (=Hierosaurus
sternbergii)
Ornithopoda
Hadrosauridae
Claosaurus agilis
Age: Coniacian–early Campanian
(Hattin 1982)
3.8.
OREGON
,
UNITED STATES
1. Cape Sebastian Sandstone (Leahy, pers.
comm.)
?Ornithopoda
?Hadrosauridae indet.
Age: late Campanian (Bourgeois and
Dott 1985)
3.9.
NEVADA
,
UNITED STATES
1. Newark Canyon Formation (Smith et al.
1976)
Theropoda indet.
Ankylosauria
Ankylosauridae indet.
Age: ?Late Cretaceous (Smit h et al. 1976)
2. Unnamed unit (Carpenter, pers. comm.)
Theropoda indet.
Ankylosauria indet.
Ornithopoda
?Hadrosauridae indet.
Ceratopsia
Neoceratopsia
?Leptoceratops sp.
Age: Late Cretaceous (Carpenter,
pers. comm.)
3.10.
CALIFORNIA
,
UNITED STATES
1. Ladd or Williams Formation (Morris
1973b)
Ornithopoda
Hadrosauridae indet.
Age: ?Campanian (Popenoe et al. 1960)
2. Chico Formation (Embree, pers. comm.;
Hilton, pers. comm.)
Theropoda indet.
Age: Campanian (Embree, pers. comm.)
3. Point Loma Formation (Morris 1982;
Ford and Kirkland 2001; Deméré,
pers. comm.)
Ankylosauria
Ankylosauridae
Aletopelta coombsi
Ornithopoda
Hadrosauridae indet.
Age: late Campanian–early
Maastrichtian (Nilsen
and Abbott 1981)
4. Moreno Formation (Morris 1973b,
1982)
Ornithopoda
Hadrosauridae
Hadrosaurinae indet.
(=?Saurolophus sp.)
Age: late Maastrichtian (Popenoe et al.
1960)
3.11.
MONTANA
,
UNITED STATES
1. Telegraph Creek Formation
(unpublished)
Ornithopoda
Hadrosauridae indet.
Age: late Santonian (Cobban 1955)
2. Lower Two Medicine Formation
(Gilmore 1930; Horner 1989, 1992, pers.
comm.; Mongelli and Varricchio 1998)
Theropoda
Theropoda incertae sedis
Ricardoestesia sp.
Tyrannosauroidea
cf. Albertosaurus sp.
cf. Aublysodon sp.
Tyrannosauridae indet.
Troodontidae
Troodon sp.
Dromaeosauridae
Dromaeosaurus sp.
582 DINOSAUR DISTRIBUTION AND BIOLOGY

Saurornitholestes sp.
Dromaeosauridae indet.
Theropoda indet.
Ankylosauria
Ankylosauridae indet.
Ornithopoda
Hadrosauridae
Gryposaurus latidens
Hadrosauridae indet.
Ceratopsia
Undescribed neoceratopsian
Undescribed ceratopsid
Dinosaur eggs
Age: ?late Santonian–early Campanian
(Horner, pers. comm.)
3. Eagle Sandstone (Marsh 1890b)
Theropoda
Tyrannosauroidea incertae sedis
“Ornithomimus” grandis
Age: early Campanian (Sahni 1972)
4. Claggett Shale (Marsh 1893; Fiorillo
1990)
Theropoda
Avialae
Hesperornithiformes
Coniornis altus
Hesperornis sp.
Ornithopoda
Hadrosauridae
cf. Maiasaura sp.
Age: late Campanian (Fiorillo 1990)
5. Judith River Formation (Marsh 1893;
Sahni 1972; Dodson 1984, 1986; Horner
1988, 1989, pers. comm.; Fiorillo and
Currie 1994; Galton 1995)
Theropoda
Tetanurae indet. (=Aublysodon
lateralis)
Coelurosauria incertae sedis
Ricardoestesia gilmorei
Tyrannosauroidea
Gorgosaurus libratus (?including
?Laelaps falculus)
Albertosaurus sarcophagus
Tyrannosauroidea indet.
(=Aublysodon mirandus,
Deinodon horridus)
Ornithomimosauria
Ornithomimus sp.
Ornithomimosauria indet.
(=Ornithomimus tenuis)
Troodontidae
Troodon formosus
?Troodontidae indet. (=Laelaps
explanatus, L. laevifrons,
Paronychodon lacustris,
Zapsalis abradens)
Dromaeosauridae
Dromaeosaurus albertensis
Saurornitholestes langstoni
Ankylosauria
Ankylosauria indet. (=Palaeoscincus
costatus)
Nodosauridae
Edmontonia longiceps
Edmontonia rugosidens
Ornithopoda
Euornithopoda
?Orodromeus makelai
(=Thescelosaurus neglectus)
Hadrosauridae
Brachylophosaurus canadensis
Brachylophosaurus sp.
Hadrosauridae indet. (=Diclonius
calamarius, D. pentagonus,
D. perangulatus, Dysganus
encaustus partim,
Hadrosaurus breviceps,
H. paucidens)
Pachycephalosauria
Pachycephalosauridae
Stegoceras validum
Stegoceras sp.
Pachycephalosaurus wyomingensis
Ceratopsia
Ceratopsidae
Monoclonius crassus
Avaceratops lammersi
Undescribed ceratopsid
Ceratopsidae indet. (=Ceratops
montanus, Diclonius
bicarinatus, D. encaustus
partim, D. haydenianus,
D. peiganus, Monoclonius
fissus, M. recurvicornis,
M. sphenocerus,)
Dinosaur eggs
Age: late Campanian (Goodwin
and Deino 1989)
6. Upper Two Medicine Formation
(Gilmore 1930; Horner and Makela
1979; Horner 1983, 1989, 1992,
1999; Horner and Weishampel 1988;
Varricchio and Horner 1993; Sampson
1995; Varricchio and Chiappe 1995;
Ryan and Currie 1998; Varricchio et al.
1999; Burnham et al. 2000; Trexler 2001)
Theropoda
Coelurosauria incertae sedis
Ricardoestesia sp.
Tyrannosauroidea
Daspletosaurus torosus
cf. Albertosaurus sp.
Tyrannosauroidea indet.
(=Aublysodon mirandus)
Oviraptorosauria
Chirostenotes pergracilis
Ornithomimidae indet.
Troodontidae
Troodon formosus
Dromaeosauridae
Bambiraptor feinbergorum
Saurornitholestes sp.
Dromaeosauridae indet.
Avialae
Enantiornithes
Avisaurus gloriae
Ankylosauria
Nodosauridae
Edmontonia rugosidens
Ankylosauridae
Euoplocephalus tutus
Ankylosauridae indet.
Ornithopoda
Euornithopoda
Orodromeus makelai
Undescribed euornithopodan
Hadrosauridae
Maiasaura peeblesorum
Prosaurolophus blackfeetensis
Gryposaurus sp.
Hypacrosaurus stebingeri
Hadrosauridae indet.
Pachycephalosauria
Pachycephalosauridae indet.
Ceratopsia
Neoceratopsia
Leptoceratops sp.
Ceratopsidae
Einiosaurus procurvicornis
Achelousaurus horneri
Brachyceratops montanensis
Ceratopsidae indet. (=Styracosaurus
ovatus)
Dinosaur eggs
Age: middle-late Campanian (Horner,
pers. comm.)
7. Lower Bearpaw Shale Formation
(Horner 1979)
Ankylosauria
Nodosauridae
Edmontonia sp.
Ornithopoda
Hadrosauridae
Unnamed hadrosaurine
(=cf. Kritosaurus sp.)
Lambeosaurinae indet.
(=?Lambeosaurus
magnicristatus)
Age: late Campanian (Rice and Shurr
1983)
8. Cokedale Formation (Horner 1989)
Theropoda
Tyrannosauridae indet.
Ornithopoda
Hadrosauridae indet.
Age: Campanian (Horner 1989)
9. Horsethief Sandstone (Bibler and
Schmidt 1986; Horner 1989, pers.
comm.)
Ornithopoda
Hadrosauridae indet.
Dinosaur eggs
Age: early Maastrichtian (Viele and
Harris 1965)
10. Livingston Formation (McMannis
1965; Skipp and McGrew 1972;
Horner, pers. comm.)
Theropoda
Tyrannosauroidea
?Tyrannosaurus rex
Tyrannosauridae indet.
Ornithopoda
Hadrosauridae indet.
Ceratopsia
Neoceratopsia
?Leptoceratops sp.
Age: ?late Campanian–late
Maastrichtian (Cobban
and Reeside 1952)
11. St. Mary River Formation (Brown and
Schlaikjer 1942; Weishampel and
Horner 1987; Chinner y and
Weishampel 1998)
DINOSAUR DISTRIBUTION 583

Theropoda
Tyrannosauridae indet.
Ornithomimidae indet.
?Dromaeosauridae indet.
Ornithopoda
Hadrosauridae
Hadrosaurinae indet.
Undescribed lambeosaurine
Lambeosaurinae indet.
Ceratopsia
Neoceratopsia
Montanoceratops cerorhynchus
Ceratopsidae indet.
Dinosaur eggs
Age: early Maastrichtian (Cobban
and Reeside 1952)
12. Billman Creek Formation (Horner
1989)
Theropoda indet.
Ornithopoda
Hadrosauridae indet.
Age: early Maastrichtian (Roberts 1972)
13. Willow Creek Formation (Russell 1968;
may be St. Mary River Formation)
Ornithopoda
Euornithopoda
?Thescelosaurus sp.
Age: late Maastrichtian (Honkala 1955)
14. Hell Creek Formation (including
questionable Lance strata) (Jepsen
1931; Estes et al. 1969; Molnar 1978,
1980d; Lupton et al. 1980; Brett
Surman and Paul 1985; Carpenter
and Breithaupt 1986; Sloan et al.
1986; Currie 1987a; Giffin et al. 1987;
Hutchinson and Chiappe 1998; Galton
1999a; Buckley and Ott 2001; Ott and
Buckley 2001; Stidham and Hutchison
2001; Varricchio 2001b; Williamson
and Carr 2002)
Theropoda
Tyrannosauroidea
Tyrannosaurus rex (including
Albertosaurus megagracilis,
Aublysodon molnari)
Nanotyrannus lancensis
Tyrannosauroidea indet.
(including Aublysodon
mirandus)
Ornithomimosauria
Ornithomimus sp.
Ornithomimidae indet.
Oviraptorosauria
Chirostenotes elegans (=Elmisaurus
elegans)
?Chirostenotes sp.
Troodontidae
Troodon formosus
(=?Saurornithoides sp.,
Stenonynchosaurus inequalis)
?Troodontidae indet.
(=Paronychodon lacustris)
Dromaeosauridae
Dromaeosaurus sp.
Dromaeosauridae indet.
(two species?)
Avialae
Unnamed alvarezsaurid
Enantiornithes
Avisaurus archibaldi
Theropoda indet.
Ornithischia indet.
Ankylosauria
Ankylosauria indet.
Nodosauridae
Unnamed Edmontonia species
Ankylosauridae
Ankylosaurus magniventris
Ankylosauridae indet.
Ornithopoda
Euornithopoda
?Bugenasaura infernalis
(=?Thescelosaurus garbanii)
?Thescelosaurus sp.
Hadrosauridae
Edmontosaurus regalis
Edmontosaurus annectens
(including Anatotitan copei)
?Parasaurolophus walkeri
Pachycephalosauria
Pachycephalosauridae
Pachycephalosaurus wyomingensis
Stegoceras edmontonense
Stygimoloch spinifer (including
Stenotholus kohleri)
Sphaerotholus buchholtzae
Sphaerotholus goodwini
Stegoceras sp.
Ceratopsia
Neoceratopsia
Leptoceratops cf. gracilis
Ceratopsidae
Triceratops horridus (=T. prorsus,
T. serratus)
Triceratops sp.
Torosaurus latus
Ceratopsidae indet. (including
Triceratops maximus,
Ugrosaurus olsoni)
Dinosaur eggs
Age: late Maastrichtian (Cobban and
Reeside 1952; Baadsgaard
and Lerbekmo 1980)
3.12.
WYOMING
,
UNITED STATES
1. Frontier Formation (Marsh 1889a; Lull
1921; Coombs 1978a; Carpenter and
Kirkland 1998)
Ankylosauria incertae sedis
Nodosaurus textilis
Stegopelta landerensis
Age: middle Cenomanian (Merewether
1983)
2. Pierre Shale Formation (Carpenter,
pers. comm.)
Ornithopoda
Undescribed hadrosaurid
Age: early Campanian (Gill and Cobban
1966)
3. Meeteetse Formation (Keefer and Troyer
1956; Keefer 1965; Storrs 1991)
Dinosauria indet.
Theropoda
Tyrannosauridae indet.
Ornithopoda
Hadrosauridae indet.
Dinosaur eggs
Age: late Campanian (Lillegraven and
McKenna 1986)
4. Almond Formation (Breithaupt 1985;
Carpenter 1992a)
Theropoda
Tyrannosauridae indet.
?Troodontidae indet. (=Paronychodon
lacustris)
Dromaeosauridae
Dromaeosaurus sp.
Theropoda indet.
Ankylosauria
Nodosauridae
Edmontonia sp.
Ornithopoda
Euornithopoda
cf. Thescelosaurus sp.
Hadrosauridae
Edmontosaurus sp.
Maiasaura sp.
Lambeosaurinae indet.
Hadrosauridae indet.
Ceratopsia
Ceratopsidae
?Anchiceratops sp.
Ceratopsidae indet.
Age: early Maastrichtian (Carpenter
1992a); late Campanian
(Breithaupt 1985)
5. Harebell Formation (McKenna 1980;
Harris et al. 1996)
Theropod tracks
Ankylosauria
?Nodosauridae indet.
Ceratopsia
Neoceratopsia indet.
(=?Leptoceratops sp.)
Age: Maastrichtian (Harris et al. 1996)
6. Pinyon Conglomerate (McKenna and
Love 1970)
Ceratopsia
Neoceratopsia
Leptoceratops sp.
Age: Maastrichtian (McKenna and Love
1970 )
7. Medicine Bow Formation (Lull 1933;
Lillegraven and Eberle 1999)
Theropoda
Tyrannosauroidea
Aublysodon mirabilis
Ornithopoda
Euornithopoda
Thescelosaurus sp.
Hadrosauridae
Hadrosaurinae indet.
Ceratopsia
Ceratopsidae
Chasmosaurinae indet.
(=Triceratops sp.)
Age: Maastrichtian (Cobban and Reeside
1952)
8. Evanston Formation (Tracy and Oriel
1959; Rubey et al. 1961; Lucas 1981;
Breithaupt 1985)
Sauropoda
Lithostrotia
Alamosaurus sp.
584 DINOSAUR DISTRIBUTION AND BIOLOGY

Ceratopsia
Ceratopsidae
Triceratops horridus
(=T. flabellatus)
Age: late Maastrichtian (Cobban
and Reeside 1952)
9. Ferris Formation (Bowen 1918; Lull 1933;
Lillegraven and Eberle 1999)
Theropoda
Theropoda incertae sedis
Ricardoestesia sp.
Tyrannosauroidea
Tyrannosaurus rex
Tyrannosauroidea indet. (including
Aublysodon mirandus)
Ornithomimosauria
Ornithomimus cf. velox
cf. Struthiomimus sp.
Ornithomimid indet.
Troodontidae
Troodon formosus
?Troodontidae indet. (=Paronychodon
lacustris)
Dromaeosauridae
Dromaeosaurus sp.
Saurornitholestes sp.
Avialae indet.
Theropod indet.
Ankylosauria
Nodosauridae
Edmontonia cf. rugosidens
Ankylosauridae
Ankylosaurus sp.
Ornithopoda
Euornithopoda
Thescelosaurus sp.
Hadrosauridae
Hadrosaurinae indet.
Pachycephalosauria
Pachycephalosauridae
Stygimoloch spinifer
Ceratopsia
Ceratopsidae
Chasmosaurinae indet.
(=Triceratops sp.)
Age: late Maastrichtian (Cobban
and Reeside 1952)
10. Lance Formation (including
questionable Lance strata) (Marsh
1889a, 1889b, 1889c, 1892e; Hatcher
1893; Hatcher et al. 1907; Bowen
1915; Gilmore 1915a; Brodkorb 1963;
Estes 1964; Ostrom 1978b; Breithaupt
1982, 1985, 1994; Carpenter 1982b;
Carpenter and Breithaupt 1986; Currie
1987a; McIntosh et al. 1992; Derstler
1995; Finlayson 1998; Stidham 1998;
Galton 1999a; Hope 1999; Elz˙anowski
et al. 2001)
Theropoda
Coelurosauria incertae sedis
Ricardoestesia sp.
Tyrannosauroidea
Albertosaurus sarcophagus
cf. Tyrannosaurus rex
Tyrannosaurus sp.
Ornithomimosauria
Ornithomimus sp.
Ornithomimosauria indet.
(=Ornithomimus sedens)
Oviraptorosauria
?Chirostenotes sp.
Troodontidae
Troodon formosus (=Pectinodon
bakkeri, ?Saurornithoides sp.,
Stenonychosaurus inequalis)
?Troodontidae indet. (=Laelaps
cristatus, Paronychodon
lacustris)
Dromaeosauridae
?Velociraptor sp.
Dromaeosaurus sp.
Saurornitholestes sp.
Dromaeosauridae indet.
Avialae
Enantiornithes indet.
?Hesperornithiformes
Potamornis skutchi
Neornithes
Galliformes
Palintropus retusus
Tetanurae indet. (including
Aublysodon amplus,
A. mirandus, ?Dryptosaurus sp.)
Sauropoda
Diplodocidae
Dyslocosaurus polyonychius
(may be from the Morrison
Formation)
Ankylosauria
Ankylosauria indet. (=Palaeoscincus
latus)
Nodosauridae
Unnamed Edmontonia species
Nodosauridae indet.
Ankylosauridae
Ankylosaurus cf. magniventris
Ankylosauridae indet.
Ornithopoda
Euornithopoda
Thescelosaurus neglectus
Bugenasaura infernalis
?Thescelosaurus sp.
Hadrosauridae
Edmontosaurus regalis
Edmontosaurus annectens
?Lambeosaurus sp.
Hadrosauridae indet.
Pachycephalosauria
Pachycephalosauridae
Pachycephalosaurus wyomingensis
cf. Pachycephalosaurus sp.
Stygimoloch spinifer
Ceratopsia
Neoceratopsia
Leptoceratops gracilis
Ceratopsidae
Diceratops hatcheri
Triceratops horridus (including
T. calicornis, T. elatus,
T. eurycephalus, T. flabellatus,
T. obtusus, T. prorsus,
T. serratus)
Triceratops sp.
Torosaurus latus (including
T. gladius)
Ceratopsidae indet. (=Agathaumas
sylvestris, Bison alticornis,
Triceratops ingens, T. sulcatus)
Age: late Maastrichtian (Cobban and
Reeside 1952)
3.14.
NORTH DAKOTA
,
UNITED STATES
1. Fox Hills Formation (Hoganson et al.
1996)
Theropoda indet.
Age: Maastrichtian (Hoganson et al. 1996)
2. Hell Creek Formation (Lull 1933; Lull
and Wright 1942; Pearson et al. 2001;
Farrar pers. comm.; Fastovsky, pers.
comm.; Williams, pers. comm.;)
Theropoda
Coelurosauria indet.
(=Ricardoestesia sp.)
Tyrannosauroidea
Tyrannosaurus rex
Tyrannosauridae indet.
Ornithomimosauria
cf. Struthiomimus sp.
?Ornithomimus sp.
Ornithomimidae indet.
Oviraptorosauria
Caenagnathidae indet.
?Troodontidae indet. (=Paronychodon
lacustris)
Dromaeosauridae
Saurornitholestes sp.
Avialae
Hesperornithiformes indet.
Avialae indet.
Theropoda indet.
Ankylosauria
Nodosauridae
Unnamed Edmontonia species
Ankylosauria indet.
Ornithopoda
Euornithopoda
cf. Thescelosaurus sp.
Hadrosauridae
Edmontosaurus regalis
Hadrosauridae indet.
Pachycephalosauria
Undescribed pachycephalosaurid
Ceratopsia
Ceratopsidae
Torosaurus latus
Triceratops horridus
Triceratops sp.
Ceratopsidae indet.
Age: late Maastrichtian (Cobban and
Reeside 1952)
3.15.
SOUTH DAKOTA
,
UNITED STATES
1. Pierre Shale Formation (Wieland 1903;
Gregory 1948; Russell 1967)
Theropoda
Avialae
Hesperornithiformes
Hesperornis sp.
Ornithopoda
Hadrosauridae indet. (=Claosaurus
affinis)
Age: early Campanian (Gill and Cobban
1966)
DINOSAUR DISTRIBUTION 585

