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Paleobiogeography and biodiversity of Late Maastrichtian dinosaurs: How

many dinosaur species went extinct at the Cretaceous-Tertiary boundary?

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Bull. Soc. géol. France, 2012, t. 183, no 6, pp. 547-559

Paleobiogeography and biodiversity of Late Maastrichtian dinosaurs: how

many dinosaur species went extinct at the Cretaceous-Tertiary boundary?
JEAN LE LOEUFF1

Key-words. – Dinosauria, Maastrichtian, Species-area relationship, Palaeobiogeography, Biodiversity, Extinction

Abstract. – The global Late Maastrichtian non-avian dinosaur apparent biodiversity is extensively surveyed for the first
time. It amounts to 104 species (including unnamed forms) in 2010. The real biodiversity being obscured by
taphonomical biases and the scarcity of the continental fossil record, a species-area relationship is used to estimate it.
The results show that several hundreds (between 628 and 1078) non-avian dinosaur species were alive in the Late
Maastrichtian, which is almost an order of magnitude above previous estimates. Because of the complex Late Creta-
ceous palaeobiogeography, discussions about dinosaur extinction should be based on this estimated real global
biodiversity, not on the apparent biodiversity of a single area. Given the mean duration of dinosaur genera (7.7 Ma), the
presence of so many dinosaur species in the Latest Cretaceous is consistent with the termination of most lineages at the
Cretaceous-Tertiary boundary (the Late Maastrichtian sub-stage is 2.8 m.y. long). The Late Maastrichtian dinosaurian
biodiversity is therefore consistent with the sudden extinction of the group following the Chicxulub impact.

Paléobiogéographie et diversité des dinosaures du Maastrichtien supérieur : combien de

genres de dinosaures ont-ils disparu à la limite Crétacé-Tertiaire ?

Mots-clés. – Dinosauria, Maastrichtien, Courbe aire-espèces, Paléobiogéographie, Biodiversité, Extinction

Résumé. – La biodiversité apparente des dinosaures au Maastrichtien supérieur est analysée en détail : 104 espèces
(comprenant des formes non-nommées) ont été découvertes en 2010. La biodiversité réelle est altérée par de nombreux
biais taphonomiques et la rareté de l’enregistrement fossile continental. Pour l’estimer une relation aire-espèces est uti-
lisée, suggérant qu’un minimum de plusieurs centaines d’espèces (entre 628 et 1078) de dinosaures non-aviens vivaient
durant le Maastrichtien supérieur. Ce résultat est un ordre de grandeur au-dessus des estimations précédentes, qui ont
négligé la complexité du contexte paléobiogéographique. Etant donnée la longévité moyenne d’un genre de dinosaure
(7.7 Ma) la plupart de ces formes ont dû s’éteindre à la limite Crétacé-Tertiaire. La biodiversité dinosaurienne à l’ex-
trême fin du Crétacé était donc remarquablement élevée et la disparition du groupe a été soudaine, ce qui est
parfaitement compatible avec les suites de l’impact de Chicxulub.

INTRODUCTION focused on the presence or absence of a decline of this di-

Although the Cretaceous-Tertiary boundary is widely
known as the dinosaur extinction, a review of the literature
suggests that only a handful of dinosaur species went ex-
tinct at the Cretaceous Tertiary Boundary, or even slightly
before it (12 to 14 species according to Sullivan [1987];
Taquet [1993]; 20 genera according to Archibald [1996,
1997]). As these authors state, these data are not consistent
with a catastrophic mass extinction and rather suggest that
dinosaurs went extinct gradually on a time span of several
million years. However, these data are only based on the ap-
parent dinosaur biodiversity in Montana, where outcrops of
the Late Maastrichtian Hell Creek Formation have been ex-
tensively searched for fossil vertebrates [see Archibald,
1997] although the biodiversity of a small part of western
North America can hardly be considered as representative
of the whole Earth.
Other points of view have been expressed about the dino-
saur diversity of this part of the world, the debate being
versity in the last million years of the Cretaceous, several
authors concluding that no decline is apparent [e.g. Sheehan
et al., 1991; Fastovsky et al. 2004; Wang and Dodson,
2006]. Fastovsky et al [2004] have recognized that the Late
Maastrichtian dinosaur diversity outside North America is a
clue to a better understanding of dinosaur extinction sce-
nario but their work is largely focused (like most of the arti-
cles cited above) on the supposed decline of dinosaur
diversity between the Campanian and the Maastrichtian,
which is not our topic here (although, again, this question
cannot be properly answered without using global data).
Archibald and Fastovsky [2004] recognized that “recent
discoveries of Late Cretaceous dinosaurs in South America,
China, Europe, Siberia, and India hold promise as addi-
tional Cretaceous-Tertiary boundary sections, but for now,
we must content ourselves with what is known from the
Western Interior”. This assertion, however, is simply un-
founded as many dinosaurs are now known from Late
Maastrichtian strata across the world. As shown below, the

1. Musée des Dinosaures, 11260 Espéraza, France (jeanleloeuff@yahoo.fr)

Manuscript accepted on April 13, 2012

Bull. Soc. géol. Fr., 2012, no 6

548 LE LOEUFF J.
Western Interior dinosaur record accounts for about 40% of
the global Late Maastrichtian record. A very few authors
have tried to assess the real dinosaur diversity from the
known record. Russell [1995] was the first to use species-
area relationships to estimate real dinosaur biodiversity. He
did not focus on Late Maastrichtian record but obtained a
total diversity of 403 genera for the Campanian stage (ap-
parent diversity: 111 genera in 1995). Wang and Dodson
[2006] used a statistical method, the abundance-based cov-
erage estimator, to suggest that between 213 and 246 dino-
saur genera were present in the whole world for the whole
Maastrichtian stage, but their attempt does not take into ac-
count the palaeobiogeographical context. Their estimate in-
cludes only genera with discoverable record however,
whereas Russell [1995] and this paper try to estimate the to-
tal biodiversity (including genera having no discoverable
record: see discussion in Wang and Dodson [2006]). The
complex palaeobiogeography at the Cretaceous-Tertiary
transition [see Le Loeuff, 1991; 1997] invalidates all hy-
potheses based exclusively on the North American record
(thus most of the previous hypotheses) as at least four major
palaeobiogeographic realms can be recognized, each of
them hosting endemic vertebrate assemblages in several
bioprovinces: the dinosaur biodiversity of less than 6% of
the Earth’s emerged land (western North America) cannot
be representative of the global diversity. The aim of this pa-
per is to provide accurate global data on Late Maastrichtian
geographical and stratigraphical dinosaur distribution, and
tools to assess the real biodiversity from these incomplete
data. The recent data on Late Maastrichtian dinosaurs col-
lected in many places of the world [e.g. Bolotzky and
Godefroit, 2004; Buffetaut et al., 2005; Candeiro et al.,
2008; Case et al., 2000; Godefroit et al., 2000, 2001, 2003,
2004, 2008, 2009; Godefroit and Motchrova-Dekova, 2010;
Laurent et al., 2002; Lockley et al., 2002; Pereda-
Suberbiola et al., 2004, 2009a,b] are summarized below.
METHODS – APPARENT BIODIVERSITY VS
REAL BIODIVERSITY
Signor and Lipps [1982] have emphasized the disparity be-
tween real and apparent biodiversity in the fossil record. In
the case of land vertebrates, taphonomic biases are so im-
portant [cf. Fastovsky and Sheehan, 1997] that they have a
strong incidence on the significance of both the sampled
biodiversity and the sampled stratigraphical distribution
(many dinosaur species are known from a single locality,
which gives an idea of how bad the fossil record is for large
land vertebrates). The apparent (or sampled) biodiversity
(i.e. the number of Late Maastrichtian dinosaur species
known so far) is thus lower than the real biodiversity (i.e.
the number of dinosaur species which really existed during
the Late Maastrichtian). Several methods have been used to
estimate real palaeobiodiversity from the sampled
biodiversity. Some are based on the relationship between
the area of a land mass and the number of animal species
that live upon it [cf. Brown, 1995]. As stated by Barnosky
et al. [2005] the species-area curve is one of the best under-
stood relationships in ecology and it predicts how many
species can be expected as the geographic area of sampling
increases. In other words when the area of a region in-
creases, so does the number of species. This relationship is
Bull. Soc. géol. Fr., 2012, no 6
expressed by S = c Az where S is the number of species, A
is the geographic area (in km2) that was sampled and c and
z are empirically derived constants that express the slope of
the power function and its intercept. Such methods were
originally transposed to palaeontology for estimations of
Late Pleistocene mammalian biodiversity (excluding bats,
rodents and insectivores: cf. Dodson [1990]) rather than for
the more fragmentary Mesozoic data. Russell [1995]
obtained the following relationship for Late Pleistocene
mammals:
Generic diversity = 0.1386 A0.36
Russell [1995] proposed to employ the exponent (0.36) to
calculate intercepts postulated to be representative of dino-
saur diversity, given that the area of emerged lands in the
Late Cretaceous can be estimated from palaeogeographic
maps [e.g. Smith et al., 1994], and that the apparent generic
diversity of the best sampled area in the world (postulated
to represent the real generic diversity) is used to calculate
the intercept.
However there is no reason to postulate that this expo-
nent 0.36 has a general value for land vertebrates. An alter-
native method is to use field data (i.e. lists of taxa per
localities) to establish a species-area relationship for Late
Maastrichtian dinosaurs like Barnosky et al. [2005] did for
Miocene mammals. We obtained the following formula
from the North American data using locality data from the
Paleobiology Database (www.paleodb.org) and faunal lists
from Weishampel et al. [2004] (see also appendix I). The
curve (fig. 1) was produced from a set of faunal lists, suc-
cessively adding species from different areas of western
United States (corresponding geographic areas are the
smallest polygons that enclosed all localities contributing to
a given list).
Species diversity = 1,8093 A 0,2259
This curve does not depend of the chosen palaeo-
geographic map as it is based on the surface of the polygons
enclosing localities, not on the whole surface of the Western
North American subcontinent. However both approaches
[Russell, 1995 and this paper] rely on the same ecological
FIG. 1. – Species-area curve for the Late Maastrichtian. Equation abbrevia-
tions: S = species; A = area.
FIG. 1. – Relation aire-espèce pour le Maastrichtien supérieur. S = espèce ;
A = aire.
 PALEOBIOGEOGRAPHY AND BIODIVERSITY OF LATE MAASTRICHTIAN DINOSAURS 549