2. Lance Formation (Lull and Wright 1942;
Bakker 1988)
Ankylosauria
Nodosauridae
Edmontonia longiceps
(=Denversaurus schlessmani)
Ornithopoda
Hadrosauridae
Edmontosaurus annectens
(=Anatotitan copei)
?Edmontosaurus regalis
Hadrosauridae indet. (=Thespesius
occidentalis)
Age: late Maastrichtian (Cobban and
Reeside 1952)
3. Hell Creek Formation (Colbert and
Bump 1947; Morris 1976; Bjork 1985b;
Carpenter and Breithaupt 1986;
Woods and Stucky 1992; Currie et al.
1993; Galton 1995; Triebold and Russell
1995; Holland 1997; Triebold 1997;
Jacobson and Sroka 1998; Triebold
et al. 1999)
Theropoda
Tyrannosauroidea
Tyrannosaurus rex
Albertosaurus sp.
Ornithomimosauria
Ornithomimus sp.
Struthiomimus sp.
Ornithomimidae indet.
Oviraptorosauria indet.
Caenagnathidae
Caenagnathus sp.
Chirostenotes pergracilis
(=Caenagnathus collinsi)
Chirostenotes sp.
Oviraptorosauria indet.
Troodontidae
Troodon sp.
Troodontidae indet. (including
Paronychodon lacustris)
Dromaeosauridae
Dromaeosaurus sp.
Saurornitholestes sp.
Dromaeosauridae indet.
Undescribed theropod
Theropod tracks
Ankylosauria
Nodosauridae
Edmontonia sp.
Ornithopoda
Euornithopoda
Bugenasaura infernalis (including
Thescelosaurus garbani)
Thescelosaurus neglectus
Euornithopoda indet.
Hadrosauridae
Edmontosaurus regalis
Edmontosaurus annectens (including
Anatotitan copei)
Edmontosaurus sp.
Hadrosauridae indet.
Pachycephalosauria
Pachycephalosauridae
Pachycephalosaurus wyomingensis
Pachycephalosaurus sp.
Stegoceras sp.
Stygimoloch sp.
Pachycephalosauria indet.
Ceratopsia
Ceratopsidae
Triceratops horridus
Triceratops sp.
Torosaurus latus
Age: late Maastrichtian (Cobban and
Reeside 1952)
3.16.
NEBRASKA
,
UNITED STATES
1. Niobrara Chalk Formation (Stidman,
pers. comm.) (not plotted)
Theropoda
Aves
Ornithurae
Apatornis celer
Age: Coniacian-early Campanian
(Hattin 1982)
2. Pierre Shale Formation (Stidman,
pers. comm.) (not plotted)
Theropoda
Aves
Hesperornithiformes
Hesperornis regalis
Age: early Campanian (Gill and Cobban
1966)
3.17.
MISSOURI
,
UNITED STATES
1. Ripley Formation (Gilmore and Stewart
1945; Horner 1979)
Ornithopoda
Hadrosauridae indet. (=Neosaurus
missouriensis)
Age: ?early Maastrichtian (Horner 1979)
3.22.
NEW JERSEY
,
UNITED STATES
1. Raritan Formation (Baird 1989;
Gallagher 1993; Gallagher et al. 1999)
Theropoda
Avialae indet.
?Theropoda indet.
Euornithopod tracks
Age: Cenomanian (Olsson 1980)
2. Merchantville Formation (Gallagher
1993)
Ornithopoda
Hadrosauridae
Hadrosaurus foulkii
Hadrosauridae indet.
Age: early Campanian (Owens 1983)
3. Woodbury Formation (Gallagher 1984,
1993)
Ornithopoda
Hadrosauridae
Hadrosaurus foulkii
Hadrosauridae indet.
Age: late early Campanian (Owens 1983)
4. Marshalltown Formation (Gallagher
1993)
Theropoda
Coelurosauria
Dryptosaurus aquilunguis
Ornithopoda
Hadrosauridae
Hadrosaurus foulkii
Hadrosauridae indet. (including
Hypsibema sp.)
Age: late Campanian (Owens 1983)
5. Mount Laurel Formation (including
Wenonah Formation) (Baird and Horner
1977; Gallagher 1993)
Theropoda
Tyrannosauroidea
Dryptosaurus aquilunguis
Ornithomimosauria indet.
(=Coelosaurus antiquus)
Ankylosauria
Nodosauridae indet.
Ornithopoda
Hadrosauridae indet.
Age: late Campanian–early
Maastrichtian (Owens 1983)
6. Navesink Formation (including New
Egypt Formation) (Olson and Parris
1987; Gallagher 1993)
Theropoda
Tyrannosauroidea
Dryptosaurus aquilunguis
Tyrannosauroidea indet. (including
Laelaps macropus, Tomodon
horrificus)
Ornithomimosauria indet.
(=Coelosaurus antiquus)
Ankylosauria
Nodosauridae indet.
Ornithopoda
Hadrosauridae
Unnamed hadrosaurine
(=“Hadrosaurus” minor)
Hadrosauridae indet. (including
Hadrosaurus cavatus,
H. minor)
Age: late Campanian–early
Maastrichtian (Owens 1983)
7. Hornerstown Formation (Olson and
Parris 1987)
Theropoda
Avialae
Aves
Procellariiformes
Tytthostonyx glauconiticus
Gastornithiformes
Graculavus velox
Telmatornis priscus
Anatalavis rex
Laornis edwardsianus
Palaeotringa littoralis
Palaeotringa vagans
Age: late Maastrichtian–early Paleocene
(Koch and Olsson 1974)
3.24.
DELAWARE
,
UNITED STATES
1. Merchantville Formation (Gallagher
1993)
Theropoda
Ornithomimidae indet.
Ornithopoda
Hadrosauridae indet.
Age: early Campanian (Owens 1983)
2. Marshalltown Formation (Gallagher
1993)
Theropoda
Ornithomimidae indet.
Ornithopoda
Hadrosauridae indet.
586 DINOSAUR DISTRIBUTION AND BIOLOGY

Age: late Campanian (Owens 1983)
3. Mount Laurel Formation (Baird and
Galton 1981; Gallagher 1993)
Ornithopoda
Hadrosauridae indet.
Age: late Campanian–early
Maastrichtian (Owens 1983)
3.25.
MARYLAND
,
UNITED STATES
1. Severn Formation (Baird 1986b)
Theropoda
Ornithomimosauria indet.
(=Coelosaurus antiquus)
Ornithopoda
Hadrosauridae indet.
Age: Maastrichtian (Baird 1986b)
3.27.
NORTH CAROLINA
,
UNITED STATES
1. Black Creek Formation (Baird and
Horner 1979)
Theropoda
Ornithomimosauria indet.
(=Coelosaurus cf. antiquus)
Theropoda indet. (=?Dryptosaurus sp.)
Ornithopoda
Hadrosauridae
?Lophorhothon atopus
Hadrosauridae indet. (=Hypsibema
crassicauda)
Age: Campanian (Brett and Wheeler
1961)
3.28.
SOUTH CAROLINA
,
UNITED STATES
1. Donoho Creek Formation (Weishampel
and Young 1996; Knight pers. comm.)
Theropoda indet.
Ornithopoda
Hadrosauridae indet.
Age: early Maastrichtian (Sohl and
Owens 1991)
3.29.
GEORGIA
,
UNITED STATES
1. Blufftown Formation (Reinhardt et al.
1981; Schwimmer and Best 1989;
Schwimmer et al. 1993)
Theropoda
Tyrannosauridae indet.
Ornithomimidae indet.
Ornithopoda
Hadrosauridae indet.
Age: early Campanian (Owens 1983)
3.30.
ALABAMA
,
UNITED STATES
1. Upper Eutaw Formation (Lamb 1996,
pers. comm.)
Theropoda indet.
Ankylosauria
?Ankylosauridae indet.
Ornithopoda
?Hadrosauridae indet.
Age: late Santonian–early Campanian
(Lamb, pers. comm.)
2. Mooreville Chalk Formation (Lull and
Wright 1942; Langston 1960a; Olson
1975; Dobie 1978; Lamb 1993, 1996,
1997, 1998, 2001; Chiappe et al. 2002a;
Carpenter, pers. comm.; Clarke, pers.
comm.)
Theropoda
Ornithomimosauria indet.
(=Coelosaurus cf. antiquus)
Avialae
Enantiornithes
Halimornis thompsoni
Ichthyornithiformes
Ichthyornis dispar (=Angelinornis
antecessor)
Theropoda indet.
Ankylosauria
Nodosauridae indet.
Ornithopoda
?Hadrosauridae
Lophorhothon atopus
?Hadrosauridae indet.
Dinosaur eggs
Age: late Santonian–early Campanian
(Mancini et al. 1984;
Puckett 1994)
3. Demopolis Chalk Formation (Lamb
1989; Carr et al., in press)
Theropoda
Unnamed tyrannosauroidean
?Tyrannosauridae indet.
Age: middle Campanian (Puckett 1994)
4. Ripley Formation (Lamb, pers. comm.)
Ornithopoda
?Hadrosauridae indet.
Age: early Maastrichtian (Lamb,
pers. comm.)
3.31.
MISSISSIPPI
,
UNITED STATES
1. Basal Eutaw or Upper McShan
Formation (Carpenter 1982c)
Dinosauria indet.
Theropoda
Ornithomimidae indet.
Ornithopoda
Hadrosauridae indet.
Age: late Coniacian–?early Santonian
(Carpenter 1982c)
2. Eutaw Formation (Kaye and Russell
1973)
Ornithopoda
Hadrosauridae indet.
Age: late Santonian (Kaye and Russell
1973)
3. Selma Group (Lull and Wright 1942;
Carpenter 1982c)
Ornithopoda
Hadrosauridae indet.
Age: Campanian (Carpenter 1982c)
3.32.
TENNESSEE
,
UNITED STATES
1. Demopolis Shale (Bryan et al. 1991)
Ornithopoda
Hadrosauridae indet.
Age: middle Campanian (Puckett 1994)
3.33.
ALASKA
,
UNITED STATES
1. Lower Matanuska Formation (Gangloff
1995, 1998; Pasch and May 1997, 2001)
Ornithopoda
Hadrosauridae indet.
Age: Turonian (Gangloff 1998)
2. Upper Matanuska Formation (Gangloff
1995, 1998; Pasch and May 1997)
Ankylosauria
Nodosauridae
Unnamed Edmontonia species
Age: late Campanian–early
Maastrichtian (Gangloff
1995); Turonian (Pasch and
May 1997)
3. Prince Creek Formation (Brouwers et al.
1987; Parrish et al. 1987; Clemens and
Nelms 1993; Upper Colville Group,
Davies 1987; Gangloff 1998; May and
Gangloff 1999; Fiorillo and Gangloff
2000)
Theropoda
Tyrannosauridae indet.
Troodontidae
Troodon formosus
Dromaeosauridae
Dromaeosaurus albertensis
Saurornitholestes langstoni
Theropoda indet.
Ornithopoda
Euornithopoda
cf. Thescelosaurus sp.
Undescribed hadrosaurid
Hadrosauridae indet.
Ceratopsia
Ceratopsidae
Pachyrhinosaurus cf. canadensis
?Anchiceratops sp.
?Brachyceratops sp.
Ceratopsidae indet.
Age: middle Maastrichtian (Frederiksen
1986); early middle
Maastrichtian (Brouwers
et al. 1987)
4.4.
ESTADO DE PUEBLA
,
MEXICO
1. Mexcala Formation (Bravo-Cuevas and
Jiménez-Hidalgo 1996)
Theropod tracks
Euornithopod tracks
Age: Campanian-Maastrichtian (Bravo-
Cuevas, V. M., and Jiménez-
Hidalgo 1996)
4.5.
ESTADO DE BAJA CALIFORNIA NORTE
,
MEXICO
1. “El Gallo” formation (Morris 1971,
1973a, 1981; Carpenter and Alf 1994;
Rodriguez de la Rosa 1998)
Theropoda
Tyrannosauridae indet.
Ornithomimidae indet.
Troodontidae indet.
Troodon formosus
Dromaeosauridae
Saurornitholestes sp.
Dromaeosauridae indet.
Ankylosauria
Ankylosauria indet.
Nodosauridae
cf. Euoplocephalus sp.
Ornithopoda
Hadrosauridae
?Lambeosaurus laticaudus
?Lambeosaurus sp.
Hadrosaurinae indet.
DINOSAUR DISTRIBUTION 587

Ceratopsia
Ceratopsidae indet.
Dinosaur eggs
Age: Campanian (Morris 1981)
2. “La Bocana Roja” formation (Molnar
1974; Brodkorb 1976)
Theropoda
?Tyrannosauroidea
Labocania anomala
?Tyrannosauridae indet.
Avialae
Enantiornithiformes
Alexornis antecedens
Ornithopoda
Hadrosauridae indet.
Age: ?Campanian (Molnar 1974)
4.6.
ESTADO DE SONORA
,
MEXICO
1. Snake Ridge Formation (Lull and Wright
1942)
Ornithopoda
Hadrosauridae indet.
Age: ?Maastrichtian (unconfirmed)
2. Corral de Enmedio Formation
(Taliaferro 1933; Lucas and González-
Leon 1996)
Theropoda
Tyrannosauridae indet.
Ornithopoda
Hadrosauridae indet. (=Trachodon
mirabilis)
Ceratopsia
Ceratopsidae indet.
Age: late Campanian–Maastrichtian
(Gonzáles-Leon and Lucas
1996)
3. Camas Formation (Taliaferro 1933;
Gonzáles-Leon and Lucas 1996
Theropoda
Tyrannosauridae indet.
Ornithopoda
Hadrosauridae indet.
Age: late Campanian–Maastrichtian
(Gonzáles-Leon and Lucas
1996)
4. Packard Formation (Taliaferro 1933;
Gonzáles-Leon and Lucas 1996)
Ornithopoda
Hadrosauridae indet.
Age: late Campanian–Maastrichtian
(Gonzáles-Leon and Lucas
1996)
5. Lomas Coloradas Formation (Taliaferro
1933; Gonzáles-Leon and Lucas 1996)
Theropoda
Tyrannosauridae indet.
Ornithopoda
Hadrosauridae indet.
Ceratopsia
Ceratopsidae indet.
Age: late Campanian–Maastrichtian
(Gonzáles-Leon and Lucas
1996)
4.7.
ESTADO DE CHIHUAHUA
,
MEXICO
1. Aguja Formation (Westgate, pers.
comm.)
Theropoda
Tyrannosauridae indet.
Maniraptora indet.
Ornithopoda
Hadrosauridae
Kritosaurus sp.
Ceratopsia
Ceratopsidae
?Chasmosaurus sp.
Age: Campanian (Westgate, pers.
comm.)
2. Ojinaga Formation (Rodriguez de la
Rosa, pers. comm.)
Ceratopsia
Ceratopsidae indet.
Age: Late Cretaceous (Rodriguez de la
Rosa, pers. comm.)
4.8.
ESTADO DE COAHUILA
,
MEXICO
1. Palau Formation (Rodriguez de la Rosa,
pers. comm.)
Ceratopsia
Ceratopsidae indet.
Age: Late Cretaceous (Rodriguez de la
Rosa, pers. comm.)
2. Cerro del Pueblo Formation (Murray
et al. 1960; Hernández et al. 1996;
Rodriguez de la Rosa and Cevallos-
Ferriz 1998; Seranno Branas and
Hernandez-Rivera 1998; Aguillon
et al. 1999; Hernandez-Rivera and
Delgado de Jesus 1999; Rodriguez
de la Rosa 2001)
Theropoda
Tyrannosauridae indet.
Ornithomimidae indet.
?Oviraptoridae indet.
?Troodontidae indet.
Dromaeosauridae indet.
Avialan tracks
Ankylosauria
Ankylosauridae indet.
Ornithopoda
Hadrosauridae
Hadrosaurinae indet.
(?=Gryposaurus sp.)
Lambeosaurinae indet.
Euornithopod tracks
Ceratopsia
Ceratopsidae
?Chasmosaurus sp.
Ceratopsidae indet. (including
Monoclonius sp.)
Dinosaur coprolites
Age: late Campanian (Hernández et al.
1996)
3. Soledad Beds (Janensch 1926; Lull 1933)
Ceratopsia indet.
Age: late Maastrichtian (=Laramie
Formation; Janensch 1926)
5.1.
DEPARTEMENTO DO COMAYAGUA
,
HONDURAS
1. Lower Valle de Angeles Redbeds (Horne
et al. 1974; Finch 1981; Horne 1994)
Ornithopoda
Hadrosauridae indet.
Age: Cenomanian (Horne 1994)
588 DINOSAUR DISTRIBUTION AND BIOLOGY
Europe
European Late Cretaceous dinosaurs are becoming increasingly
well known (84 locations, amounting to a 110% increase). Areas
in southern France, northern Spain, eastern Austria, The Nether-
lands, and the Transylvania region of Romania have produced
remarkable dinosaur faunas thanks to the early work of F. Nopcsa,
P. Matheron, L. Dollo, A. F. de Lapparent, E. Bunzel, and H. G.
Seeley. More recently, research in France by E. Buffetaut, P. Taquet,
P. Kerourio, B. Sigé, J. Le Loeuff, and Y. Laurent, among others,
has made this region one of the best known for its Late Cretaceous
dinosaur faunas. In The Netherlands, A. W. F. Meijer, E. W. A.
Mulder, and collaborators have extended our knowledge of the
dinosaurs from the type localities of the Maastrichtian Stage.
Knowledge of the Late Cretaceous dinosaurs of Spain, one of the
most promising European areas for new discoveries, has been
increased through the important faunistic work of M. L. Casa-
novas Cladellas, J. V. Santafé-Llopis, J. L. Sanz, C. Pol, X. Pereda-
Suberbiola, A. Buscalioni, J. J. Moratalla, W. Brinkmann, A.Schulp,
L. Ardévol, J. Company, and A. Galobart. A preliminary account
of new hadrosaurid material from the Late Cretaceous of north-
ern Italy has been reported by F. M. Dalla Vecchia. Finally, study
of the important Romanian dinosaur fauna has been recently
renewed by D. Grigorescu, Z. Csiki, V. Codrea, C.-M. Jianu, and
D. B. Weishampel. All these Late Cretaceous faunas of Europe
are indicated in figures 24.5 and 24.19.
6.2.
SCANIA
,
SWEDEN
1. Unnamed unit (Persson 1959)
Theropoda indet.
Age: early Campanian (Persson 1959)
2. Unnamed unit (Nessov 1993)
Ornithopoda
Euornithopoda indet.
Age: early Campanian (Nessov 1993)
3. Unnamed unit (Nessov and Yarkov 1993;
Kurochkin 2000)
Theropoda
Avialae