relationship and the results obtained by the two methods are New Zealand, Antarctica); a Euramerican (East northern
of the same order of magnitude. We present below (tables I-IV) America, western Europe) community; a Laurasiatic associ-
the results obtained from our species-area curve, which is ation (West northern America, Asia). This geographical
supposed to better reflect an ecological reality. The crucial splitting led to an important endemism in the different is-
point, as is shown below, rather lies in the recognition of lands, with different families occupying the same ecologi-
the bioprovinces, their surface area and the degree of cal niches. A now classical example for the top predators
endemism in the various provinces. was given by Bonaparte and Kielan-Jaworowska [1987]:
large Late Cretaceous meat-eating dinosaurs in northern
America as well as in Asia are tyrannosaurids, i.e. giant
DATA Coelurosaurians whereas in southern America this ecologi-
cal niche is occupied by the family Abelisauridae which are
Late Cretaceous palaeobiogeography Ceratosaurians. Recent investigations have shown that
During the Cretaceous period, original faunal associations abelisaurids were also the top Late Cretaceous predators in
had differentiated on the different landmasses [cf. Russell, India, Africa and Europe [Buffetaut et al., 1988; Le Loeuff
1995; Rage, 1981, 1988; Le Loeuff, 1991, 1997]: a West- and Buffetaut, 1991; Wilson et al., 2003]. Even inside the
Gondwanan association (Africa, South America, India, paleorealms, the different provinces show marked differ-
Madagascar); an East-Gondwanan association (Australia, ences at the generic and/or specific levels, a consequence of
their separation since many million years at the end of the
Mesozoic. Thus, it is irrelevant to try to use data from a sin-
gle landmass to estimate the latest Cretaceous dinosaur
TABLE I. – Low endemism hypothesis based on the paleomap of Smith paleobiodiversity. Late Maastrichtian dinosaurs have been
et al. [1994]. Emerged surfaces of the 10 main Late Maastrichtian biopro- discovered in most areas of the world, as far as correct
vinces, calculated real species diversity; the right column indicates the ap-
parent biodiversity in 2010. datations for continental strata can be obtained. The fossil
TABL. I. – Hypothèse de faible endémisme basée sur la carte paléogéogra- record is very different however for these different Late
phique de Smith et al. [1994]. Surfaces émergées des 10 principales bio- Maastrichtian landmasses: in some places, a very few or
provinces, diversité réelle calculée (nombre d'espèces) ; la colonne de
droite indique la biodiversité apparente en 2010. even no dinosaur bone has been unearthed for different
reasons (lack of Late Maastrichtian continental strata,
Province Surface real app div
2
(km ) div 2010
unexplored strata, etc.).
W. North America (WNA) 6 537 000 63 42
Asia (AS) 2 6149 000 86 23
Europe (EA) 4 503 000 58 18
E. North America (ENA) 8 135 000 66 3 TABLE III. – High endemism hypothesis based on the paleomap of Smith et
al. [1994]. Emerged surfaces of the 21 Late Maastrichtian bioprovinces,
Madagascar (MA) 435 000 34 0 calculated real species diversity.
India (IN) 2 324 000 50 6 TABL. III. – Hypothèse d'endémisme élevé, basé sur la carte paléogéogra-
phique de Smith et al. [1994]. Surfaces émergées des 21 bioprovinces du
South America (SA) 12 493 000 73 7
Maastrichtien supérieur, biodiversité réelle estimée.
Antarctica-Australia (AA) 22 227 000 83 2 2
Province Surface (km )
Africa (AF) 23 389 000 84 2
W. North America (WNA) 6 537 000 63
New Zealand (NZ) 290 000 31 1
Mainland Asia (AS) 26 149 000 86
TOTAL 106 482 000 628 104
Japan (JP) 290 000 31
N. West Asia (NWA) 145 000 26
Scandinavia (SC) 3 777 000 55
Ibero-Armorican Island (IAI) 435 000 34
TABLE II. – Low endemism hypothesis based on the paleomap of Ziegler
et al. [1997]. Emerged surfaces of the 10 Late Maastrichtian bioprovinces, Central Europe (CE) 290 000 31
calculated real species diversity. Hateg region (HA) 100 000 24
TABL. II. – Hypothèse de faible endémisme basée sur la carte paléogéogra-
phique de Ziegler et al. [1997]. Surfaces émergées des 10 bioprovinces du S.E. North America (SENA) 3 922 000 56
Maastrichtien supérieur, biodiversité réelle estimée. N.E. North America (NENA) 2 033 000 48
Province Surface (km2) real diversity Greenland (GR) 2 179 000 49
W. North America (WNA) 8 814 000 67 Madagascar (MA) 435 000 34
Asia (AS) 38 886 000 94 India (IN) 2 324 000 50
Europe (EA) 3 629 000 55 South America (SA) 12 348 000 72
E. North America (ENA) 11 795 000 72 N.W. South America (NSA) 145 000 26
Madagascar (MA) 259 000 30 Antarctica (ANT) 11 476 000 71
India (IN) 2 981 000 52 Antarctic peninsula (ANT2) 290 000 31
South America (SA) 15 814 000 77 Australia (AUS) 10 459 000 70
Antarctica-Australia (AA) 21 517 000 82 West Africa (WA) 5 665 000 61
Africa (AF) 24 369 000 84 East-South Africa (ESA) 17 723 000 79
New Zealand (NZ) 518 000 35 New Zealand (NZ) 290 000 31
TOTAL 128 586 000 648 TOTAL 106 462 000 1 028