?Hesperornithiformes indet.
(=Hesperornis rossicus)
Age: early Campanian (Nessov and
Yarkov 1993)
10.17.
ISLE OF WIGHT
,
ENGLAND
1. Upper Greensand (Lydekker 1888b;
Le Loeuff 1993) (not plotted)
Sauropoda indet. (=Titanosaurus
lydekkeri)
Age: early Cenomanian (Rawson et al.
1978)
10.21.
KENT
,
ENGLAND
1. Upper Greensand (=Chalk Marl; Huxley
1867; Pereda-Suberbiola 1993a; Pereda-
Suberbiola and Barrett 1999)
Ankylosauria
Nodosauridae indet. (=Acanthopholis
horridus)
Age: early Cenomanian (Rawson et al.
1978)
10.22.
HERTFORDSHIRE
,
ENGLAND
1. Lower Chalk (=Chalk Marl; Newton 1892)
Ornithopoda
Hadrosauridae indet. (=Iguanodon hilli)
Age: early Cenomanian (Rawson et al.
1978)
11.3.
PROVINCE DE NAMUR
,
BELGIUM
1. Glauconie argileuse (Dollo 1883c, 1903)
Theropoda indet. (=Megalosaurus
lonzeensis)
Ornithopoda
Iguanodontia
Craspedodon lonzeensis
Age: Santonian (Taquet 1976)
11.4.
PROVINCE DE LIMBURG
,
BELGIUM
1. Maastricht Formation (Buffetaut et al.
1985b; Mulder et al. 1997, 1998)
Ornithopoda
Hadrosauridae indet. (including
Orthomerus dolloi)
Age: late Maastrichtian (Felder 1975a,
1975b)
12.1.
PROVINCIE LIMBURG
,
THE NETHERLANDS
1. Maastricht Formation (Seeley 1883;
Buffetaut et al. 1985b; Mulder et al.
1998; Jagt et al. 1999; Weishampel et al.
1999)
Theropoda
Tetanurae indet. (=Megalosaurus
bredai)
Ornithopoda
Hadrosauridae
Euhadrosauria indet.
Hadrosauridae indet. (including
Orthomerus dolloi)
Age: late Maastrichtian (Felder 1975a,
1975b)
14.1.
DÉPARTEMENT DE LA VENDÉE
,
FRANCE
1. Unnamed unit (“Grès calcaires”; Buffetaut
et al. 1991; Buffetaut and Pouit 1994)
Theropoda indet.
Age: Turonian (Buffetaut et al. 1991)
2. Unnamed unit (Buffetaut et al. 1991)
Theropoda indet.
Age: Santonian (Buffetaut and Pouit
1994)
14.7.
DÉPARTEMENT DU GARD
,
FRANCE
1. Unnamed unit (Gervais 1859; Dumas
1876; Buffetaut et al. 1991)
Theropoda indet. (=Megalosaurus sp.)
Age: Cenomanian (Buffetaut et al. 1991)
2. Unnamed unit (Buffetaut et al. 1986;
Sigé et al. 1997)
Theropoda
?Dromaeosauridae indet.
Sauropoda
Lithostrotia indet. (=Titanosauridae
indet.)
Ornithopoda
Euornithopoda
cf. Rhabdodon priscus
Age: Maastrichtian (Buffetaut et al.
1986)
3. Unnamed unit (Buffetaut et al. 1986)
Theropoda
?Dromaeosauridae indet.
Age: late Cretaceous (Buffetaut et al.
1986)
14.8.
DÉPARTEMENT DU VAR
,
FRANCE
1. Unnamed unit (Le Loeuff and Buffetaut
1991 )
Theropoda
Ceratosauria indet. (=Tarascosaurus
salluvicus)
Age: early Campanian (Buffetaut et al.
1991, 1997a; Le Loeuff and
Buffetaut 1991)
2. Grès à Reptiles Formation (Lapparent
1947; Le Loeuff et al. 1992; Buffetaut
et al. 1995b, 2000a; Le Loeuff 1998)
Theropoda
?Abelisauridae indet.
?Dromaeosauridae
Variraptor mechinorum
Avialae
Enantiornithes indet.
Ornithurae
Gargantuavis philoinos
Theropoda indet. (=Megalosaurus
pannoniensis, Avialae indet.)
Sauropoda
Sauropoda indet. (=Hypselosaurus
priscus, Titanosaurus indicus)
Ankylosauria indet.
Ornithopoda
Euornithopoda
Rhabdodon priscus
Age: early Maastrichtian (=Rognacien;
Lapparent 1947; Westphal
and Durand 1990)
3. Unnamed unit (Kérourio1982; Williams
et al. 1984)
Dinosaur eggs
Age: early Maastrichtian (Williams et al.
198 4)
4. Unnamed unit (Williams et al. 1984)
Dinosaur eggs
Age: late Maastrichtian (Williams et al.
198 4)
14.12.
DÉPARTEMENT DE L
’
HÉRAULT
,
FRANCE
1. Unnamed unit (“Fuvélien”; Le Loeuff
1992, 1998; Buffetaut et al. 1996b,
1997a; Garcia and Pereda-Suberbiola
2003)
Theropoda
Dromaeosauridae indet.
Ankylosauria
Nodosauridae
Struthiosaurus languedocensis
Nodosauridae indet.
Ornithopoda
Euornithopoda
cf. Rhabdodon priscus
Age: early Campanian (Buffetaut et al.
1996b, 1997a)
2. Grès de Saint-Chinian (Lapparent 1947;
Buffetaut et al. 1986, 1999; Buffetaut
and Le Loeuff 1991b; Vianey-Liaud and
Crochet 1993; Garcia et al. 2000)
Theropoda
?Abelisauridae indet.
?Dromaeosauridae
Variraptor mechinorum
Avialae
Enanthiornithes indet.
Theropoda indet. (=Megalosaurus
pannoniensis)
Sauropoda
Lithostrotia
Ampelosaurus atacis
Ankylosauria indet. (=Rhodanosaurus
lugdunensis)
Ornithopoda
Euornithopoda
Rhabdodon septimanicus
Dinosaur eggs
Age: late Campanian–early
Maastrichtian (Le Loeuff,
unpubl. data)
3. Unnamed unit (Garcia et al. 2000)
Dinosaur eggs
Age: Maastrichtian (Garcia et al. 2000)
14.13.
DÉPARTEMENT DE LA DORDOGNE
,
FRANCE
1. Unnamed unit (=Lignites du Sarladais;
Arnaud 1866; Buffetaut et al. 1991)
Dinosauria indet. (=Megalosaurus sp.)
Age: Cenomanian (Arnaud 1866;
Buffetaut et al. 1991)
2. Craie tufau (Gervais 1852, 1859;
Buffetaut et al. 1991; Laurent et al.
2001)
Sauropoda indet. (=?Aepysaurus sp.)
Age: Campanian (Buffetaut et al. 1991;
Le Loeuff 1992, 1993;
Laurent et al. 2001)
3. Unnamed unit (Platel 1989; Buffetaut et
al. 1991; Le Loeuff 1992, 1993, 1998)
Sauropoda
Lithostrotia indet. (=Titanosauridae
indet.)
Age: late Campanian (Buffetaut et al.
1991 )
DINOSAUR DISTRIBUTION 589

14.16.
DÉPARTEMENT DE L
’
ORNE
,
FRANCE
1. Craie chloritée (Morière 1877; Bigot
1904; Buffetaut et al. 1991)
?Dinosauria indet. (doubtfully
dinosaurian)
Age: Cenomanian (Buffetaut et al. 1991)
14.19.
DÉPARTEMENT DE LA CHARENTE
-
MARITIME
,
FRANCE
1. Unnamed unit (Lapparent 1967;
Buffetaut et al. 1991)
Sauropoda indet. (=Megalosaurus sp.)
Age: Cenomanian (Lapparent 1967)
2. Sables de Lussant (Buffetaut et al. 1991;
Buffetaut and Pouit 1994)
Theropoda indet.
Sauropoda
?Lithostrotia (=?Titanosauridae
indet.)
Age: Cenomanian (Buffetaut et al. 1991)
14.23.
DÉPARTEMENT DE LA SARTHE
,
FRANCE
1. Unnamed unit (Buffetaut 1989a)
Sauropoda
Lithostrotia indet. (=Titanosauridae
indet.)
Age: Cenomanian (Buffetaut 1989a;
Buffetaut et al. 1991)
14.24.
DÉPARTEMENT DE LA VIENNE
,
FRANCE
1. Unnamed unit (Buffetaut and Pouit
1994)
Theropoda indet.
Age: Cenomanian (Buffetaut and Pouit
1994; reworked into
Miocene)
14.31.
DÉPARTEMENT DU VAUCLUSE
,
FRANCE
1. Unnamed unit (Depéret 1894; Buffetaut
et al. 1991)
Sauropoda indet.
Age: early Cenomanian (Buffetaut et al.
1991 )
14.33.
DÉPARTEMENT DU MAINE
-
ET
-
LOIRE
,
FRANCE
1. Unnamed unit (Buffetaut et al. 1980)
Theropoda indet.
Ornithopoda
Iguanodontia indet.
Age: Cenomanian (Buffetaut et al. 1980;
reworked into Miocene)
2. Sables de Brézé (Buffetaut and Pouit
1994)
Theropoda indet.
Age: Cenomanian (Buffetaut and Pouit
1994)
14.34.
DÉPARTEMENT DES LANDES
,
FRANCE
1. Unnamed unit (Laurent et al. 2001)
Sauropoda
Lithostrotia indet. (=Titanosauridae
indet.)
Age: late Campanian (Laurent et al.
2001)
14.35.
DÉPARTEMENT DE LA HAUTE
-
GARONNE
,
FRANCE
1. Calcarintes du Jadet Formation
(=Calcaires nankin Formation; Paris
and Taquet 1973; Le Loeuff et al. 1994a;
Laurent 1996; Laurent et al. 1999, 2001,
2002)
Ornithopoda
Hadrosauridae indet.
Age: late Maastrichtian (Buffetaut et al.
1991; Laurent et al. 2001,
2002)
2. Lestaillats Marls Formation (Laurent
et al. 1999, 2001, 2002)
Theropoda.
?Dromaeosauridae indet.
Theropoda indet.
Ankylosauria
Nodosauridae indet.
Ornithopoda
Hadrosauridae indet.
Age: late Maastrichtian (Laurent et al.
1999)
3. Marnes d’Auzas Formation (Le Loeuff
et al. 1994a; Laurent 1996; Gheerbrant
et al. 1997)
Theropoda indet.
Ankylosauria indet.
Ornithopoda
Hadrosauridae indet.
Age: late Maastrichtian (Le Loeuff et al.
1994a)
14.36.
DÉPARTEMENT DE L
’
ARIÈGE
,
FRANCE
1. Marnes d’Auzas (Le Loeuff 1991b;
Le Loeuff et al. 1994a; Laurent 1997)
Undescribed theropod
Ornithopoda
Hadrosauridae indet.
Age: late Maastrichtian (Laurent 1997)
2. Grès de Labarre Formation (Villatte et al.
1985; Le Loeuff 1991b, 1998)
Sauropoda
Lithostrotia
cf. Ampelosaurus atacis
Ankylosauria
Nodosauridae indet.
Ornithopoda
Euornithopoda
Rhabdodon priscus
Age: Maastrichtian (Villatte et al. 1985)
14.37.
DÉPARTEMENT DE L
’
AUDE
,
FRANCE
1. Grès d’Alet Formation (Le Loeuff,
unpubl. data)
Dinosaur eggs
Age: Campanian (Bilotte 1985)
2. Grès de Saint-Chinian Formation (Le
Loeuff, unpubl. data)
Theropoda
?Dromaeosauridae
Variraptor mechinorum
Ornithopoda
Euornithopoda
Rhabdodon cf. priscus
Age: late Campanian–early
Maastrichtian (Le Loeuff,
unpubl. data)
3. Marnes Rouges Inférieures Formation
(Lapparent and Lavocat 1955; Lapparent
1967; Clottes and Raynaud 1983;
Beetschen 1985; Breton et al. 1986;
Buffetaut et al. 1997a; Cousin et al.
1989; Le Loeuff 1991a, 1992, 1993,
1995, 1998; Le Loeuff et al. 1994a,
1994b, 1994c)
Theropoda
Dromaeosauridae indet.
Avialae
Ornithurae
Gargantuavis philoinos
Theropoda indet.
Sauropoda
Lithostrotia
Ampelosaurus atacis
Ankylosauria indet.
Ornithopoda
Euornithopoda
cf. Rhabdodon priscus
Dinosaur eggs
Age: early Maastrichtian (Bessiere et al.
1989)
4. Marnes rouges de Roquelongue
Formation (Le Loeuff et al. 1993;
Buffetaut et al. 1997a; Laurent et al.
1997; Le Loeuff 1998)
Theropoda
Dromaeosauridae indet.
Ornithopoda
Hadrosauridae indet.
Dinosaur eggs
Age: late Maastrichtian-Danian
(Laurent et al. 1997)
5. Calcaires et argiles de Vignevieille
Formation (Lautier and Le Loeuff 1992)
Dinosaur eggs
Age: late Maastrichtian (Bilotte 1985)
14.38.
DÉPARTEMENT DES BOUCHES
-
DU
-
RHÔNE
,
FRANCE
1. Grès à Reptiles Formation (Lapparent
1947; Buffetaut et al. 1986; Le Loeuff
et al. 1992; Le Loeuff and Buffetaut
1998; Allain and Taquet 2000)
Theropoda
?Abelisauridae indet.
?Dromaeosauridae
Variraptor mechinorum
Pyroraptor olympius
Theropoda indet.
Sauropoda
Sauropoda indet. (=Hypselosaurus
priscus)
Ankylosauria indet. (=Rhodanosaurus
lugdunensis)
Ornithopoda
Euornithopoda
Rhabdodon priscus
Age: early Maastrichtian (Westphal and
Durand 1990)
2. Unnamed unit (Kérourio 1982)
Dinosaur eggs
Age: late Campanian (Kérourio 1982)
3. Unnamed unit (Penner 1985)
Dinosaur eggs
Age: early Maastrichtian (Vianey-Liaud
et al. 1994)
590 DINOSAUR DISTRIBUTION AND BIOLOGY

4. Unnamed unit (Kérourio 1981b)
Dinosaur eggs
Age: early Maastrichtian (Kérourio
1981b)
5. Unnamed unit (Kérourio 1981b, 1982;
Williams et al. 1984; Cousin et al. 1987)
Dinosaur eggs
Age: late Maastrichtian (Williams et al.
198 4)
15.1.
PROVINCIA DE SORIA
,
SPAIN
1. Unnamed unit (Sanz et al. 1992)
Dinosaur eggs
Age: Maastrichtian (Sanz et al. 1992)
15.3.
PROVINCIA DE VALENCIA
,
SPAIN
1. Unnamed unit (Sanz 1986)
Ankylosauria
Nodosauridae indet.
Age: Santonian-Maastrichtian (Sanz 1986)
2. Sierra Perenchiza Formation (Company
et al. 1998)
Ornithopoda
Hadrosauridae
Euhadrosauria indet.
Age: Campanian-Maastrichtian
(Gutierrez et al. 1984)
3. Unnamed unit (Company, pers. comm.)
Theropoda indet.
Sauropoda
Lithostrotia indet. (=Titanosauridae
indet.)
Ankylosauria
Nodosauridae indet.
Ornithopoda
Euornithopoda
Rhabdodon sp.
Dinosaur eggs
Age: Campanian-Maastrichtian
(Company, pers. comm.)
4. Unnamed unit (Moratalla and Frances
1991 )
Dinosaur eggs
Age: Maastrichtian (Moratalla and
Frances 1991)
5. Unnamed unit (Casanovas-Cladellas
1992)
Ornithopoda
Hadrosauridae indet.
Age: Late Cretaceous (Casanovas-
Cladellas 1992)
6. Unnamed unit (Casanovas-Cladellas
1992)
Undescribed sauropod
Age: Late Cretaceous (Casanovas-
Cladellas 1992)
15.4.
PROVINCIA DE TERUEL
,
SPAIN
1. Unnamed unit (Cuenca-Bescós et al.
1999b)
Theropods tracks
Age: Cenomanian (Cuenca-Bescós et al.
1999a)
15.6.
PROVINCIA DE BURGOS
,
SPAIN
1. Unit S3U1 (Sanz 1986; Astibia et al. 1987,
1990; Pereda-Suberbiola 1993, 1999;
Pereda-Suberbiola and Astibia 1993;
Pereda-Suberbiola et al. 1995, 2000;
Le Loeuff 1998; Pereda-Suberbiola and
Sanz 1999; Sanz et al. 1999)
Theropoda
?Abelisauridae indet.
Ornithomimosauria indet.
?Troodontidae indet.
(=cf. Euronychodon sp.)
Dromaeosauridae indet.
(several species)
Sauropoda
Lithostrotia
Lirainasaurus astibiae
Lithostrotia indet.
(=Titanosauridae indet.)
Ankylosauria
Nodosauridae
Struthiosaurus sp.
Nodosauridae indet.
Ornithopoda
Euornithopoda
Rhabdodon sp.
Hadrosauridae indet.
Euornithopoda indet.
Age: late Campanian (Pereda-Suberbiola
et al. 2000)
2. Unnamed unit (Pereda-Suberbiola et al.
1999)
Sauropoda
Titanosauridae indet.
Ankylosauria indet.
Ornithopoda indet.
Age: late Campanian-early
Maastrichtian (Pereda-
Suberbiola et al. 1999)
3. Calizas de Lychnus Formation
(Moratalla 1990; Moratalla and Frances
1991; Pol et al. 1992)
Theropoda
?Troodontidae indet. (=cf.
Paronychodon sp.)
Dromaeosauridae indet.
Ankylosauria indet.
Dinosaur eggs
Age: Maastrichtian (Pol et al. 1992)
15.8.
PROVINCIA DE LLEIDA
,
SPAIN
1. Aren Formation (Lapparent and Aguirre
1956a, 1956b; Bataller 1958, 1960;
Lapparent 1958, 1959; Erben et al. 1979;
Llompart and Kraus 1982; Brinkmann
1984; Casanovas-Cladellas et al. 1984,
1985a, 1985b, 1986, 1987, 1988, 1993d,
1995d; Llompart et al. 1984; Kohring
1989; Casanovas-Cladellas 1992; Le
Loeuff 1993; Ardévol et al. 1995; Sanz
et al. 1995; Sander et al. 1998; Priéto-
Marquez et al. 2000)
Theropoda
Coelurosauria incertae sedis
cf. Ricardoestesia sp.
Theropoda indet. (=Megalosaurus sp.)
Theropod tracks
Sauropoda
Sauropoda indet. (including
Hypselosaurus sp.,
Titanosaurus cf. indicus)
Sauropod tracks
Ornithopoda
Euornithopoda
Rhabdodon priscus
Iguanodontia indet.
Hadrosauridae
Pararhabdodon isonensis
Hadrosauridae indet.
(=Orthomerus sp.)
Euornithopod tracks
Dinosaur eggs
Age: Maastrichtian (Senz 1998)
2. Lower Tremp Formation (Llompart 1979;
Casanovas-Cladellas 1992; Casanovas-
Cladellas and Santafé-Llopis 1993;
Casanovas Cladellas et al. 1993b; Vianey-
Liaud and López-Martínez 1997; López-
Martínez et al. 1998a, 1998b, 1999;
Sander et al. 1998; López-Martínez
1999; Martinell et al. 2001)
Sauropoda
Sauropoda indet. (including
Hypselosaurus sp.,
Titanosaurus indicus)
Sauropod tracks (may be feeding
traces of rays)
Ornithopoda
Euornithopoda
Rhabdodon priscus
Hadrosauridae
Euhadrosauria indet.
Euornithopod tracks
Dinosaur eggs
Age: late Maastrichtian (Galbrun et al.
1993)
3. Unnamed unit (Le Loeuff 1998)
Sauropoda
Lithostrotia indet. (=Titanosauridae
indet.)
Age: late Maastrichtian (Le Loeuff 1998)
4. Unnamed unit (Casanovas-Cladellas
1992)
Sauropoda
Lithostrotia indet. (=Titanosauridae
indet.)
Ornithopoda
Hadrosauridae indet.
Age: Late Cretaceous (Casanovas-
Cladellas 1992)
5. Unnamed unit (Casanovas-Cladellas
et al. 1988)
Theropoda
Tetanurae indet. (=Megalosauridae
indet.)
Age: Late Cretaceous (Casanovas-
Cladellas et al. 1988)
15.9.
PROVINCIA DE CUENCA
,
SPAIN
1. Villalba de la Sierra Formation
(Gutierrez and Robles 1979; Moratalla
and Frances 1991)
Dinosaur eggs
Age: Maastrichtian (Moratalla 1992)
15.11.
PROVINCIA DE SEGOVIA
,
SPAIN
1. Unnamed unit (Sanz 1986; Sanz and
Buscalioni 1987; Buscalioni and
Martínez -Salanova 1990)
Theropoda indet.
Sauropoda
Lithostrotia indet. (=Titanosauridae
indet.)
DINOSAUR DISTRIBUTION 591