Bull. Soc. géol. Fr., 2012, no 6

550 LE LOEUFF J.

TABLE IV. – High endemism hypothesis based on the paleomap of Ziegler in the same province. The ichnological data are taken into
et al. [1997]. Emerged surfaces of the 21 Late Maastrichtian bioprovinces,
calculated real species diversity. account in the same way. A complete list of the taxa is
TABL. IV. – Hypothèse d'endémisme élevé, basée sur la carte paléogéogra- given in appendix I.
phique de Ziegler et al. [1997]. Surfaces émergées de 21 bioprovinces du
Maastrichtien supérieur, biodiversité réelle estimée.
Laurasia
Province Surface (km2)
W. North America (WNA) 8 814 000 67
During the Late Cretaceous, Laurasia comprised North
America and Asia (despite the regression of the Western In-
Mainland Asia (AS) 38 109 000 93
terior Sea during the Late Maastrichtian, eastern North
Western Asia (WAS) 777 000 39
America was possibly still a different palaeobioprovince
N.W. Asia (NWA) 129 000 26 linked to the European archipelago: see Carpenter et al.
Scandinavia (SC) 2 981 000 52 [1997]). In western North America, various Late Maastrichtian
Ibero-Armorican Island (IAI) 259 000 30 formations have yielded 36 dinosaur species, according to
Central Europe (CE) 129 000 26 Weishampel et al. [2004]. This apparent biodiversity of west-
Hareg region (HA) 259 000 30 ern North America is therefore clearly greater than that of
S.E. North America (SENA) 557 3000 60 Montana alone (20 species). Since Weishampel et al.’s paper
N.E. North America (NENA) 583 3000 61 in 2004, a few new taxa have been described [e.g. Bakker
Greenland (GR) 387 000 33
et al., 2006; Farke and Williamson, 2006; Wu et al., 2007;
Madagascar (MA) 259 000 30
Longrich and Currie, 2009]. The 6.537.000 km2 [from Smith
et al., 1994] western North American province has thus
India (IN) 2 981 000 52
yielded 42 dinosaur species (including unnamed species: see
N. South America (NSA) 13 610 000 74
appendix I) in 2010 and is the best sampled area in the world
S. South America (SSA) 2 203 000 49 for this time interval.
Antarctica (ANT) 9 592 000 68
Eastern North American Late Maastrichtian faunas, east
Antarctic peninsula (ANT2) 1 693 000 46 of the Western Interior Sea, are known from the late
Australia (AUS) 10 240 000 69 Maastrichtian New Egypt Formation [Carpenter et al., 1997],
West Africa (WA) 4 277 000 57 which has yielded the theropod Dryptosaurus aquilunguis,
East-South Africa (ESA) 20 091 000 81 Coelosaurus antiquus (Ornithomimidae) and “Hadrosaurus”
New Zealand (NZ) 518 000 35 minor (Hadrosauridae). Thus three species only are known
TOTAL 128 714 000 1 078 from this large part of northern America.
In Asia (the largest part of Laurasia with 26,149,000 to
38,886,00 km2 according to Smith et al. [1994] and Ziegler
et al. [1997], respectively), most of the richest Mongolian
localities are Late Campanian or Early Maastrichtian in age
A short review of Late Maastrichtian dinosaurs
[Jerzykiewicz, 1997; Osmolska, 1997a,b,c]. No Late
Maastrichtian dinosaurs are now known from all continents. Maastrichtian locality is known so far in Mongolia. In
The richest continental deposits spanning the Creta- South China (Nanxiong basin, Guangdong Province), Dong
ceous-Tertiary boundary are exposed in western North [1979] described four dinosaur species from the Yuanpu
America (Canada and United States of America). However Formation, a unit close to the Cretaceous-Tertiary boundary
only some of these Maastrichtian localities can be confi- as a basaltic flow near its top yielded dates averaging
dently assigned to the Late Maastrichtian, either on the ba- 67.4 ma [cf. Russell et al., 1993]: the taxa comprise the
sis of palynological associations or because of their tyrannosaurid Tarbosaurus, the nemegtosaurid Nemegtosaurus
intercalation inside marine deposits. In a few cases, dino- pachi, the therizinosaurid Nanshiungosaurus brevispinus
saur bones have been found in marine rocks of Late and the hadrosaurid Microhadrosaurus nanshiungensis. The
Maastrichtian age: hadrosaurid and theropod bones are overlying Pingling Formation (which contains the Creta-
known from different localities of the Maastricht Formation ceous-Tertiary boundary) yielded at least 6 different dino-
of the Netherlands [see Jagt et al., 2003 for a recent synthe- saurs according to Russell et al. [1993]: small theropods,
sis] or its equivalents in Germany [see Wellnhofer, 1994]. A tyrannosaurids, therizinosaurids, dicraeosaurids (= nemeg-
few radiometric datations have also been used in some parts tosaurids), small ornithopods and hadrosaurids. The precise
of the world such as in the Nanxiong basin of China to pre- age of the Nanxiong dinosaurs has been disputed, some au-
cise the age of Late Cretaceous dinosaur-bearing forma- thors [e.g. Zhao et al., 2002] suggesting a Paleocene age for
tions. Although there are sometimes claims for Paleocene some of the Pingling dinosaurs; however it seems that this
non-avian dinosaurs, all these assertions have been refuted formation is rather well dated of the Late Maastrichtian
[see Ocampo et al., 2006 for a recent review]. The follow- [cf. Russell et al., 1993] but that some reworked Cretaceous
ing review includes the skeletal record with additional data material is mixed with Early Paleocene material [e.g. Buck
from the ichnological record when available (i.e. in South et al., 2004]. We retain a total of 6 dinosaur species for the
America); dinosaur eggs parataxa are not taken into ac- Late Maastrichtian of the Nanxiong basin although studies
count. Unnamed forms are treated as independant “species” on eggshell microstructure suggest up to 8 different forms
when they belong to an unrepresented family in the fossil [cf. Zhao et al., 2002].
record of the palaeobioprovince (e.g. the single theropod Since the late 1990s fieldwork by Pascal Godefroit
tooth from the Late Maastrichtian of Africa is considered as from the Brussels Natural History Museum on the banks of
representing an original species) or when it can be shown the Amur River both on Chinese (Yuliangze Formation of
that they do not belong to a well established species present Jiayin and Wulaga) and Russian (Udurchukan Formation of

Bull. Soc. géol. Fr., 2012, no 6

 PALEOBIOGEOGRAPHY AND BIODIVERSITY OF LATE MAASTRICHTIAN DINOSAURS
Blagoveschensk and Kundur) sides, in far eastern Siberia,
has revealed an extraordinary new dinosaur fauna, which is
dated of the Late Maastrichtian by a palynological assem-
blage [Godefroit et al., 2008]. Following early twentieth
century Russian researchers, Godefroit and colleagues have
described a new Late Maastrichtian hadrosaur-dominated
fauna comprising at least seven distinct hadrosaurids, a
theropod and an ankylosaur [Tumanova et al., 2004].
Alifanov and Bolotsky [2010] have described the alleged
sauropod Arkharavia from the same beds but this species is
apparently based on a few caudal vertebrae which belong to
a very large hadrosaur, not a sauropod. However, rare sauropod
remains have been found from the Udurchkan Formation
(P. Godefroit, pers. comm.). The localities belong to the
Wodehouseia spinata – Aquipollenites subtilis paleozone.
As stated by Godefroit et al. [2004], Wodehouseia spinata
is regarded as a good biostratigraphic indicator for Late
Maastrichtian formations in North America.
In 2009 Godefroit et al. yielded the preliminary results
of their excavations at Kakanaut in northeastern Russia. A
diversified dinosaur assemblage of Late Maastrichtian age
has been recognized with at least seven different species of
dromaeosaurids, tyrannosaurids, troodontids, hadrosaurids,
ornithopods, neoceratopsians and ankylosaurians.
The apparent diversity reaches 23 dinosaur species for
Late Maastrichtian Asia. The whole Asiamerican record
reaches 68 species.
Europe
Europe was an archipelago during the Late Cretaceous with
an important degree of endemism on the different islands as
most dinosaurs are generically different between Romania
and western Europe for example. Palaeogeographic maps
show that 4 main islands existed in the Maastrichtian: an
Ibero-Armorican island on France and Spain, a Central Eu-
ropean island, a British island and a very large island on
Scandinavia and northeastern Europe (Fenno-Scandian
island). Additional smaller islands were located closer to
the Black Sea in Crimea, Bulgaria and Romania, which pre-
cise extension is obscured by the subsequent Alpine thrusts
[see Jianu and Boekshoeten, 1999].
Late Maastrichtian marine deposits of northern Europe
(The Netherlands and Germany) have yielded the theropod
Betasuchus bredai (possibly related to Dryptosaurus
aquilunguis) and the hadrosaurid “Orthomerus dolloi” [see
Jagt et al., 2003; Wellnhofer, 1994 for a discussion on the
age of the specimens]. In a recent review, Jagt et al. [2003]
concluded that “at least one type of theropod and more than
one taxon of non-lambeosaurine hadrosaurid as well as a
possible euhadrosaurian are represented in this material”.
Thus the remains of at least 4 different taxa are represented
close to the Central European island.
Isolated finds from Crimea (a hadrosauroid dinosaur
named “Orthomerus weberi” by Riabinin [1945]; see Le
Loeuff [1992]) and recently from Bulgaria (a possible
ornithomimosaur according to Mateus et al. [2009] and a
hadrosaur described by Godefroit and Motchurova-Dekova
in 2010) show that there is some potential for more Late
Maastrichtian dinosaur discoveries around the Black Sea
where three taxa are represented by isolated bones.
In southern France and northern Spain (a third island of
the European archipelago), the richest Late Cretaceous
551
localities are Late Campanian and Early Maastrichtian
[Buffetaut and Le Loeuff, 1991; Buffetaut et al., 1991, 1997;
Le Loeuff, 1992; Le Loeuff and Buffetaut, 1995] and are
characterized by Titanosaur-Rhabdodon dominated assem-
blages. However, recent excavations of Late Maastrichtian
localities have led to the discovery of another hadrosaur-
dominated assemblage [Le Loeuff et al., 1994] comprising
at least five hadrosauroid taxa (including Pararhabdodon
isonense, Koutalisaurus kohlerorum and Arenysaurus
ardevoli [Pereda-Suberbiola et al., 2009a,b; Le Loeuff et al.,
1993; Laurent et al., 1997]), a dromaeosaurid [Le Loeuff
and Buffetaut, 1996] and at least two additional small
theropods including “Euronychodon”-like teeth [Lopez-
Martinez et al., 2001], a large theropod, an ankylosaur and
at least one saltasaurid sauropod [Laurent et al., 2002 ;
Canudo et al., 1999]. Localities in southwestern France are