Sauropoda indet.
Ornithischia indet.
Ornithopoda
Euornithopoda
Rhabdodon sp.
Age: Campanian-Maastrichtian (Sanz
and Buscalioni 1987)
15.12.
PROVINCIA DE HUESCA
,
SPAIN
1. Tremp Formation (López-Martínez et al.
2001; Cuenca-Bescós and Canudo, pers.
comm.)
Theropoda
Coelurosauria indet.
?Troodontidae indet.
(=cf. Euronychodon sp.)
Dromaeosauridae indet. (two species)
Sauropoda indet.
Ornithopoda
Euhadrosauria indet.
Hadrosauridae indet.
Dinosaur eggs
Age: late Maastrichtian (Cuenca-Bescós
and Canudo, pers. comm.)
15.13.
PROVINCIA DE BARCELONA
,
SPAIN
1. Tremp Formation (Le Loeuff and
Martínez -Rius 1997; Schulp and Brokx
1999)
Sauropod tracks
Age: Maastrichtian (Le Loeuff and -Rius
1997; Schulp and Brokx 1999)
16.3.
DISTRITO DO COIMBRA
,
PORTUGAL
1. Unnamed unit (Lapparent and
Zbyszewski 1957; Antunes and Pais
1978)
Theropoda
Coelurosauria indet.
Theropoda indet. (=?Megalosaurus
pannoniensis)
Age: late Campanian–?Maastrichtian
(Lapparent and Zbyszewski
1957; Antunes and Pais 1978)
2. Unnamed unit (Antunes and Sigogneau-
Russell 1991, 1992; Galton 1996)
Theropoda
Coelurosauria indet. (=A, B, C;
Antunes and Sigogneau-
Russell 1992)
?Troodontidae indet. (=Euronychodon
portucalensis)
Dromaeosauridae indet.
Theropoda indet. (=“Carnosauria”,
Galton 1996; Ther. indet.
A, B, Antunes and
Sigogneau-Russell 1992)
Sauropoda indet.
Ornithopoda indet. (=Taveirosaurus
costai)
Age: late Campanian–Maastrichtian
(Galton 1996)
17.3.
BAYERN
,
GERMANY
1. Gerhartsreiter Schichten (Wellnhofer
1994)
Ornithopoda
Hadrosauridae indet.
Age: late Maastrichtian (Wellnhofer 1994)
19.4.
REGIONE FRIULI
–
VENEZIA GIULIA
,
ITALY
1. Calcari di Aurisina (Brazzatti and
Calligaris 1995; Dalla Vecchia 1997a)
Ornithopoda
Unnamed hadrosaurid
Age: late Santonian (Tarlao et al. 1994;
Tentor et al. 1994)
19.8.
REGIONE PUGLIA
,
ITALY
1. Calcare di Altamura (Nicosia et al. 2000;
Dalla Vecchia 2001b)
?Sauropod tracks
?Ankylosaur tracks
Hadrosaurid tracks
Age: late Coniacian–early Santonian
(Dalla Vecchia 2001b)
23.1.
SEVEROCESKY KRAJ
,
CZECH REPUBLIC
1. Korycaner Formation (Fritsch 1905)
?Dinosauria indet. (=Iguanodon
exogirarus)
Age: Cenomanian (Fritsch 1905)
23.2.
VYCHODOCESKY KRAJ
,
CZECH
REPUBLIC
1. Preiesener Formation (Fritsch 1905)
?Dinosauria indet. (=Albisaurus scutifer)
Age: ?Coniacian-?Santonian (=?early
Senonian; Fritsch 1905)
24.1.
NIEDERÖSTERREICH
,
AUSTRIA
1. Gosau Formation (Bunzel 1871; Seeley
1881; Pereda-Suberbiola 1993b; Le
Loeuff 1998)
Theropoda indet. (=Megalosaurus
pannoniensis)
Ankylosauria
Nodosauridae
Struthiosaurus austriacus (including
Crataeomus lepidophorus,
C. pawlowitschii partim,
Danubiosaurus anceps partim,
Hoplosaurus ischyrus,
Leipsanosaurus noricus,
Pleuropeltis suessi partim)
Ankylosauria indet. (=Danubiosaurus
anceps partim)
?Ankylosauria indet. (=Rhadinosaurus
alcinus)
Ornithopoda
Euornithopoda
?Rhabdodon priscus (=Mochlodon
suessi)
Age: Campanian (Jeletzky 1960; Thenius
1974)
26.2.
VESZPRÉM
,
HUNGARY
1. Csehbány Formation (Ösi, in press)
Theropoda
Coelurosauria incertae sedis
Ricardoestesia cf. gilmorei
Maniraptora indet.
Ankylosauria
Unnamed nodosauridae
Ornithopoda
Euornithopoda
?Rhabdodon priscus
Age: late Santonian (Ösi, in press)
27.2.
OBCINA DIVACA
,
SLOVENIA
1. ?Liburnian Formation (Debeljak et al.
1999; Dalla Vecchia 2001b)
Theropoda
?Dromaeosauridae indet.
Theropoda indet.
Ornithopoda
Hadrosauridae indet.
Ornithopoda indet.
Age: early Campanian–late
Maastrichtian (Debeljak et
al. 1999); late Maastrichtian
(Dalla Vecchia 2001b)
28.1.
ISTARSKA ZUPANIJA
,
CROATIA
1. Unnamed unit (Gogala 1975; Leghissa
and Leonardi 1990; Dalla Vecchia et al.
1993; Dalla Vecchia and Tarlao 1995;
Dalla Vecchia and Venturini 1996; Dalla
Vecchia 1997b, 1998b, 2001c; Mezga and
Bajraktarevic 1999; Dalla Vecchia et al.
2001)
Sauropod tracks
Theropod tracks
Age: late Cenomanian (Dalla Vecchia
et al. 2001)
29.1.
JUDET
¸
UL BIHOR
,
ROMANIA
1. Unnamed unit (Nopcsa 1901; Csiki and
Grigorescu 1998; =“Gosau coal”)
Tetanurae indet. (=Megalosaurus
hungaricus)
Age: ?early Campanian (Csiki and
Grigorescu 1998)
29.3.
JUDET
¸
UL HUNEDOARA
,
ROMANIA
1. Sânpetru Formation (Nopcsa 1915,
1923a; Huene 1929b, 1932; Harrison and
Walker 1975; Grigorescu 1983a, 1983b,
1984a, 1984b; Grigorescu et al. 1985;
Weishampel et al. 1991, 1993, 2003;
Weishampel and Jianu 1996; Csiki 1997,
1999; Csiki and Grigorescu 1998;
Grigorescu et al. 1999)
Theropoda
Ceratosauria indet.
?Troodontidae indet. (including
Bradycneme dracullae,
Elopteryx nopcsai,
Heptasteornis andrewsi,
cf. Euronychodon sp.)
Dromaeosauridae indet.
Theropoda indet. (=Megalosaurus sp.)
Sauropoda
Lithostrotia
Magyarosaurus dacus (including
M. hungaricus,
M. transsylvanicus)
Lithostrotia indet.
(=Titanosauridae indet.)
Ankylosauria
Nodosauridae
Struthiosaurus transylvanicus
Ornithopoda
Euornithopoda
Zalmoxes robustus
Zalmoxes shqiperorum
Hadrosauridae
Telmatosaurus transsylvanicus
592 DINOSAUR DISTRIBUTION AND BIOLOGY

?Dinosaur coprolites
Age: ?late Maastrichtian (Grigorescu
1983a, 1987; 1992)
2. Densu¸s Ciula Formation (Weishampel
et al. 1991, 2003; Csiki 1997; Csiki and
Grigorescu 1998; Grigorescu et al.
1999)
Theropoda
?Troodontidae indet.
Dromaeosauridae indet.
Unnamed theropod
Theropoda indet.
Sauropoda
Lithostrotia
Magyarosaurus dacus
Ornithopoda
Euornithopoda
Zalmoxes robustus
Zalmoxes shqiperorum
Age: ?late Maastrichtian
(Grigorescu 1983a,
1987, 1992)
29.4.
JUDET
¸
UL ALBA
,
ROMANIA
1. Unnamed unit (=“Boze¸s strata”; Nopcsa
1905c; Grigorescu 1987; Weishampel et
al. 2003)
Theropoda indet.
Ankylosauria
Nodosauridae
Struthiosaurus transylvanicus
Ornithopoda
Euornithopoda
Zalmoxes shqiperorum
Age: ?late Maastrichtian (Antonescu
1973; Ianovici et al. 1976;
Antonescu et al. 1983)
30.1.
KRYMSKAYA OBLAST
,
UKRAINE
1. Unnamed unit (Riabinin 1945)
Ornithopoda
Hadrosauridae indet. (=Orthomerus
weberi)
Age: Maastrichtian (Jeletzky 1962)
37.6.
KURSKAYA OBLAST
,
RUSSIA
1. Unnamed unit (Kiprijanow 1883)
Theropoda indet. (=Poekilopleuron
schmidti)
Age: Cenomanian (Rozhdestvensky
1973b)
DINOSAUR DISTRIBUTION 593
Asia
The Asian record of Late Cretaceous dinosaurs (110 locations, an
increase of more than 80%; figs. 24.6, 24.16, 24.20, 24.21, and
24.22) includes the famous discoveries in the Mongolian
Gobi Desert (Lavas 1993) by the Central Asiatic Expeditions (An-
drews 1932), the Joint Soviet-Mongolian Expeditions (Efremov
1956), and the Polish-Mongolian Expeditions (Kielan-Jaworowska
1969). These explorations continue to the present day through
the important research of the Joint Mongolian Academy of
Science–American Museum of Natural History expeditions (No-
vacek 1996) and the Hayashibara Museum of Natural Sciences–
Geological Institute of the Academy of Science of Mongolia Joint
Paleontological Expeditions (Ishii et al. 1995).
Dinosaur material has also been recovered from early Late
Cretaceous horizons of Kazakhstan, Uzbekistan, Tadzhikistan,
and Kirghizia by S. M. Kurzanov, A. N. Riabinin, A. K. Rozhdest-
vensky, and most recently by J. D. Archibald, A. O. Averianov,
L. A. Nessov, H.-D. Sues, and colleagues. The dinosaurs of central
India are known through the work of C. A. Matley and F. von
Huene and later work by S. L. Jain, D. M. Mohabey, K. N. Prasad,
A. Sahni, and P. Yadagiri. Known from important locations in
the People’s Republic of China, principally Nei Mongol Zizhiqu,
Henan, and Shandong, these dinosaur assemblages have been
studied by C. Wiman and C.-C. Young, and later by Hu C.-C.,
P. J. Currie, D. A. Eberth, P. Godefroit, Zhao X.-J., Pang Q.-P., and
Chen Z.-W. Finally, through recent efforts, the dinosaur faunas
of Japan are beginning to be understood, thanks to the efforts
of Y. Azuma, Y. Tomida, Y. Hasegawa, and colleagues.
31.1.
MECHOOZ YERUSHALAYIM
,
ISRAEL
1. Unnamed unit (Avnimelech 1962a,
1966)
Theropod tracks
Age: early Cenomanian (Avnimelech
1962a, 1966)
32.1.
MUHAFAZAT DIMASHQ
,
SYRIA
1. Unnamed unit (Hooijer 1968)
Theropoda indet.
Age: ?Cenomanian (Hooijer 1968)
33.1.
HASHEMITE KINGDOM
,
JORDAN
1. Phosphorite Unit, Balqa Group (Martill
et al. 1996b)
?Ornithopoda indet.
Age: Maastrichtian (Martill et al. 1996)
34.1.
MUSQAT
,
SULTANATE OF OMAN
1. Al-Khod Conglomerate (Schulp et al.
2000)
Theropoda indet.
Ornithopoda indet.
Age: Maastrichtian (Nolan et al. 1990)
37.7.
YEVREYSKAYA AVTONOMNAYA OBLAST
,
RUSSIA
1. Tsagaan Svita (Riabinin 1930b; Bolotsky
and Kurzanov 1991; Godefroit et al.
2003; Godefroit, pers. comm.; Kurzanov,
pers. comm.)
Theropoda
?Tyrannosauridae indet.
Ornithopoda
Hadrosauridae
Amurosaurus riabinini
Olorotitan arharensis
Hadrosaurinae indet. (=Trachodon
amurensis)
Age: late Maastrichtian (Godefroit,
pers. comm.)
37.8.
CHUKOTKA AUTONOMOUS OKRUG
,
RUSSIA
1. Kakanaut Formation (Nessov and
Golovneva 1990; Rich et al. 1997)
Theropoda
Troodontidae
Troodon cf. formosus
?Avialae indet.
Theropoda indet.
Ornithopoda
Hadrosauridae indet.
Age: middle Maastrichtian (Nessov and
Golovneva 1990)
37.9.
VOLGOGRAD OBLAST
,
RUSSIA
1. Unnamed unit (Kurochkin 2000)
Theropoda
Avialae
?Hesperornithiformes indet.
(=Hesperornis rossicus)
Hesperornis sp.
Age: early Campanian (Kurochkin 2000)
2. Unnamed unit (Kurochkin 2000)
Theropoda
Avialae
Aves
?Pelicanniformes
Volgavis marina
Age: late Maastrichtian (Kurochkin
2000)
3. Unnamed unit (Averianov and Yarkov
2004) (not plotted)
?Tetanurae indet.
?Dromaeosauridae indet.
Age: early Maastrichtian (Averianov and
Yarkov 2004)
37.10.
AMURSKAYA OBLAST
,
RUSSIA
1. Tsagaan Svita (Bolotsky and Moiseenko
1988; Bolotsky et al. 1989)
Theropoda indet.
Ornithopoda
Undescribed hadrosaurid
Age: Campanian-Maastrichtian
(Bolotsky et al. 1989)
37.11.
SAKHALINSKAYA OBLAST
,
RUSSIA
1. Ryugase Group (Nagao 1936; Suzuki
et al., 2004)
Ornithopoda
Hadrosauridae

Nipponosaurus sachalinensis
Age: late Santonian–early Campanian
(Matsumoto and Obata
1979)
39.1.
ONGTUSTIK QAZAQSTAN OBLYSY
,
KAZAKHSTAN
1. Yalovach Svita (Rozhdestvensky and
Khozatsky 1967)
Theropoda indet.
Sauropoda indet.
Ornithopoda
Hadrosauridae
Lambeosaurinae indet.
(?=Jaxartosaurus sp.)
Age: Turonian–early Santonian
(Rozhdestvensky 1977)
2. Dabrazinskaya Svita (Riabinin 1931b,
1937, 1938; Rozhdestvensky 1968a;
Nessov 1995; Kurzanov, pers. comm.)
Sauropoda indet. (=Antarctosaurus
jaxartensis)
Ornithopoda
Hadrosauridae
Jaxartosaurus aralensis (including
Procheneosaurus convincens)
Hadrosauridae indet.
(=Bactrosaurus prynadai,
Cionodon kysylkumensis)
Age: Turonian-Santonian
(Rozhdestvensky 1968a;
Kurzanov, pers. comm.)
3. Unnamed unit (Nurumov 1964;
Kurzanov, pers. comm.)
Dinosaur eggs
Age: ?Late Cretaceous (Rozhdestvensky
1973b)
39.3.
QYZYLORDA OBLYSY
,
KAZAKHSTAN
1. Beleuta Svita (Rozhdestvensky 1968a;
Kurzanov, pers. comm.)
Theropoda
Tyrannosauridae indet.
?Therizinosauridae indet.
Sauropoda indet.
Ankylosauria
Ankylosauridae indet.
Ornithopoda
Hadrosauridae
Aralosaurus tuberiferus
Hadrosauridae indet.
Age: Turonian-Santonian
(Rozhdestvensky 1968a)
2. Bostobe Svita (Shilin and Suslov 1982;
Nessov et al. 1994; Kordikova et al.
1996; Kurzanov, pers. comm.)
Theropoda
Tyrannosauroidea incertae sedis
Alectrosaurus sp.
Tyrannosauridae indet.
Therizinosauroidea indet.
Dromaeosauridae indet.
Theropoda indet.
Ankylosauria
Ankylosauridae indet.
Ornithopoda
Hadrosauridae
Aralosaurus tuberiferus
cf. Lophorhothon sp.
Hadrosauridae indet. (including
Arstanosaurus akkurganensis)
Age: Santonian-early Campanian
(Nessov et al. 1994)
3. Manrakskaya Svita (Bazhanov 1961;
Sochava 1971; Kurzanov, pers. comm.)
Dinosaur eggs
Age: Late Cretaceous (Rozhdestvensky
1973b)
39.4.
ZHEZQAZGHAN OBLYS
,
KAZAKHSTAN
1. Kankazgan Formation (Kordikova et al.
1996)
Theropoda
Tyrannosauridae indet.
Ornithomimidae indet.
Dromaeosauridae indet.
Age: Cenomanian-Turonian (Kordikova
et al. 1996)
39.5.
QOSTANAY OBLYSY
,
KAZAKHSTAN
1. Eginsai Svita (Nessov and Prizemlin
1991; Kurochkin 2000)
Theropoda
Avialae
Hesperornithiformes
Asiahesperornis bazhanovi
Age: late Santonian–early Campanian
(Kurochkin 2000)
40.3.
QORAQALPOGHISTON RESPUBLIKASI
,
UZBEKISTAN
1. Upper Khodzhakul Formation (Nessov
et al. 1994; Nessov 1995)
?Troodontidae indet. (including
Pectinodon asiamericanus,
Troodon asiamericanus)
Ankylosauria indet.
Ornithopoda
Hadrosauridae
?Gilmoreosaurus sp.
Ceratopsia
Neoceratopsia indet. (=Asiaceratops
salsopaludalis, Kulceratops
kulensis)
Age: early Cenomanian (Nessov et al.
1994)
40.4.
BUKHARA WILOYATI
,
UZBEKISTAN
1. Unnamed unit (Nessov and Kaznachkin
1986)
Dinosaur eggs
Age: Late Cretaceous (Nessov and
Kaznachkin 1986)
40.5.
NAWOIY WILOYATI
,
UZBEKISTAN
1. Beleuta Svita (Kurzanov 1976a)
Theropoda
?Tyrannosauroidea
Itemirus medullaris (may be from
Bissekty Formation;
Archibald et al. 1998)
Ornithomimidae indet.
Ornithopoda
Hadrosauridae indet. (=Cionodon
kysylkumensis)
Age: Turonian-Santonian
(Rozhdestvensky 1968a)
40.6.
BUKHORO WILOYATI
,
UZBEKISTAN
1. Bissekty Formation (Nessov 1984, 1986,
1992, 1995; Nessov and Yarkov 1989;
Currie et al. 1993; Nessov and Pantheleev
1993; Nessov et al. 1994; Kurochkin
1996, 1999, 2000; Archibald et al. 1998;
Averianov 2002)
Theropoda
Tyrannosauridae indet.
Therizinosauroidea indet.
Ornithomimosauria indet. (including
Archaeornithomimus
bissektensis)
Oviraptorosauria
Caenagnathasia martinsoni
?Troodontidae indet. (=Euronychodon
asiaticus)
Dromaeosauridae indet.
Avialae
Ornithothoraces
Kuszholia mengi
Enantiornithes
Enantiornis walkeri
Enantiornis martini
Kizylumavis cretacea
Lenesornis maltshevskyi
Sazavis prisca
Zhyraornis kashkarovi
Zhyraornis logunovi
?Avialae indet. (=Ichthyornis
minusculus, Platannavis nana)
Sauropoda indet.
Ankylosauria incertae sedis
Amtosaurus archibaldi
Ankylosauridae indet.
Ornithopoda
Hadrosauridae
Bactrosaurus kysylkumensis
Undescribed hadrosaurid
Hadrosauridae indet.
Ceratopsia
Neoceratopsia indet. (including
Turanoceratops tardabilis)
Ceratopsidae indet.
Age: late Turonian-Coniacian (Nessov
et al. 1994)
42.1.
OSH OBLAST
,
KIRGHIZIA
1. Ialovachsk Formation (Nessov and
Kaznachkin 1986)
Dinosaur eggs
Age: Turonian-Santonian (Nessov and
Kaznachkin 1986)
2. Nichkesaisk Formation (Nessov and
Kaznachkin 1986)
Dinosaur eggs
Age: Santonian-Campanian (Nessov and
Kaznachkin 1986)
42.2.
DZHALAL
-
ABAD OBLAST
,
KIRGHIZIA
1. Nichkesaisk Formation (Nessov and
Kaznachkin 1986)
Dinosaur eggs
Age: Santonian-Campanian (Nessov and
Kaznachkin 1986)
43.1.
VILOYATI LENINOBOD
,
TADZHIKISTAN
1. Unnamed unit (Zakharov 1964;
Rozhdestvensky 1973b)
594 DINOSAUR DISTRIBUTION AND BIOLOGY

Theropod tracks
Age: Cenomanian (Rozhdestvensky
1973b)
2. Yalovach Svita (Rozhdestvensky 1977)
Theropoda
Tyrannosauridae indet.
Ornithomimidae indet.
?Troodontidae indet. (=Troodon
isfarensis)
Ornithopoda
Hadrosauridae indet.
Age: Turonian–early Santonian
(Rozhdestvensky 1977)
45.1.
BALOCHISTAN
,
PAKISTAN
1. Pab Formation (Malkani et al. 2001)
Sauropoda
Undescribed titanosaurians
Age: Maastrichtian (Malkani et al. 2001)
46.1.
MADHYA PRADESH
,
INDIA
1. Lameta Formation (DasGupta 1931;
Huene and Matley 1933; Chatterjee
1978b; Berman and Jain 1982; Jain and
Sahni 1983; Vianey-Liaud et al. 1987;
Sahni 1989; Sahni and Tripathi 1990;
Sahni and Khosla 1994; Joshi 1995;
Khosla and Sahni 1995; Loyal et al.
1996; Mohabey 2000, 2001)
Theropoda
Ceratosauria
Indosaurus matleyi
Indosuchus raptorius
Laevisuchus indicus
Ceratosauria indet. (=Coeluroides
largus, Dryptosauroides
grandis, Jubbulpuria tenuis,
Lametasaurus indicus
partim, Ornithomimoides
mobilis, O. barasimlensis,
Orthogoniosaurus matleyi)
?Carnosauria
Compsosuchus solus
Ornithomimidae indet.
Sauropoda
Lithostrotia
Titanosaurus indicus
Titanosaurus blanfordi
Jainosaurus septentrionalis
(=Antarctosaurus
septentrionalis)
?Titanosaurus sp.
?Antarctosaurus sp.
Titanosauria indet. (=Lametasaurus
indicus partim, Titanosaurus
cf. madagascariensis)
Ankylosauria indet.
(=Brachypodosaurus gravis;
Lametasaurus indicus partim)
Dinosaur eggs
Age: Maastrichtian (Buffetaut 1987;
Sahni and Tripathi 1990;
Dogra et al. 1994; Khosla
and Sahni 1995; Loyal et al.
1996)
46.2.
ANDHRA PRADESH
,
INDIA
1. Intertrappean Beds (Sahni et al. 1984a,
1984b; Vianey-Liaud et al. 1987; Prasad
1989; Prasad and Khajura 1995;
Mohabey 2000, 2001)
Theropoda indet.
Dinosaur eggs
Age: late Maastrichtian (Prasad 1989;
Prasad and Khajura 1995)
46.3.
GUJARAT
,
INDIA
1. Lameta Formation (Infratrappean
Limestone; Dwivedi et al. 1982;
Mohabey 1983, 1984, 1986, 1990, 1998,
2000, 2001; Srivastava 1983; Srivastava
et al. 1986; Mathur and Srivastava 1987;
Bajpai et al. 1993; Sahni and Khosla
1994; Khosla and Sahni 1995; Chatterjee
and Rudra 1996; Loyal et al. 1996; Jain
and Bandyopadhyay 1997; Roy-
Chowdhury et al. 1998; Wilson
et al. 2003)
Theropoda
Ceratosauria
Rajasaurus narmadensis
Ceratosauria indet.
(=Majungasaurus
crenatissimus)
Undescribed abelisaurids
(two species)
Theropoda indet. (=Megalosaurus sp.)
Sauropoda
?Sauropod tracks
Sauropoda indet. (=Titanosaurus
rahioliensis)
Ankylosauria
Undescribed ?ankylosaurid
Dinosauria eggs
Age: Maastrichtian (Buffetaut 1987;
Sahni and Tripathi 1990;
Dogra et al. 1994; Khosla and
Sahni 1995; Loyal et al. 1996)
2. Wagad Formation (Intertrappean Beds;
Ghevariya and Srikarni 1990)
?Sauropod tracks
?Euornithopod tracks
Age: Maastrichtian (Ghevariya and
Srikarni 1990)
3. Anjar Formation (Intertrappean Beds;
Ghevariya and Srikarni 1990; Mohabey
2000, 2001)
Undescribed saurischians
Undescribed ornithischians
Dinosaur eggs
Age: Maastrichtian (Ghevariya and
Srikarni 1990)
4. Katrol Formation (Intertrappean Beds;
Ghevariya and Srikarni 1990)
?Sauropod tracks
?Euornithopod tracks
Age: Maastrichtian (Ghevariya and
Srikarni 1990)
5. Bhuj Formation (Intertrappean Beds;
Ghevariya and Srikarni 1990)
?Sauropod tracks
?Euornithopod tracks
Age: Maastrichtian (Ghevariya and
Srikarni 1990)
46.5.
MAHARASHTRA
,
INDIA
1. Lameta Formation (Lydekker 1879;
Matley 1941; Prasad and Verma 1967;
Berman and Jain 1982; Sahni et al.
1984a, 1984b; Jain and Sahni 1985;
Vianey-Liaud et al. 1987; Mohabey et al.
1993; Mohabey 1996, 2000a, 2000b,
2001a, 2001b; Jain and Bandyopadhyay
1997; Ghosh et al. 2003; Mohabey and
Samant 2003)
Sauropoda
Lithostrotia
Titanosaurus indicus
Titanosaurus blanfordi
“Titanosaurus” colberti
Antarctosaurus sp.
Sauropoda indet. (=Titanosaurus
cf. madagascariensis)
Dinosaur eggs
Dinosaur coprolites
Age: Maastrichtian (Buffetaut 1987;
Sahni et al. 1984b; Dogra
et al. 1994)
2. Takli Formation (Lydekker 1890b;
Huene and Matley 1933; Sahni et al.
1984a, 1984b, 1984c); Mohabey 2000,
2001)
Theropoda indet. (=Massospondylus
rawesi)
Undescribed sauropod
Dinosaur eggs
Age: late Maastrichtian (Sahni et al.
1984b)
46.6.
TAMIL NADU
,
INDIA
1. Trichinopoly Group (Anonymous 1978;
Yadagiri and Ayyasami 1979, 1989)
Theropoda indet.
Sauropoda indet. (=Bruhathkayosaurus
matelyi)
Stegosauria
Stegosauridae
Dravidosaurus blanfordi
Age: Coniacian (Sastry et al. 1969)
2. Kallankurichi Formation (Huene and
Matley 1933; Rao 1956; Prasad 1968;
Mohabey 2000, 2001)
Theropoda indet.
Sauropoda
Lithostrotia
Titanosaurus indicus
Stegosauria
Stegosauridae indet.
Dinosaur eggs
Age: Maastrichtian (Sastry et al. 1969)
46.7.
MEGHALAYA
,
INDIA
1. Mahadek Formation (Mishra and Sen
2001)
Dinosauria indet.
Age: Maastrichtian (Mishra and Sen
2001)
47.3.
BAYANKHONGOR
,
MONGOLIA
1. Baruungoyot Formation (Elz˙anowski
1974, 1977; Nessov and Kaznachkin
1986; Kurochkin 2000)
Theropoda
Avialae
Enantiornithes
Gobipteryx minuta
Dinosaur eggs
DINOSAUR DISTRIBUTION 595

Age: ?middle Campanian (Gradzí´nski
et al. 1977); Maastrichtian
(Fox 1978; Lillegraven and
McKenna 1986)
2. Nemegt Svita (Barsbold 1983b;
Kurochkin 2000; =Beds of Bugeen Tsav,
Gradzí´nski et al. 1977; Kurochkin 2000)
Theropoda
Tyrannosauroidea
Tarbosaurus bataar
Ornithomimosauria
Anserimimus planinychus
Oviraptorosauria
Ingenia yanshini
Troodontidae
Saurornithoides junior
Dromaeosauridae
Adasaurus mongoliensis
Avialae
Hesperornithiformes
Judiornis nogontsavensis
Ornithopoda
Hadrosauridae
Saurolophus angustirostris
Age: early Maastrichtian (Fox 1978;
Lillegraven and McKenna
1986)
3. Beds of Nogoon Tsav (Kurzanov 1976b)
Theropoda
Tyrannosauroidea incertae sedis
Alioramus remotus
Age: ?Maastrichtian (Kurzanov 1976b)
47.4. Ö
MNÖGOV
’,
MONGOLIA
1. Bayanshiree Svita (Perle 1977, 1979;
Kurzanov 1976b; Kurzanov and
Tumanova 1978; Barsbold and Perle
1980, 1983; Nessov and Kaznachkin
1986; Tumanova 1987; Watabe et al.
1997)
Theropoda
Tyrannosauroidea incertae sedis
Alectrosaurus olseni
Therizinosauroidea
Segnosaurus galbinensis
Erlikosaurus andrewsi
Ornithomimosauria
Garudimimus brevipes
Ankylosauria
Ankylosauridae
Talarurus plicatospineus
Ankylosauria incertae sedis
Amtosaurus magnus
Ornithopoda
Hadrosauridae
Bactrosaurus mongoliensis (including
Arstanosaurus sp.)
Hadrosauridae indet.
Dinosaur eggs
Age: Cenomanian-Santonian (Hicks et
al. 1999)
2. Djadokta Formation (Gilmore 1933b;
Marya´nska and Osmólska 1975;
Gradzí´nski et al. 1977; Barsbold and
Perle 1983; Elz˙ anowski and Wellnhofer
1992, 1993; Kurzanov 1992; Mikhailov
et al. 1994, 1996; Norell et al. 1994, 1995,
2000, 2001a; Dashzeveg et al. 1995;
Norell and Makovicky 1997; Makovicky
and Norell 1998a, 1998b; Ishigaki 1999;
Suzuki et al. 1999; Clark et al. 2001;
Norell and Clarke 2001; Suzuki et al.
2001; Tereschenkov and Alifanov 2003;
Watabe, pers. comm.)
Theropoda
Coelurosauria incertae sedis
Archaeornithoides deinosauriscus
Tyrannosauroidea
?Tarbosaurus sp.
Ornithomimosauria
Undescribed ornithomimid
Oviraptorosauria
Oviraptor philoceratops
Citipati osmolskae
Unnamed oviraptorine
Khaan mckennai
Oviraptor sp.
cf. Ingenia sp.
Troodontidae
Saurornithoides mongoliensis
Byronosaurus jaffei
Dromaeosauridae
Velociraptor mongoliensis
cf. Velociraptor sp.
Undescribed dromaeosaurid
Dromaeosauridae indet.
Avialae
Alvarezsauridae
Mononykus olecranus
Shuvuuia deserti
Unnamed enantiornithine
Ornithurae
Apsaravis ukhaana
Theropoda indet.
Sauropoda indet.
Ankylosauria
Ankylosauridae
Pinacosaurus grangeri
Ankylosauridae indet.
Ankylosaur tracks
Ornithopoda
Hadrosauridae indet.
Ceratopsia
Neoceratopsia
Protoceratops andrewsi
Bagaceratops sp.
Udanoceratops tschizhovi
Bainoceratops efremovi
Dinosaur eggs
Dinosaur tracks
Age: ?middle Campanian (Jerzykiewicz
and Russell 1991)
3. Beds of Alag Teeg (Gradzí´nski et al. 1977)
Theropoda indet.
Sauropoda indet.
Ankylosauria
Ankylosauridae
Pinacosaurus grangeri
Ceratopsia
Neoceratopsia
?Protoceratops andrewsi
Age: ?late Santonian or early Campanian
(Gradzí´nski et al. 1977)
4. Djadokta Svita (Kurzanov 1987;
Fastovsky et al. 1998)
Theropoda
Oviraptorosauria
Avimimus portentosus
Ornithomimosauria
Gallimimus sp.
Theropod tracks
Sauropoda
Diplodocoidea
Quaesitosaurus orientalis
Sauropod tracks
Age: ?middle Campanian (Jerzykiewicz
and Russell 1991)
5. Baruungoyot Formation (Marya´nska
and Osmólska 1975; Gradzí´nski et al.
1977; Osmólska 1982; Kurochkin
1988, 2000; Kurzanov 1990a, 1990b;
Mikhailov et al. 1994; Karhu and
Rautian 1996; Alifanov 2003)
Theropoda
Dromaeosauridae
Hulsanpes perlei
Velociraptor sp.
Avialae
Alvarezsauridae
Parvicursor remotus
Enantiornithes
Gobipteryx minuta
Hesperornithiformes indet.
Theropoda indet.
Sauropoda indet.
Ankylosauria
Ankylosauridae
Saichania chulsanensis
Tarchia gigantea (=T. kielanae)
Pachycephalosauria
Pachycephalosauridae
Tylocephale gilmorei
Ceratopsia
Neoceratopsia
Bagaceratops rozhdestvenskyi
(=Breviceratops kozlowskii)
Lamaceratops tereschenkoi
Platyceratops tatarinovi
Dinosaur eggs
Age: ?middle Campanian (Gradzí´nski
et al. 1977); Maastrichtian
(Fox 1978; Lillegraven and
McKenna 1986)
6. Red Beds of Hermiin Tsav (Gradzí´nski
et al. 1977; Kurochkin 1996; Mikhailov
1996)
Theropoda
Oviraptorosauria
Ingenia yanshini
Conchoraptor gracilis
Dromaeosauridae
Velociraptor sp.
Avialae
Enantiornithes
Gobipteryx minuta (=Nanantius
valifanovi)
Theropoda indet.
Sauropoda indet.
Ankylosauria
Ankylosauridae
Saichania chulsanensis
Ceratopsia
Neoceratopsia
Bagaceratops rozhdestvenskyi
596 DINOSAUR DISTRIBUTION AND BIOLOGY