well dated with benthic foraminifers [Laurent et al., 2002]
and can be safely assigned to the Late Maastrichtian. There
is no consensus about the age of some Spanish localities
considered as Early Maastrichtian by Lopez-Martinez et al.
[2001], an opinion refuted by Laurent et al. in 2002. Eleven
different species are thus known from this island.
In Romania, 11 different dinosaurs species are known
from the Hateg basin during the Maastrichtian. Although
the age of the Sanpetru Formation was usually thought to be
Late Maastrichtian, recent studies [e.g. Panaiotu and Panaiotu,
2002] suggest a slightly older age (Early Maastrichtian),
thus these data are not used in this paper.
The Fenno-Scandian island has yielded only Campanian
dinosaurs referred by Lindgren et al. [2007] to the family
Leptoceratopsidae, a clade previously unknown in Europe.
This suggests a dinosaur assemblage quite different from
the South-European fauna.
The European apparent dinosaur biodiversity in the
Late Maastrichtian therefore amounts to 18 species (includ-
ing unnamed species).
Western Gondwana
In Africa, most Late Cretaceous localities are Cenomanian
in age [Cavin et al., 2010]. Maastrichtian Formations from
Niger have yielded a few dinosaur bones [Taquet, 1976] but
their precise age (Early or Late Maastrichtian) is unknown.
The recent discovery of dinosaur bones in the Late
Maastrichtian phosphates of the Ouled Abdoun basin of
Morocco [Pereda-Suberbiola et al., 2004; Buffetaut et al.,
2005] is thus so far the only evidence of latest
Maastrichtian African dinosaurs. The age of these deposits
is well constrained by selachian teeth [Noubhani and Cappetta,
1997]. The dinosaurian remains consist of fragmentary re-
mains of a sauropod referred to the Titanosauriformes by
Pereda-Suberbiola et al. [2004] and a theropod dinosaur
tooth referred by Buffetaut et al. [2005] to the family
Abelisauridae. This is so far the meagre record for African
Late Maastrichtian dinosaurs.
In India, the Lameta Formation is directly overlain by
the Deccan Trapps, the emplacement of which seems to
have begun in magnetozone 29R [see Buffetaut, 1987]. The
Lameta Formation is thus supposed to be of Late
Maastrichtian age. Its dinosaur assemblage described by Huene
and Matley in 1933 has been revised in recent years. Two
sauropods species are considered as valid: the saltasaurids
Isisaurus colberti (JAIN & BANDYOPADHYAY, 1997) and
Bull. Soc. géol. Fr., 2012, no 6
552 LE LOEUFF J.
Jainosaurus septentrionalis (HUENE & MATLEY 1933); Wil-
son et al. [2003] described the new abelisaurid theropod
Rajasaurus narmadensis. Wilson and Mohabey [2006] con-
sider that three large theropod species (including the
abelisaurids Indosaurus and Indosuchus), one small-bodied
theropod (possibly a fourth abelisaurid) and at least two
sauropods are present in these horizons.
In South America, most Late Cretaceous localities are
ante-Maastrichtian in age. However, Dias-Brito et al.
[2001] have referred the uppermost formation of the Bauru
Group of Minas Gerais State in Brazil (the Marilia For-
mation) to the Late Maastrichtian on the basis of a
charophytes-ostracods association. This unit has yielded
several dinosaur taxa, including three saltasaurids
(Baurutitan britoi KELLNER et al., 2005; Trigonosaurus
pricei CAMPOS et al., 2005; Uberabatitan riberoi SALGADO
& CARVALHO, 2008) and three theropods: a possible
elmisaurid [Novas et al., 2005], an abelisaurid [Candeiro
et al., 2008; Novas et al., 2008] and a possible
carcharodontosaurid [Candeiro et al., 2008; Candeiro and
Tanke, 2008; Candeiro, 2009]. Meyer and Lockley [1999]
have described a dinosaur megatracksite from the Late
Maastrichtian El Molino Formation of Bolivia [see also
Lockley et al., 2002]. They recognized at least five different
ichnotaxa (two theropods, one ankylosaur, two sauropods).
From these ichnotaxa, the ankylosaur is retained here as
this group has no skeletal record in the Late Maastrichtian
of South America.
This gives a total of 15 dinosaur species for West
Gondwanan provinces (including one ichnotaxon) with
Abelisauridae, Carcharodontosauridae,?Elmisauridae, Salta-
sauridae, Ankylosauria.
East Gondwana
The first dinosaur bone from the Late Maastrichtian of Aus-
tralia is a theropod humerus from the marine Miria Forma-
tion of Giralia range, western Australia [Long, 1992, 1998].
Its age is well calibrated by foraminifers and ammonites