Dinosaur eggs
Age: ?middle Campanian (Gradzí´nski
et al. 1977)
7. White Beds of Hermiin Tsav (Gradzí´nski
et al. 1977)
Theropoda
Tyrannosauroidea
Tarbosaurus bataar
Therizinosauroidea
Therizinosaurus cheloniformis
Theropoda indet.
Ankylosauria
Ankylosauridae
Tarchia gigantea
Ornithopoda
Hadrosauridae
Saurolophus angustirostris
Age: early Maastrichtian (Fox 1978;
Lillegraven and McKenna
1986)
8. Nemegt Formation (Gradzí´nski et al.
1977; Kurzanov and Osmólska 1991;
Perle et al. 1994; Osmólska 1996;
Barsbold et al. 2000a, 2000b; Ishii et al.
2000; Kurochkin 2000; Kurochkin et al.
2002; Lü et al. 2002; Currie et al. 2003)
Theropoda
Tyrannosauroidea
Bagaraatan ostromi
Tarbosaurus bataar (including
Maleevosaurus novojilovi)
Tyrannosauridae indet.
Therizinosauroidea
Therizinosaurus cheloniformis
Ornithomimosauria
Gallimimus bullatus
Deinocheirus mirificus
Oviraptorosauria
Oviraptoridae
Rinchenia mongoliensis
Nomingia gobiensis
?Elmisaurus rarus
Ingenia sp.
Oviraptoridae indet.
Troodontidae
Saurornithoides junior
Borogovia gracilicrus
Tochisaur us nemegtensis
Avialae
Alvarezsauridae
Mononykus olecranus
Enantiornithes
Gurilynia nessovi
Hesperornithiformes
cf. Parahesperornis sp.
Hesperornithiformes indet.
Presbyornithidae
Teviornis gobiensis
Theropoda indet.
Theropod tracks
Sauropoda
Diplodocoidea
Nemegtosaurus mongoliensis
Saltasauridae
Opisthocoelicaudia skarzynskii
Sauropod tracks
Ankylosauria
Ankylosauridae
Tarchia gigantea
Ornithopoda
Hadrosauridae
Saurolophus angustirostris
Barsboldia sicinskii
Hadrosauridae indet.
Hadrosaurid tracks
Pachycephalosauria
Homalocephaloidea
Homalocephale calathocercos
Pachycephalosauridae
Prenocephale prenes
Age: early Maastrichtian (Fox 1978;
Lillegraven and McKenna
1986)
9. Shireegiin Gashuun Formation
(=Bayanshiree Svita; Marya´nska and
Osmólska 1975; Mikhailov et al. 1994;
Sereno 2000)
Theropoda
Ornithomimidae indet.
Theropoda indet.
Ankylosauria indet. (=Maleevus
disparoserratus)
Ornithopoda
Hadrosauridae indet.
Ceratopsia
Neoceratopsia
Graciliceratops mongoliensis
(=Microceratops gobiensis)
Dinosaur eggs
Age: Cenomanian-Santonian (Hicks
et al. 1999)
47.5. Ö
VÖRKHANGAI
,
MONGOLIA
1. Unnamed unit (Perle et al. 1982)
Pachycephalosauria
Goyocephala
Goyocephale lattimorei
Age: ?late Santonian or early Campanian
(Osmólska, pers. comm.)
2. “Djadokta” Svita (Kurzanov 1987)
Theropoda
Oviraptorosauria
Avimimus portentosus
Age: ?middle Campanian (Jerzykiewicz
and Russell 1991)
3. Djadokhta Formation (Mikhailov et al.
1994)
Dinosaur eggs
Age: ?middle Campanian (Jerzykiewicz
and Russell 1991)
47.6.
DUNDGOV
’,
MONGOLIA
1. Dohoin Usu Formation (Gilmore 1933b)
Theropoda indet.
Ornithopoda
Hadrosauridae indet.
Age: ?Campanian (equivalent to Iren
Dabasu Formation; Morris
1936)
47.7.
DORNOGOV
’,
MONGOLIA
1. Bayanshiree Svita (Maleev 1952a, 1952b;
Barsbold 1972; Nessov and Kaznachkin
1986; Tumanova 1993; Mikhailov et al.
1994; Perle et al. 1999)
Theropoda
Therizinosauroidea
Enigmosaurus mongoliensis
Segnosaurus galbinensis
Dromaeosauridae
Achillobator giganticus
Theropoda indet.
Ankylosauria
Ankylosauridae
Talarurus plicatospineus
Tsagantegia longicranialis
Ornithopoda
Hadrosauridae indet.
Dinosaur eggs
Age: Cenomanian-Santonian (Hicks
et al. 1999)
2. Djadokhta Formation (Mikhailov et al.
1994)
Dinosaur eggs
Age: ?middle Campanian (Jerzykiewicz
and Russell 1991)
3. Baruungoyot Formation (Mikhailov
et al. 1994)
Dinosaur eggs
Age: ?middle Campanian (Gradzí´nski et
al. 1977; Maastrichtian, Fox
1978; Lillegraven and
McKenna 1986)
48.1.
XIZANG ZIZHIQU
,
PEOPLE
’
S REPUBLIC
OF CHINA
1. Zonggo Formation (Zhao unpubl. data)
Theropoda
Ornithomimosauria
Ornithomimus sp.
Sauropoda
?Lithostrotia
Megacervixosaurus tibetensis
Ornithopoda
Hadrosauridae indet.
Age: Late Cretaceous (Zhao, unpubl.
data)
2. Margyang Formation (Zhao, unpubl.
data)
Pachycephalosauria
cf. Homalocephale calathocercos
Age: Late Cretaceous (Zhao, unpubl.
data)
3. Quwu Formation (Zhen et al. 1989)
Dinosauria tracks
Age: Late Cretaceous (Mateer, pers.
comm.)
48.5.
XINJIANG UYGUR ZIZHIQU
,
PEOPLE
’
S REPUBLIC OF CHINA
1. Subashi Formation (Dong 1977; Chen
et al. 1982)
Theropoda
Tyrannosauroidea
?Tarbosaurus bataar
(?=Shanshanosaurus
huoyanshanensis)
Tyrannosauroidea indet.
(=Tyrannosaurus turpanensis)
Sauropoda
Lithostrotia indet. (=Titanosauridae
indet.)
Sauropoda indet. (=Nemegtosaurus
pachi)
DINOSAUR DISTRIBUTION 597

Ankylosauria indet.
Dinosaur eggs
Age: Maastrichtian (Shen and Mateer
1992)
2. Wulungo Formation (Wu 1973)
Ornithopoda
Hadrosauridae indet. (=Jaxartosaurus
fuyunensis)
Age: Late Cretaceous (Wu 1973)
48.6.
SHAANXI
,
PEOPLE
’
S REPUBLIC
OF CHINA
1. Shanyang Formation (Xue et al. 1996)
Theropoda
Tyrannosauridae indet.
Avetheropoda incertae sedis
Shanyangosaurus niupanggouensis
Sauropoda indet. (=Qinlingosaurus
luonanensis)
Ornithopoda
Hadrosauridae
Shantungosaurus cf. giganteus
Ornithopoda indet.
Dinosaur eggs
Age: Late Cretaceous (Xue et al. 1996)
48.7.
JILIN
,
PEOPLE
’
S REPUBLIC OF CHINA
1. Quan-Tou Formation (Anonymous
1989) (not plotted)
Ornithopoda
Hadrosauridae indet.
Late Cretaceous (Anonymous 1989)
48.8.
GANSU
,
PEOPLE
’
S REPUBLIC
OF CHINA
1. Minhe Formation (Bohlin 1953;
Marya´nska 1977; Mateer, pers. comm.)
Theropoda indet.
?Ankylosauria indet. (=Peishansaurus
philemys)
Ceratopsia
Neoceratopsia
Protoceratops andrewsi
Neoceratopsia indet.
(=Microceratops gobiensis,
M. sulcidens)
?Pachycephalosauria indet.
(=Heishansaurus
pachycephalus)
Age: Campanian-Maastrichtian (Mateer,
pers. comm.)
2. Unnamed unit (Xie 1980)
Theropoda indet.
Sauropoda
Lithostrotia indet. (=Titanosauridae
indet.)
Ornithopoda
Hadrosauridae indet.
Age: early Maastrichtian (i.e.,
contemporary with the
Subashi Formation; Dong
1977; Chen et al. 1982)
48.9.
SHANDONG
,
PEOPLE
’
S REPUBLIC
OF CHINA
1. Wangshi Group (Wiman 1929; Chow
1951; Young 1958a; Dong 1978; Buffetaut
1995a; Buffetaut and Tong 1995)
Theropoda
Tyrannosauroidea
?Tarbosaurus bataar
(?=Chingkankousaurus fragilis)
Theropoda indet.
(=cf. Szechuanosaurus campi)
Ankylosauria
Ankylosauridae
Pinacosaurus cf. grangeri
Ornithopoda
Hadrosauridae
Tanius sinensis (including
T. chingkankouensis)
Shantungosaurus giganteus
Tsintaosaurus spinorhinus (including
Tanius laiyangensis)
Hadrosauridae indet.
Lambeosaurinae indet.
Pachycephalosauria incertae sedis
Micropachycephalosaurus
hongtuyanensis
Dinosaur eggs
Age: Campanian (Buffetaut and Tong
1995)
48.10.
HEBEI
,
PEOPLE
’
S REPUBLIC
OF CHINA
1. Huiquanpu Formation (Pang and Cheng
2001)
Theropoda indet. (=cf. Szechuanosaurus
campi)
Sauropoda
Titanosauria
Huabeisaurus allocotus
Ankylosauria
Ankylosauridae
Tianzhenosaurus youngi
Ornithopoda
Hadrosauridae
cf. Shantungosaurus sp.
Age: Late Cretaceous (Pang and Cheng
2001)
48.11.
NEI MONGOL ZIZHIQU
,
PEOPLE
’
S REPUBLIC OF CHINA
1. Iren Dabasu Formation (=Iren Nor
Formation, Reline Dabasu Formation;
Van Straelen 1925, 1928; Gilmore 1933a;
Rozhdestvensky 1966; Weishampel and
Horner 1986; Dong et al. 1989; Mader
and Bradley 1989; Currie and Eberth
1993; Godefroit et al. 1998b; Currie and
Dong 2001a; Ryan et al. 2001a; X. H.
Zhang et al. 2001; Xu et al. 2002d)
Theropoda
Tyrannosauroidea incertae sedis
Alectrosaurus olseni
Therizinosauroidea
Neimongosaurus yangi
Erliansaurus bellamanus
Therizinosauroidea indet.
Ornithomimosauria
Archaeornithomimus asiaticus
?Garudimimus sp.
Oviraptorosauria
Avimimus portentosus
Troodontidae
Saurornithoides sp.
Troodontidae indet.
Dromaeosauridae
Velociraptor sp.
Dromaeosauridae indet.
Avialae
Alvarezsauridae
Mononykus olecranus
Sauropoda indet. (=?Camarasauridae
indet.)
Ankylosauria
Undescribed ankylosaurid
Ornithopoda
Euornithopoda
Hadrosauridae
Bactrosaurus johnsoni
Gilmoreosaurus mongoliensis
?Euornithopoda indet.
Dinosaur eggs
Age: ?Campanian (Currie and Eberth
1993)
2. Djadokhta Formation (Dong et al. 1989;
Currie and Peng 1993; Jerzykiewicz
et al. 1993; Zhao and Li 1993; Dong
and Currie 1996; Godefroit et al. 1999;
Currie and Dong 2001a; Lambert et al.
2001; You and Dong 2003)
Theropoda
Tyrannosauroidea
Tarbosaurus sp.
Oviraptorosauria
Oviraptor philoceratops
Troodontidae
Saurornithoides mongoliensis
Dromaeosauridae
Velociraptor mongoliensis
Sauropoda indet.
Ankylosauria
Ankylosauridae
Pinacosaurus grangeri
Pinacosaurus mephistocephalus
Ornithopoda
Hadrosauridae indet.
Ceratopsia
Neoceratopsia
Protoceratops andrewsi
Protoceratops hellenikoorhinus
Magnirostris dodsoni
cf. Bagaceratops sp.
Udanoceratops sp.
Dinosaur eggs
Age: ?middle Campanian (Jerzykiewicz
and Russell 1991)
3. Minhe Formation (Bohlin 1953;
Marya´nska 1977; Mateer, pers. comm.)
Theropoda
Dromaeosauridae
cf. Velociraptor mongoliensis
Theropoda indet.
Sauropoda indet.
Ankylosauria
Nodosauridae indet.
Ceratopsia
Neoceratopsia
Protoceratops andrewsi
Neoceratopsia indet.
(=Microceratops gobiensis)
Pachycephalosauria
Pachycephalosauridae
598 DINOSAUR DISTRIBUTION AND BIOLOGY

“Troodon” bexelli
Age: Campanian-Maastrichtian (Mateer,
pers. comm.)
48.12.
HENAN
,
PEOPLE
’
S REPUBLIC
OF CHINA
1. Quiba Formation (Dong 1979)
Theropoda
Tyrannosauroidea
?Tarbosaurus bataar
(?=Tyrannosaurus
luanchuanensis)
Age: Late Cretaceous (Dong 1979)
2. Nanchao Formation (Zhao 1979; Tong
and Wang 1980; Cohen et al. 1995;
Manning et al. 1997)
Theropoda
Therizinosauroidea indet.
?Euornithopod tracks
Dinosaur eggs
Age: Late Cretaceous (Zhen et al. 1985)
3. Zoumagang Formation (Li et al. 1995;
Fang et al. 1998)
Sauropoda indet.
Dinosaur eggs
Age: Late Cretaceous (Fang et al. 1998)
4. Sangping Formation (Fang et al. 1998;
Xu unpubl. data)
Theropoda indet.
Sauropoda indet.
?Ankylosauria indet.
Ornithopoda
Iguanodontia indet.
Dinosaur eggs
Age: Late Cretaceous (Fang et al. 1998)
5. Zhaoyin Formation (Fang et al. 1998)
Dinosaur eggs
Age: Late Cretaceous (Fang et al. 1998)
6. Unnamed unit (Fang et al. 1998)
Dinosaur eggs
Age: Late Cretaceous (Fang et al. 1998)
7. Gaogou Formation (Whyte et al. 2000)
Dinosaur eggs
Age: Late Cretaceous (Whyte et al. 2000)
8. Mayiacun Formation (Whyte et al. 2000;
Zhang et al. 2000)
Sauropoda indet.
Ornithopoda
Hadrosauridae indet.
Dinosaur eggs
Age: Late Cretaceous (Whyte et al. 2000)
9. Sigou Formation (Whyte et al. 2000)
Dinosaur eggs
Age: Late Cretaceous (Whyte et al.
2000)
48.13.
ZHEJIANG
,
PEOPLE
’
S REPUBLIC
OF CHINA
1. Tangshang Formation (Dong 1979;
Mateer 1989)
Theropoda
Therizinosauroidea indet.
(=“Chilantaisaurus”
zheziangensis)
Dinosaur eggs
Age: Cenomanian-Turonian (Chen and
Chang 1994); Santonian-
Campanian (Dong 1979)
2. Quxian Formation (Mateer 1989)
Dinosaur eggs
Age: Campanian (Chen and Chang
1994)
3. Wangi Formation (Zhen et al. 1989)
Dinosauria tracks
Age: Late Cretaceous (Mateer, pers.
comm.)
48.14.
NINGXIA HUIZU ZIZHIQU
,
PEOPLE
’
S REPUBLIC OF CHINA
1. Unnamed unit (Young 1935b;
Marya´nska 1977)
Ankylosauria
Ankylosauridae
Pinacosaurus grangeri (=Pinacosaurus
ninghsiensis)
Age: ?middle Campanian (i.e.,
?contemporary with
Djadokhta Formation;
Marya´nska 1977)
2. Unnamed unit (Zhao and Ding 1976)
Dinosauria eggs
Age: ?early Late Cretaceous (Zhao and
Ding 1976)
48.15.
SHANXI
,
PEOPLE
’
S REPUBLIC
OF CHINA
1. Zhumabao Formation (Mateer,
pers. comm.)
Ceratopsia
Neoceratopsia
Microceratops sp.
Age: early Late Cretaceous (Mateer,
pers. comm.)
2. Huiquanpu Formation (Cheng and
Pang 1996; Pang et al. 1996; Barrett
et al. 1998; Pang and Cheng 1998,
2000, 2001)
Theropoda indet. (including
cf. Szechuanosaurus campi)
Sauropoda
Titanosauria
Huabeisaurus allocotus
Lithostrotia indet.
(=Titanosauridae indet.)
Ankylosauria
Ankylosauridae
Tianzhenosaurus youngi
Ankylosauridae indet.
Ankylosauria indet. (=Shanxia
tianzhenensis)
Ornithopoda
Hadrosauridae
cf. Shantungosaurus sp.
Hadrosauridae indet.
Age: Late Cretaceous (Cheng and Pang
1996)
48.18.
HEILONGJIANG
,
PEOPLE
’
S REPUBLIC
OF CHINA
1. Tsagaan Svita (Riabinin 1930a)
Ornithopoda
Hadrosauridae
Tanius sinensis
Hadrosaurinae indet. (=Trachodon
amurensis)
Hadrosauridae indet.
Age: ?Coniacian-?Maastrichtian
(=?Senonian;
Rozhdestvensky 1973b)
2. Yongantun Formation (Yang 1987)
Dinosauria indet.
Age: Cenomanian (Yang 1987)
3. Yuliangze Formation (Yang et al. 1986;
Yang 1987; Godefroit et al. 2000, 2001)
Theropoda
Tyrannosauridae indet.
(=Tyrannosaurus sp.)
Ornithomimosauria indet.
(=Ornithomimus sp.)
Ornithopoda
Hadrosauridae
Charonosaurus jiayinensis
Hadrosauridae indet.
(=Mandschurosaurus
amurensis)
Age: Turonian (Yang 1987)
4. Unnamed unit (Riabinin 1930a)
Theropoda
Tyrannosauroidea
?Tarbosaurus bataar (=Albertosaurus
periculosus)
Ornithopoda
Hadrosauridae indet. (=Saurolophus
kryschtofovici)
Age: Late Cretaceous (Riabinin 1930a)
48.19.
ANHUI
,
PEOPLE
’
S REPUBLIC
OF CHINA
1. Xiaoyan Formation (Hou 1977)
Pachycephalosauria
Wannanosaur us yansiensis
Age: Campanian (Chen and Chang
1994)
48.20.
HUBEI
,
PEOPLE
’
S REPUBLIC
OF CHINA
1. Unnamed unit (Zhao unpubl. data)
Ornithopoda indet.
Dinosaur eggs
Age: ?Late Cretaceous (Zhao unpubl.
data)
48.21.
HUNAN
,
PEOPLE
’
S REPUBLIC
OF CHINA
1. Red Beds (Zeng and Zhang 1979)
Dinosaur eggs
Age: Late Cretaceous (Zeng and Zhang
1979)
2. Jinjiang Formation (Zheng 1982)
Dinosaur tracks
Age: Late Cretaceous (Zheng 1982)
48.22.
GUANGDONG
,
PEOPLE
’
S REPUBLIC
OF CHINA
1. Yuanpu Formation (=Nanxiong
Formation; Zhao 1975; Dong 1979;
Zhen et al. 1989)
Theropoda
Tyrannosauroidea
Tarbosaurus bataar
Tyrannosauridae indet.
Therizinosauroidea
Nanshiungosaurus brevispinus
Theropoda indet.
DINOSAUR DISTRIBUTION 599

Ornithopoda
Hadrosauridae
Hadrosaurinae indet.
(=Microhadrosaurus
nanshiungensis)
Dinosaur eggs
Undescribed dinosaur tracks
Age: Campanian (Chen and Chang
1994)
2. Unnamed unit (Qiu and Huang 2001)
Theropoda
Unnamed oviraptorosaurian
Dinosaur eggs
Age: Late Cretaceous (Qiu and Huang
2001)
3. Dalangshan Formation (Lü 2002)
(not plotted)
Theropoda
Oviraptorosauria
Heyuannia huangi
Age: Maastrichtian (Lü 2002)
48.23.
JIANGSU
,
PEOPLE
’
S REPUBLIC
OF CHINA
1. Wangshi Formation (Zhen et al. 1989)
Undescribed dinosaur tracks
Age: Campanian (Buffetaut and Tong
1995)
50.3.
CH
ˇ
OLLA NAMDO
,
SOUTH KOREA
1. Uhangri Formation (Yang et al. 1995;
Lee et al. 2001; Lee and Huh 2002)
Theropod tracks (including avialan
tracks)
Euornithopod tracks
Sauropod tracks
Age: Campanian (Lee and Huh 2002)
2. Seonso Conglomerate (Huh and
Zelenitsky 2002)
Dinosaur eggs
Age: ?Late Cretaceous (Huh and
Zelenisky 2002)
53.3.
FUKUSHIMA PREFECTURE
,
JAPAN
1. Futaba Group (Hasegawa et al. 1987;
Azuma and Tomida 1997)
Theropoda indet.
Ornithischia indet.
Ornithopoda
Hadrosauridae indet.
Age: Coniacian-Santonian (Azuma and
Tomida 1997)
2. Ashizawa Formation (Hasegawa et al.
1987; Tanimoto and Suzuki 1998)
Theropoda indet.
Ornithopoda
Hadrosauridae indet.
Age: Coniacian (Tanimoto and Suzuki
1998)
3. Tamayama Formation (Tanimoto and
Suzuki 1997, 1998)
Sauropoda indet. (=cf. Nemegtosaurus sp.)
Age: Santonian (Tanimoto and Suzuki
1997, 1998)
53.4.
KUMAMOTO PREFECTURE
,
JAPAN
1. Mifune Group (Tamura et al. 1991;
Hasegawa et al. 1992; Ikegami and
Tamura 1997; Chure et al. 1999)
Theropoda
?Carcharodontosauridae indet.
Theropoda indet.
Ornithopoda
Iguanodontia indet.
?Ceratopsia indet.
Age: Cenomanian (Matsukawa and
Obata 1994; Azuma and
Tomida 1997; Chure et al.
1998)
53.6.
TOKUSHIMA PREFECTURE
,
JAPAN
1. Akazaki Formation (Hasegawa et al.
1987 )
Theropoda indet.
Ornithopoda
Hadrosauridae indet.
Age: Coniacian (Matsukawa and Obata
1994)
2. Tamayama Formation (Hasegawa 1988)
Sauropoda indet. (=Diplodocidae
indet.)
Age: late Santonian (Matsukawa and
Obata 1994)
53.14.
HOKKAIDO PREFECTURE
,
JAPAN
1. Yezo Group (Azuma and Tomida 1997;
Hayakawa, pers. comm; ?=Ezo Group)
Ankylosauria
Undescribed nodosaurid
Ornithopoda
Hadrosauridae indet.
Age: Coniacian (Azuma and Tomida
1997)
53.15.
KYUSHU PREFECTURE
,
JAPAN
1. Mitsuse Formation (Matsuo 1967; Obata
et al. 1972)
Ornithopoda
Hadrosauridae indet.
Age: early Santonian or late Coniacian
(Matsuo 1967)
600 DINOSAUR DISTRIBUTION AND BIOLOGY
South America
The South American record of Late Cretaceous dinosaurs (61 lo-
cations, an increase of nearly 50%; figs. 24.8 and 24.9) consists
of a number of early discoveries by F. Ameghino, R. Lydekker,
and F. Huene in Argentina. Recent work throughout the conti-
nent, however, has eclipsed these efforts. In Brazil, R. J. Bertini,
D. A. Campos, A. W. A. Kellner, and L. I. Price have unearthed
a wealth of dinosaur material, much of which is still to be pub-
lished. Bolivian, Peruvian, Uruguayan, and Chilean dinosaurs
are now known through the work of F. de Broin, G. Faccio, J. Iri-
arte, C. A. Meyer, B. Sigé, and others. Finally, the pace of discov-
eries of Late Cretaceous dinosaurs from Argentina has dramati-
cally increased, thanks to the work of a host of investigators,
including J. F. Bonaparte, J. Powell, G. del Corro, R. A. Coria, J. O.
Calvo, L. M. Chiappe, F. E. Novas, R. D. Martinez, and L. Salgado,
R. M. Casamiquela, and O. Alcober.
54.3.
DEPARTAMENTO DE TOLIMA
,
COLOMBIA
1. “Ortega” formation (Langston 1965)
Theropoda indet.
Age: Maastrichtian (Langston 1965)
56.1.
DEPARTAMENTO DE PUNO
,
PERU
1. Upper Vilquechico Formation (Sigé
1968; Kérourio and Sigé 1984; Jaillard
et al. 1993)
Theropod tracks
Euornithopod tracks
Dinosaur eggs
Age: late Campanian–late
Maastrichtian (Sempere
et al. 1997)
56.2.
DEPARTAMENTO DE AMAZONAS
,
PERU
1. Bagua Formation (Mourier et al. 1986,
1988)
Theropoda indet.
Sauropoda
Lithostrotia indet. (=Titanosauridae
indet.)
Dinosaur eggs
Age: late Campanian–Maastrichtian
(Mourier et al. 1988)
56.3.
DEPARTAMENTO DE CUZCO
,
PERU
1. Soncco Formation (Noblet et al. 1995)
Theropod tracks
Euornithopod tracks
Age: Maastrichtian (Mourier et al. 1994)
57.2.
ESTADO DO SÃO PAULO
,
BRAZIL
1. Adamantina Formation (Arid and
Vizotto 1971; Powell 1986; Bertini et al.
1993; Kellner 2000; Santucci and Bertini
2001)
Dinosauria indet.
Theropoda
Abelisauridae indet.
Theropoda indet. (including
?Megalosaurus sp.)
Sauropoda
Lithostrotia indet. (including
Antarctosaurus brasiliensis,
Titanosaurus cf. australis,
Titanosauridae indet.)
Undescribed titanosaurian

Age: Cenomanian-Campanian (Soares
et al. 1980)
2. Marilia Formation (Bertini et al. 1993)
Theropoda indet. (=?Ceratosaurus sp.)
Sauropoda
Lithostrotia indet. (=Titanosauridae
indet.)
Age: Campanian-Maastrichtian (Soares
et al. 1980)
3. Baurú Group (Chiappe 1991b; Kellner
and Azevedo 1999)
Theropoda
Avialae indet.
Sauropoda
Lithostrotia
Antarctosaurus brasiliensis
Gondwanatitan faustoi
Unnamed lithostrotian
Lithostrotia indet.
(=Titanosauridae indet.)
Age: Late Cretaceous (Kellner and
Azevedo 1999)
57.3.
ESTADO DO PARANÁ
,
BRAZIL
1. Baurú Group (Bonaparte 1978a)
Theropoda indet.
Sauropoda
Lithostrotia indet.
(=cf. Titanosaurus sp.)
Age: Campanian-Maastrichtian (Petri
and Campanha 1981)
57.4.
ESTADO DO MINAS GERAIS
,
BRAZIL
1. Marília Formation (Price 1951;
Bonaparte 1978a; Campos and Bertini
1985; Powell 1986; Bertini et al. 1993;
Kellner and Campos 1998; Campos
and Kellner 1999; Kellner et al. 1999;
=Uberaba Formation, Kellner 2000)
Theropoda
Abelisauridae indet.
Theropoda indet.
Sauropoda
Lithostrotia
Antarctosaurus brasiliensis
Undescribed titanosaurian
Dinosaur eggs
Age: Campanian-Maastrichtian (Soares
et al. 1980)
2. Uberaba Formation (Santucci and
Bertini 2001)
Sauropoda
Undescribed titanosaurian
Titanosauridae indet.
Age: Campanian-Maastrichtian
(Santucci and Bertini
2001)
57.5.
ESTADO DO GOIÁS
,
BRAZIL
1. Baurú Group (Bonaparte 1978a)
Theropoda indet.
Sauropoda
Lithostrotia
Antarctosaurus brasiliensis
Lithostrotia indet.
(=cf. Titanosaurus sp.)
Age: Campanian-Maastrichtian
(Petri and Campanha
1981 )
57.9.
ESTADO DO MATTO GROSSO
,
BRAZIL
1. Baurú Group (Price 1960; Kellner et al.
1995)
Theropoda indet.
Sauropoda
Lithostrotia indet. (=Titanosauridae
indet.)
Age: Campanian-Maastrichtian (Petri
and Campanha 1981)
57.10.
ESTADO DO MARANHÃO
,
BRAZIL
1. Itapecuru Formation (Price 1947;
Carvalho and Pedrão 1998; Carvalho
et al. 2003)
Theropoda indet.
Theropod tracks
Sauropoda
Diplodocoidea
Amazonsaurus maranhensis
Sauropoda indet.
Age: Cenomanian-Santonian (Petri and
Mendes 1981); Aptian-Albian
(Rossetti et al. 2001)
2. Unnamed unit (Kellner 2000)
Dinosauria indet.
Theropoda
Spinosauridae indet.
Theropoda indet.
Sauropoda indet.
Age: Late Cretaceous (Kellner 2000)
57.11.
ESTADO DO AMAZONAS
,
BRAZIL
1. Sucunduri Formation (Price 1960)
Theropoda indet.
Age: Late Cretaceous (Price 1960)
58.1.
DEPARTEMENT DE SORIANO
,
URUGUAY
1. Mercedes Formation (Faccio 1994)
Dinosaur eggs
Age: Coniacian-Maastrichtian
(=Senonian; Bossi and
Navarro 1990)
58.2.
DEPARTAMENTO PAYSANDÚ
,
URUGUAY
1. Guichon Formation (Huene 1934b,
Bonaparte 1978a)
Theropoda
?Ornithomimidae indet.
?Ornithopoda
?Iguanodontia indet.
Age: Coniacian-Maastrichtian
(=Senonian; Bonaparte
1978a)
2. Mercedes Formation (Faccio 1994)
Dinosaur eggs
Age: Coniacian-Maastrichtian
(=Senonian; Bossi and
Navarro 1990)
58.3.
DEPARTAMENTO PALMITAS
,
URUGUAY
1. Asencio Formation (Huene 1929a; Bossi
1966)
Sauropoda
Titanosauria
Lithostrotia
Antarctosaurus wichmannianus
Laplatasaurus araukanicus
Saltasauridae
Neuquenosaurus australis
Titanosauria indet.
(=Argyrosaurus superbus)
Age: Late Cretaceous (Bossi 1966)
58.4.
DEPARTAMENTO COLONIA
,
URUGUAY
1. Asencio Formation (Mones 1980)
Dinosaur eggs
Age: Late Cretaceous (Mones 1980)
58.5.
DEPARTAMENTO TACUAREMBÓ
,
URUGUAY
1. Asencio Formation (Mones 1980)
Dinosaur eggs
Age: Late Cretaceous (Mones 1980)
59.1.
DEPARTAMENTO DE POTOSI
,
BOLIVIA
1. El Molino Formation (=Toro Toro
Formation; Branisa 1968; Muizon et al.
1983; Marshall et al. 1985; Leonardi
1989, 1994; Gayet et al. 2001; McCrea
et al. 2001a)
Theropoda
Coelurosauria indet.
Theropod tracks
Sauropoda indet.
Sauropod tracks
?Ankylosaur tracks
?Euornithopod tracks
Age: middle Maastrichtian (Muizon
et al. 1983; Leonardi 1989;
Gayet et al. 1993, 2001)
59.2.
DEPARTAMIENTO DE COCHABAMBA
,
BOLIVIA
1. Santa Lucia Formation (Leonardi 1989)
Theropod tracks
Euornithopod tracks
Age: Maastrichtian (Leonardi 1989)
59.3.
DEPARTAMIENTO DE CHUQUISACA
,
BOLIVIA
1. Chaunaca Formation (Mazzetta and
Markley 2000)
Theropod tracks
Sauropod tracks
Age: Campanian (Mazzetta and Markley
2000)
2. El Molino Formation (Meyer et al. 2001)
Theropod tracks
Sauropod tracks
Ankylosaur tracks
Euornithopod tracks
Age: Maastrichtian (Muizon et al. 1983;
Leonardi 1989; Gayet et al.
1993)
62.4.
PROVINCIA DE CONCEPCIÓN
,
CHILE
1. Unnamed unit (Lambrecht 1929;
S. L. Olson 1992)
Theropoda
Avialae
Neornithes
Gaviiformes
Neogaeornis wetzeli
Age: Maastrichtian (Chiappe 1996a)
DINOSAUR DISTRIBUTION 601

62.5.
PROVINCIA DE COQUIMBO
,
CHILE
1. Viñita Formation (Casamiquela et al.
1969; Vargas et al. 2000)
Sauropoda
Lithostrotia
Antarctosaurus cf. wichmannianus
Lithostrotia indet.
(=Titanosauridae indet.)
Age: Campanian-Maastrichtian
(Gasparini 1979)
62.6.
PROVINCIA DE BIOBÍO
,
CHILE
1. Quiriquina Formation (Lambrecht 1929;
Chiappe 1991b)
Theropoda
Avialae
Neornithes
Gaviiformes
Neogaeornis wetzeli
Age: Campanian-Maastrichtian
(Riccardi 1988)
62.7.
PROVINCIA DE COPAIPO
,
CHILE
1. Los Hornitos Formation (Iriarte,
pers. comm.)
Sauropoda
Titanosauria indet.
Age: Late Cretaceous (Iriarte, pers.
comm.)
62.8.
PROVINCIA DE LIMARI
,
CHILE
1. Unnamed unit (Iriarte, pers. comm.)
Sauropoda
Lithostrotia
Antarctosaurus wichmanianus
Antarctosaurus sp.
Lithostrotia indet.
(=Titanosauridae indet.)
Age: Late Cretaceous (Iriarte,
pers. comm.)
63.1.
PROVINCIA DE LA RIOJA
,
ARGENTINA
1. Unnamed unit (Arcucci, pers. comm.)
Sauropoda
Lithostrotia indet. (=Titanosauridae
indet.)
Age: Maastrichtian (Sempere et al. 1997)
63.3.
PROVINCIA DE SANTA CRUZ
,
ARGENTINA
1. Chorrillo Formation (Powell 1986)
Theropoda indet.
Sauropoda
Lithostrotia indet. (=Titanosauridae
indet.)
Age: Maastrichtian (Powell 1986)
2. Cardiel Formation (=“Guaranitic”
formation; Ameghino 1898, 1899a;
Huene 1929a; Molnar 1980b; Powell
1986; Coria and Salgado 1996a)
Sauropoda
Lithostrotia indet. (=Titanosauridae
indet.)
Sauropoda indet. (=Clasmodosaurus
spatula)
Ornithopoda indet. (=Loncosaurus
argentinus)
Age: Late Cretaceous (Huene 1929a)
3. Unnamed unit (Huene 1929a)
Sauropoda
Titanosauria indet. (=Argyrosaurus
superbus)
Age: Late Cretaceous (Huene 1929a)
4. Unnamed unit (Dibenedetto, pers.
comm., in Bonaparte and Gasparini
1979)
Sauropoda
Titanosauria
Argyrosaurus superbus
Age: Late Cretaceous (Bonaparte
and Gasparini 1979)
63.4.
PROVINCIA DE CHUBUT
,
ARGENTINA
1. Laguna Palacios Formation (Bonaparte
1979a)
Theropoda indet.
Sauropoda
Titanosauria indet. (=Argyrosaurus
superbus)
Age: Coniacian-Maastrichtian
(=Senonian; Bonaparte
1979a)
2. Bajo Barreal Formation (Bonaparte and
Gasparini 1979; Martínez et al. 1986,
1993, 1999; Powell 1986; Powell et al.
1989; Martínez and Novas 1997;
Martínez 1998, 1999; Lamanna, pers.
comm.; =“San Jorge” formation, Brett-
Surman 1979)
Theropoda
Ceratosauria
Xenotarsosaurus bonapartei
Abelisauridae indet.
Undescribed tetanuran
?Coelurosauria
Aniksosaurus darwini
Theropoda indet.
Sauropoda
Undescribed diplodocoid
Titanosauria
Epachthosaurus sciuttoi
Undescribed titanosaurian
Lithostrotia indet.
(=Titanosauridae indet.)
Sauropoda indet. (including
Campylodon ameghinoi)
?Ornithischian tracks
Ornithopoda
Euornithopoda
Notohypsilophodon comodorensis
Hadrosauridae
Secernosaurus koerneri
Age: Cenomanian-Coniacian (Bridges
et al. 2000)
3. Castillo, Bajo Barreal, and Laguna
Palacios formations (undifferentiated;
Huene 1929a; Powell 1986)
Theropoda
?Abelisauridae indet.
Sauropoda
Titanosauria
Argyrosaurus superbus
Argyrosaurus sp.
Lithostrotia
Antarctosaurus cf. wichmannianus
Laplatasaurus araukanicus
cf. Laplatasaurus sp.
Sauropoda indet. (=Campylodoniscus
ameghinoi)
Age: Cenomanian-Coniacian (Bridges
et al. 2000); Campanian-
Maastrichtian (Powell
1986)
4. La Colonia Formation (Bonaparte 1985;
Ardolino and Delpino 1987)
Theropoda
Ceratosauria
Carnotaurus sastrei
Age: Campanian-Maastrichtian (Pascual
et al. 2000)
5. Unnamed unit (Casamiquela vide
Leonardi 1994)
Euornithopod tracks
Age: late Maastrichtian (Casamiquela
vide Leonardi 1994)
6. Cerro Castillo Formation (Corro 1966;
Bonaparte, pers. comm.)
Theropoda
Tetanurae indet. (=Megalosaurus
chubutensis, M. inexpectatus)
Age: Late Cretaceous (Bonaparte 1978a)
7. Unnamed unit (Woodward 1901;
Bonaparte 1978a)
Theropoda
Ceratosauria indet. (=Genyodectes
serus)
Age: Late Cretaceous (Bonaparte 1978a)
63.6.
PROVINCIA DE NEUQUÉN
,
ARGENTINA
1. Río Limay Formation (Leonardi 1989;
Calvo 1991, 1999; Calvo and Bonaparte
1991; Bonaparte and Coria 1993; Coria
and Salgado 1993b, 1998a, 1998b,
1999b, 2001; Leonardi 1994; Calvo and
Salgado 1995; Coria and Salgado 1995;
Bonaparte 1996b; Coria and Currie
1997; Calvo and Coria 1998; Mazzetta
and Blanco 2001; Leanza and Hugo
2001; Coria and Calvo 2002; includes
Candeleros, Huincul, and Cerro
Lisandro formations)
Theropoda
Ceratosauria
Ilokelesia aguadagrandensis
Unnamed abelisaur
Carnosauria
Giganotosaurus carolinii
Unnamed carcharodontosaurid
Theropod tracks
Sauropoda
Diplodocoidea
Rayososaurus tessonei (=
Rebbachisaurus tessonei)
Titanosauria
Argentinosaurus huinculensis
Andesaurus delgadoi
Titanosauridae indet.
Sauropod tracks
Ornithopoda
Euornithopoda
Anabisetia saldiviai
Euornithopod tracks
602 DINOSAUR DISTRIBUTION AND BIOLOGY