[Bennett and Long, 1991]. In New Zealand, the Upper
Maungatuniwka Sandstone is of Early Maastrichtian age
and yielded various dinosaurs [Wiffen, 1996; Molnar,
1997]. A recent find in the Takatika Grift of the Chatham
island, 900 km east of New Zealand was reported by
Stilwell et al. [2006] who have described several bones of a
large unidentified theropod in this unit spanning the Creta-
ceous-Tertiary boundary. The discovery of a Late
Maastrichtian hadrosaur tooth from Antarctica was reported
by Case et al. [2000] in the Lopez de Bertodano Formation,
a marine unit of Vega island (Antarctic peninsula) well
dated by ammonites and palynomorphs. Other Late Creta-
ceous finds from Antarctica come from older strata of
Campanian to early Maastrichtian age [Salgado and
Gasparini, 2006; Hooker et al., 1991] with the exception of
a partial hadrosaur bone [Rich et al., 1999]. The East
Gondwanan provinces have thus yielded only three Late
Maastrichtian dinosaur species from unexpectedly remote
places.
Late Maastrichtian apparent biodiversity
Four palaeobiogeographic realms and a minimum of 11 prov-
inces ( Asia, eastern North America, western North
Bull. Soc. géol. Fr., 2012, no 6
America, Europe, India, Madagascar, Africa, South Amer-
ica, Antarctica, Australia, New-Zealand) existed in the Late
Maastrichtian. Their area was comprised between 0.3 and
28 million square kilometres. All these provinces have now
yielded Late Maastrichtian dinosaurs totalising 104 species
(including yet unnamed species and one unnamed ichno-
species). This is the apparent global dinosaur biodiversity at
the end of the Maastrichtian in 2010 (in 1975 it amounted to
about 35 genera; in 2000 to 59 genera according to Le
Loeuff and Laurent [2000]).
INTERPRETATION OF THE DATA – ESTIMATION
OF THE REAL BIODIVERSITY
We have applied our species-area relationship to two hy-
potheses of the palaeobiogeography of the Latest Creta-
ceous. The surfaces of landmasses have been calculated
from the Maastrichtian palaeogeographic maps of Smith
et al. [1994] (see fig. 2) and Ziegler et al. [1997] (see fig. 3)
using the software PDF-XChange Viewer (tool “area”).
Both maps are Mollweide equal area projections which sac-
rifices fidelity to angle and shape in favour of accurate de-
piction of area. These reconstructions differ from each
other in many details with a total of emerged lands of
106,462,000 km2 in Smith et al’s map versus
128,582,000 km2 in Ziegler et al. [1997]. However the re-
sults do not change significantly whether one map or the
other is used (compare tables I and II and tables III and IV).
Low endemism hypothesis
In this hypothesis ten bioprovinces only are recognized
(tables I and II; figures 2a and 3a) during the Late
Maastrichtian. Our relationship gives a real biodiversity of
628 species with the map of Smith et al. [1994] and 648
species using the map of Ziegler et al. [1997].
If all late Maastrichtian dinosaurs species have been
found from Montana and the Dakotas (an unlikely hypothe-
sis) the global Late Maastrichtian dinosaur generic
biodiversity would be situated between 628 and 648 spe-
cies. The Late Maastrichtian fossil record would thus be
16% complete, which is in good agreement with former es-
timations of the total dinosaur fossil record (10 to 28%
complete according to Holmes and Dodson [1997]). It can
also be noticed that the northern hemisphere records ac-
counts for 83% of the observed biodiversity and that south-
ern data are still extremely scanty.
High endemism hypothesis
It is likely that these are under-estimates, as we have
over-simplified the Late Maastrichtian palaeogeography
(note that a 24,000,000 square kilometres area would con-
tain 84 species, but two 12,000,000 square kms areas would
contain 72 species each, i.e. a total of 144 species). It may
be more realistic to consider that the Late Maastrichtian
world was more partitioned. The example of western Eu-
rope suggests that this may be correct as the data from the
different islands show a strong endemism: most if not all
species are different between the “Hateg island” and the
“Ibero-Armorican island”. The few available data from
Scandinavia [e.g. Lindgren et al., 2007] also suggest a dino-
saur assemblage extremely different from what is known in
 PALEOBIOGEOGRAPHY AND BIODIVERSITY OF LATE MAASTRICHTIAN DINOSAURS 553