Age: Cenomanian–early Turonian
(Leanza and Hugo 2001);
Albian-Cenomanian (Calvo
and Bonaparte 1991)
2. Río Neuquén Formation (Huene 1929a;
Bonaparte and Gasparini 1979; Novas
et al. 1996; Novas and Puerta 1997;
Novas 1998; Calvo et al. 2001; Coria
2001; Calvo and González Riga 2003;
=Portezuelo Formation, Leanza and
Hugo 2001)
Theropoda
Tetanurae indet.
?Dromaeosauridae
Troodontidae indet.
Dromaeosauridae
Unenlagia comahuensis
Megaraptor namunhuaiquii
Avialae
Alvarezsauridae
Patagonykus puertai
Theropoda indet.
Sauropoda
Lithostrotia
Antarctosaurus wichmannianus
Rinconsaurus cadamirus
Titanosaurus sp.
Undescribed lithostrotian
Sauropoda indet. (=Antarctosaurus
giganteus)
Ornithopoda indet.
Age: Turonian-Coniacian (Cruz et al.
1989; Leanza and Hugo
2001)
3. Río Colorado Formation (Bonaparte
and Gasparini 1979; Powell 1986;
Chiappe and Calvo 1989; Chiappe 1990,
1992; Bonaparte 1991a; Alvarenga and
Bonaparte 1992; Salgado and Calvo
1993; Calvo et al. 1997b; Chiappe et al.
1998b, 2001a, 2001b; González Riga and
Calvo 2001; Coria et al. 2000, 2002a)
Theropoda
Ceratosauria
Velocisaurus unicus
Aucasaurus garridoi
Carnosauria
Unnamed carcharodontosaurid
Avialae
Alvarezsauridae
Alvarezsaurus calvoi
Ornithothoraces
Patagopteryx deferrariisi
Enantiornithes
Neuquenornis volans
Enantiornithes indet.
Theropoda indet.
Sauropoda
Titanosauria
Lithostrotia
Saltasauridae
Neuquenosaurus australis
(including ?Loricosaurus
scutatus, Microcoelus
patagonicus, Titanosaurus
nanus)
Titanosaurus sp.
Undescribed lithostrotian
Lithostrotia indet.
(=Titanosauridae indet.)
Titanosauria indet.
(=cf. Argyrosaurus sp.)
Titanosaurid eggs
Age: Santonian (Leanza and Hugo
2001); Santonian–early
Campanian (Heredia and
Salgado 1999)
4. Unnamed unit (Leonardi 1989)
Theropod tracks
Age: ?Campanian-Maastrichtian
(=Senonian; Leonardi 1989)
5. ?Laguna Palacios Formation (Tapia 1918;
Huene 1929a)
?Ceratopsia indet. (=Notoceratops
bonarellii)
Age: Late Cretaceous (Bonaparte 1996b)
63.7.
PROVINCIA DE SALTA
,
ARGENTINA
1. Los Blanquitos Formation (Powell 1978,
1979, 1986; Bonaparte 1996b)
Theropoda
?Dromaeosauridae
Unquillosaurus ceibalii
Theropoda indet.
Sauropoda
Sauropoda indet.
Titanosauria
Lithostrotia
Laplatasaurus cf. araukanicus
?Titanosaurus sp.
Lithostrotia indet.
(=Titanosauridae indet.)
Titanosauria incertae sedis
Antarctosaurus sp.
Dinosaur eggs
Age: ?Campanian (Bonaparte et al. 1977)
2. Yacoraite Formation (Bonaparte and
Bossi 1967; Alonso and Marquillas 1976,
1986; Alonso 1980, 1989; Leonardi
1989)
Theropoda indet.
Theropod tracks (including avialan trac ks)
Ornithopoda
Euornithopod tracks
Hadrosauridae indet.
Age: Maastrichtian (Sempere et al. 1997)
3. Lecho Formation (Bonaparte and Powell
1980; Walker 1981; Brett-Surman and
Paul 1985; Powell 1986; Chiappe 1991a,
1993; Frankfurt and Chiappe 1999)
Theropoda
Ceratosauria
Noasaurus leali
?Abelisauridae indet.
?Oviraptorosauria indet.
Avialae
Enantiornithes
Avisaurus archibaldi
Enantiornis leali
Soroavisaurus australis
Lectavis bretincola
Yungavolucris brevipedalis
Enantiornithes indet.
Sauropoda
Saltasauridae
Saltasaurus loricatus
Age: ?late Campanian-Maastrichtian
(Bonaparte et al. 1977;
Powell 1986)
63.8.
PROVINCIA DE ENTRE RÍOS
,
ARGENTINA
1. Puerto Yerua Formation (=Tacuarembo
Formation; Huene 1929a; Powell 1986)
Sauropoda
Lithostrotia indet. (=Argyrosaurus
superbus)
Age: Late Cretaceous (Powell 1986)
63.9.
PROVINCIA DE MENDOZA
,
ARGENTINA
1. Río Colorado Formation (Alcober et al.
1995, 1998a; Wilson et al. 1999a; Leanza
and Hugo 2001; includes Bajo de la
Carpa and Anacleto formations)
Theropoda
Undescribed carcharodontosaurid
Sauropoda
Lithostrotia indet. (=Titanosauridae
indet.)
Age: Santonian (Leanza and Hugo 2001);
Santonian–early Campanian
(Heredia and Salgado 1999)
2. Loncoche Formation (González Riga
1999b)
Theropoda indet.
Sauropoda indet.
Age: late Campanian–early
Maastrichtian (González
Riga 1999b)
63.10.
PROVINCIA DE RÍO NEGRO
,
ARGENTINA
1. Angostura Colorada Formation (J. E.
Powell 1986, 1987c; Bonaparte 1996b)
Sauropoda
Lithostrotia
Aeolosaurus rionegrinus
Lithostrotia indet.
(=Titanosauridae indet.)
Dinosaur eggs
Age: Campanian (Powell 1986;
Bonaparte 1996b)
2. Los Alamitos Formation (Bonaparte
et al. 1984; J. E. Powell 1986, 1987c;
Chiappe 1996a; Salgado et al. 1997b)
Theropoda
Avialae indet.
Theropoda indet.
Sauropoda
Lithostrotia
Aeolosaurus rionegrinus
Ornithopoda
Hadrosauridae
Unnamed hadrosaurine
(=“Kritosaurus” australis)
Dinosaur eggs
Age: late Campanian–early Maastrichtian
(Bonaparte et al. 1984)
3. Coli Toro Formation (Casamiquela
1964a; Powell 1986)
Sauropoda
Lithostrotia indet. (=Titanosauridae
indet.)
DINOSAUR DISTRIBUTION 603

Ornithopoda
Hadrosauridae indet.
Age: Campanian-Maastrichtian (Powell
1986)
4. Río Colorado Formation (Lydekker
1893a; Huene 1929a; Bonaparte and
Gasparini 1978; Bonaparte and Novas
1985; Coria and Salgado 1996b; Leanza
and Hugo 2001; Calvo, pers. comm.;
includes Bajo de la Carpa and Anacleto
formations)
Theropoda
Ceratosauria
Abelisaurus comahuensis
?Abelisauridae indet.
Theropoda indet.
Sauropoda
Lithostrotia
Antarctosaurus wichmannianus
Lithostrotia indet.
(=cf. Macrurosaurus sp.)
Saltasauridae
Neuquenosaurus australis
(including ?Loricosaurus
scutatus, Titanosaurus nanus
Saltasaurus robustus
?Titanosaurus sp.
Ornithopoda
Euornithopoda
Gasparinisaura cincosaltensis
Age: Santonian (Leanza and Hugo 2001);
Santonian–early Campanian
(Heredia and Salgado 1999)
5. Allen Formation (Huene 1929a; Powell
1986, 1996; Salgado and Coria 1993a,
1993b, 1996; Bonaparte 1996; Salgado
1996; Salgado and Azpilicueta 2000;
Clarke and Chiappe 2001; Coria 2001;
Priéto-Marquez 2002)
Theropoda
Avetheropoda incertae sedis
Quilmesaurus curriei
Avialae
Ornithurae
Limenavis patagonica
Sauropoda
Titanosauria
Undescribed titanosaurian
Lithostrotia
Aeolosaurus sp.
Laplatasaurus araukanicus
Pellegrinisaurus powelli
Rocasaurus muniozi
Saltasauridae
Saltasaurus robustus
Ankylosauria
?Nodosauridae indet.
Ornithopoda
Hadrosauridae
Undescribed hadrosaurid
Lambeosaurinae indet.
Dinosaur eggs
Age: middle Campanian (Heredia and
Salgado 1999)
6. Unnamed unit (Casamiquela 1987;
Chiappe 1991b)
Dinosaur tracks
Avialan tracks
Age: late Maastrichtian (Casamiquela
1987 )
7. Unnamed unit (Chiappe 1992)
Theropoda
Avialae
Undescribed ornithurines
Age: Maastrichtian (Chiappe 1996a)
63.11.
PROVINCIA DE LA PAMPA
,
ARGENTINA
1. Allen Formation (González Riga and
Casadio 2000)
Ornithopoda
Hadrosauridae
Hadrosaurinae indet.
Age: middle Campanian (Heredia and
Salgado 1999)
2. Colorado Formation (Casadío et al.
2002) (not plotted)
Dinosaur eggs
Age: Campanian-Maastrichtian
(Casadío et al. 2002)
604 DINOSAUR DISTRIBUTION AND BIOLOGY
Africa
African Late Cretaceous dinosaur locations now number 20, a
110% increase since 1992. Among the best come from the Western
Desert of Egypt; these were collected and studied by E. Stromer.
Although this material was destroyed at the close of World
War II, new collecting efforts have been made there by O. W. M.
Rauhut, and most recently by J. B. Smith and co-workers. Ad-
ditional African sites of significance include Morocco (worked
by A. F. de Lapparent and co-workers, and P. C. Sereno and co-
workers), Niger (worked by P. Taquet), and Madagascar (worked
by M. C. Deperét, A. Thevenin, R. Lavocat, I. Obata and co-
workers, and P. Taquet and co-workers, and by D. W. Krause,
S. D. Sampson, C. A. Forster, and co-workers. Late Cretaceous
dinosaur sites from Africa are shown in figures 24.10, 24.11,
and 24.23.
64.7.
PROVINCE D
’
AGADIR
,
MOROCCO
1. Unnamed unit (Ambroggi and
Lapparent 1954)
Theropod tracks
Age: Maastrichtian (Ambroggi and
Lapparent 1954)
64.6.
PROVINCE DE KSAR
-
ES
-
SOUK
,
MOROCCO
1. Upper Kem-Kem Beds (Lavocat 1954;
Jacobs et al. 1996; Sereno et al. 1996;
Sadleir 1998)
Theropoda
Carnosauria
Carcharodontosaurus saharicus
Coelurosauria incertae sedis
Deltadromeus agilis
Dromaeosauridae indet.
Theropod tracks
Sauropoda
Undescribed sauropods
(two species?)
Sauropoda indet.
(=Rebbachisaurus sp.)
Sauropod tracks
Ornithopoda
Undescribed iguanodontian
Euornithopod tracks
Age: Cenomanian (Sereno et al. 1996)
2. Continental Red Beds (=Grès rouge
infracénomaniens; Monbaron 1978;
Buffetaut 1989a, 1989b; Russell 1996;
Sereno et al. 1996, 1998)
Theropoda
Ceratosauria
Abelisauridae indet. (including
cf. Majungasaurus sp.)
Tetanurae incertae sedis
Sigilmassasaurus brevicollis
Sigilmassasaurus sp.
Spinosauroidea
Spinosaurus maroccanus
Spinosaurus cf. aegyptiacus
Spinosaurus sp.
Carnosauria
Carcharodontosaurus saharicus
Carcharodontosaurus sp.
?Carcharodontosaurus sp.
Theropoda indet. (including
cf. Elaphrosaurus sp.)
Sauropoda
Lithostrotia indet. (=Titanosauridae
indet.)
Sauropoda indet. (including
Rebbachisaurus sp.)
Age: Cenomanian (Buffetaut 1989a,
1989b; Russell 1996)
65.6.
WILAYA LAGHOUAT
,
ALGERIA
1. Unnamed unit (Bellair and Lapparent
1949)
Theropod tracks
Age: Cenomanian (Bellair and
Lapparent 1949)
67.1.
REGION DE GAO
,
MALI
1. Unnamed unit (Mateer et al. 1992)
Theropoda indet. (=Megalosaurus sp.)
Sauropoda
Lithostrotia indet. (=Titanosauridae
indet.)
Ornithopoda
Iguanodontia indet. (=Dryosaurus sp.)
Age: Late Cretaceous (Mateer et al.
1992)

68.2.
DÉPARTEMENT DE TAHOUA
,
NIGER
1. In Beceten Formation (Greigert et al.
1954; Broin et al. 1974)
Sauropoda
Lithostrotia indet. (=Titanosauridae
indet.)
Age: Coniacian-Santonian (Taquet 1976;
Moody and Sutcliffe 1991)
2. Unnamed unit (Greigert et al. 1954;
Broin et al. 1974; Moody and Sutcliffe
1991; Jacobs et al. 1996)
Theropoda
Avetheropoda incertae sedis
Bahariasaurus ingens
Sauropoda
Lithostrotia indet. (=Titanosauridae
indet.)
Age: Coniacian-Santonian (Moody and
Sutcliffe 1991)
3. Dukamaje Formation (Moody and
Suttcliffe 1991)
Theropoda
Avetheropoda incertae sedis
?Bahariasaurus ingens
Age: Maastrichtian (Kogbe 1981)
69.1.
AL
-
BAHR AL
-
AHMAR
,
EGYPT
1. Abu Agag Formation (Demathieu and
Wycisk 1990)
?Ornithischian tracks
Age: Turonian (Hendriks et al. 1987)
69.2.
MARSA MATR
¯
UH
,
EGYPT
1. Baharija Formation (Stromer 1934a;
El-Khashab 1977; Rauhut 1995; Jacobs
et al. 1996; Smith et al. 2001b)
Theropoda
Spinosauridae
Spinosaurus aegyptiacus
Spinosaurus sp.
Avetheropoda incertae sedis
Bahariasaurus ingens
Allosauroidea
Carcharodontosaurus saharicus
Undescribed theropod
Theropoda indet. (=Elaphrosaurus sp.,
cf. Erectopus sauvagei)
Sauropoda
Dicraeosauridae
cf. Dicraeosaurus sp.
Titanosauria incertae sedis
Aegyptosaurus baharijensis
Lithostrotia
Paralititan stromeri
Sauropoda indet.
Age: ?early Cenomanian (Said 1962)
2. Variegated Shale of the Nubian
Formation (Awad and Ghobrial 1966;
Hotton, pers. comm., in El-Khashab
1977 )
Ornithischia indet.
Age: pre-Maastrichtian Late Cretaceous
(El-Khashab 1977)
3. Nubian Sandstone (Stromer and Weiler
1930)
Theropoda indet.
Age: ?Campanian (Said 1962)
4. Phosphate Beds (Stromer and Weiler
1930; Molnar 1980b; Jacobs et al. 1996)
Theropoda
?Abelisauridae indet. (=Majungasaurus
crenatissimus)
Age: Campanian-Maastrichtian (Said
1962)
5. Dahkla Formation (Rauhut and Werner
1997)
Sauropoda
Titanosauriformes indet.
(=?Brachiosauridae indet.)
Age: Maastrichtian (Barthel and
Herrmann-Degen 1981)
71.1.
NORTHERN PROVINCE
,
SUDAN
1. Wadi Milk Formation (Broin et al. 1974;
Buffetaut et al. 1990; Werner 1993;
Rauhut and Werner 1995; Jacobs et al.
1996; Rauhut 1999)
Theropoda
Avetheropoda incertae sedis
Bahariasaurus sp.
Allosauroidea
Carcharodontosaurus sp.
Carcharodontosauridae indet.
Dromaeosauridae indet.
Theropoda indet.
Sauropoda
Dicraeosauridae indet.
Lithostrotia indet. (=Titanosauridae
indet.)
Ornithopoda
Euornithopoda indet.
Iguanodontia
cf. Ouranosaurus sp.
Iguanodontia indet.
Ornithopoda indet.
Age: ?Albian-Cenomanian (Schrank
1989)
2. Shendi Formation (Rauhut 1999)
(not plotted)
Theropoda indet.
Age: ?Albian-Cenomanian (Schrank 1989)
74.1.
RIFT VALLEY PROVINCE
,
KENYA
1. Turkana Grits (Arambourg and Wolff
1969; Jacobs et al. 1996; Harris and
Russell 1986)
Theropoda
Spinosauridae
cf. Spinosaurus sp.
Undescribed theropod
Theropoda indet.
Sauropoda
Lithostrotia indet.
(=cf. Titanosaurus sp.)
Sauropoda indet.
?Ornithopoda indet.
Age: Turonian-Santonian (Westcott
et al. 1993)
81.4.
NATAL
,
SOUTH AFRICA
1. Unnamed unit (Kennedy et al. 1987)
Sauropoda indet. (=?Titanosauridae
indet.)
Age: early Santonian (Kennedy et al.
1987 )
83.2.
FARITANY MAJUNGA
,
MADAGASCAR
1. Ankarafantsika Formation (Mateer et al.
1992)
Sauropoda indet.
Age: Cenomanian (Mateer et al. 1992)
2. Ankazomihaboka Formation (Curry
1997; Rogers and Hartman 1998)
Theropoda indet. (including
Majungasaurus crenatissimus)
Sauropoda
Lithostrotia
“Titanosaurus” madagascariensis
Age: Coniacian (Curry 1997; Rogers and
Hartman 1998)
3. Maevarano Formation (=Grès de
Maevarano, Berivotro Formation;
Depéret 1896; Thévenin 1907; Lavocat
1955a; Russell et al. 1976; Sues and
Taquet 1979; Sues 1980b; Forster 1996c;
Forster et al. 1996, 1998; Jacobs et al.
1996; Sampson et al. 1996, 1997, 1998,
2001; Dodson et al. 1998; Curr y Rogers
and Forster 2001; Carrano et al. 2002)
Theropoda
Ceratosauria
Majungatholus atopus
Masiakasaurus knopfleri
Majungasaurus crenatissimus
?Spinosauridae indet.
Avialae
Alvarezsauridae
Rahonavis ostromi
Ornithothoraces
Vorona berivotrensis
Enantiornithes indet.
Sauropoda
Lithostrotia
Rapetosaurus krausei
“Titanosaurus” madagascariensis
Undescribed lithostrotian
Lithostrotia indet.
(=Titanosauridae indet.)
?Ornithischia indet. (=Stegosaurus
madagascariensis)
Age: Campanian (Obata and Kanie
1977; Rogers and Hartman
1998)
DINOSAUR DISTRIBUTION 605
Australasia
Over the past decade, the number of Australasian dinosaur faunas
has doubled to four (figs. 24.1 and 24.12). A single Late Cretaceous
location is known from Queensland, Australia (in large part
an important ichnofauna being worked by R. A. Thulborn and

M. Wade. Two newly discovered faunas, so far based on intrigu-
ing but as yet poorly known theropod skeletal material, have
been reported from Western Australia by J. A. Long. The last site
comes from North Island, New Zealand (described by R. E. Mol-
nar, R. J. Scarlett, and J. Wiffen).
606 DINOSAUR DISTRIBUTION AND BIOLOGY
84.1.
QUEENSLAND
,
AUSTRALIA
1. Winton Formation (Coombs and
Molnar 1981; Thulborn and Wade 1984;
Rich and Vickers Rich 1997; Salisbury,
pers. comm.)
Saurischia indet.
Theropod tracks
Sauropoda
Titanosauria
Austrosaurus sp.
Undescribed sauropod
Sauropod tracks
Ornithischia indet.
Euornithopod tracks
Age: Cenomanian (Thulborn and Wade
198 4)
84.2.
WESTERN AUSTRALIA
,
AUSTRALIA
1. Molecap Greensand (Long 1995)
Theropoda indet.
Age: Cenomanian-?Santonian
(McWhae et al. 1958)
2. Miria Formation (Long 1992)
?Theropoda indet.
Age: late Maastrichtian (Long 1992)
85.1.
NORTH ISLAND
,
NEW ZEALAND
1. Tahora Formation (Molnar
1980c; Scarlett and Molnar 1984;
Wiffen and Molnar 1989;
Molnar and Wiffen 1994;
Wiffen 1996)
Theropoda indet.
Sauropoda indet.
Ankylosauria
Nodosauridae indet.
Ornithopoda
Iguanodontia indet.
Age: Campanian (Moore 1987)
Antarctica
Despite the remoteness and arduous collecting conditions of the
continent, three important discoveries have been made from
the Upper Cretaceous of Antarctica over the past fifteen years
(fig. 24.1). Described by J. A. Case, Z. de Gasparini, J. J. Hooker,
A. C. Milner, R. E. Molnar, and colleagues, these occurrences
come from James Ross and Vega islands.
86.0.
ANTARCTICA
1. Hidden Lake Formation (Molnar et al.
1996)
Theropoda indet.
Age: Coniacian-Santonian (Molnar et al.
1996)
2. Santa Marta Formation (Gasparini et al.
1987, 1996)
Ankylosauria
Nodosauridae indet.
Age: late Campanian (Olivero et al.
1991 )
3. López de Bertodano Formation
(Chatterjee 1989, 2002; Hooker et al.
1991; Milner et al. 1992; Case and
Tambussi 1999; Rich et al. 1999a; Case et
al. 2000; Milner, pers. comm.)
Theropoda
Avialae
Aves
Gaviidae
Polarornis gregorii
Undescribed avialans
Ornithopoda
Euornithopoda
Undescribed euornithopodan
Hadrosauridae indet.
Age: late Campanian–early
Maastrichtian (Pirrie et al.
1991; Milner et al.1992)