southern and eastern Europe. Thus in a second hypothesis
we have opted for a more provincialized world, shifting
from 10 palaeobioprovinces to 21, a pattern consistent with
the palaeogeographical maps (figs 2b and 3b). The species-
area relationship presented above is not altered.
Based on the map of Smith et al. [1994] we obtain a
real diversity of 1028 species. An alternative calculation
was also made after the larger landmasses of Ziegler et al.
[1997] and suggests a very slightly higher diversity of 1078
species.

FIG. 2. – Late Maastrichtian palaeogeography [from Smith et al., 1994]

Abbreviations for top: WNA: western North America; AS: Asia; EA: European archipelago; ENA: eastern North America; AF: Africa; MA: Madagascar;
IN: India; SA: South America; AA: Antarctica-Australia; NZ: New Zealand.
Abbreviations for bottom: WNA: West North America; SENA: southeastern North America; NENA: northeastern North America; GR: Greenland; AS:
Mainland Asia; JP: Japan; NWA: North West Asia; SC: Scandinavia; IAI: Ibero-Armorican Island; CE: Central Europe; HA: Hateg Island; WA: western
Africa; ESA: East and South Africa; MA: Madagascar; IN: India; SA: South America; NSA: northern South America; ANT: Antarctica; ANT2: Antarctic
Peninsula; AUS: Australia; NZ: New Zealand.
FIG. 2. – Paléogéographie du Maastrichtien supérieur [d'après Smith et al., 1994].

Bull. Soc. géol. Fr., 2012, no 6

554 LE LOEUFF J.

In this second hypothesis the dinosaur fossil record
would be 10% complete. Our results are an order of magni-
tude above previous estimates [Wang and Dodson, 2006;
Russell, 1995]. The lessons of modern studies on the
areography of recent species suggest that the majority of
species are absent from most places and where they do oc-
cur they are usually quite rare [e.g. Gaston, 2003]. We
suppose that the same pattern was true during the Mesozoic
and thus strongly suggest that our hypotheses are more real-
istic than previous estimates. As remarked by P. Godefroit
(pers. comm.) it is remarkable that within a relatively small
area the tree sedimentary basins he sampled in eastern Asia
have yielded three different hadrosaur associations from the
same palynozone. It can also be noted that the total surface

FIG. 3. – Late Maastrichtian palaeogeography [from Ziegler et al., 1997]

Abbreviations for top: same than fig. 1.
Abbreviations for bottom: WNA: West North America; SENA: southeastern North America; NENA: northeastern North America; GR: Greenland;
AS: mainland Asia; WAS: western Asia; NWA: northwestern Asia; SC: Scandinavia; IAI: Ibero-Armorican island; CE: Central Europe; HA: Hateg island;
WA: western Africa; ESA: East and South Africa; MA: Madagascar; IN: India; NSA: northern South America; SSA: southern South America; ANT:
Antarctica; ANT2: Antarctic Peninsula; AUS: Australia; NZ: New Zealand.
FIG. 3. – Paléogéographie du Maastrichtien supérieur [d'après Ziegler et al., 1997].

Bull. Soc. géol. Fr., 2012, no 6

 PALEOBIOGEOGRAPHY AND BIODIVERSITY OF LATE MAASTRICHTIAN DINOSAURS
of emerged land does not significantly change the real
biodiversity but that the number of provinces strongly influ-
ences the global counts (compare tables I and III and tables
II and IV, respectively).
Our results are testable in two ways: further analyses of
the North American record may allow to precise the real
biodiversity of this crucial province and may help to estab-
lish a stronger species-relationship area. Excavations at
poorly known localities in every province may help to assess
the degree of endemism in the various islands to test our hy-
pothesis of a 100% endemic assemblage in each province.
CONCLUSIONS
104 dinosaur species have been reported from Late
Maastrichtian strata around the world in 2010. Real dino-
saur biodiversity in the Late Maastrichtian (i.e. in a 2.8 Ma
time interval between 68.3 and 65.5 m.y.) would be situated
between 628 and 1078 species. Therefore all discussions
about dinosaur extinction should be based on this estimated
real biodiversity and not on the apparent biodiversity of a
single area. One important conclusion is that the North
American record represents only 43% of the apparent dino-
saur biodiversity in the Late Maastrichtian, thus analyses
based on North American data will miss almost two-thirds
of the available data. It is suggested that the over-represen-
tation of the North American dinosaurs is simply linked to a
collecting bias, as the Late Cretaceous localities from this
part of the world have been extensively excavated since the
1870s. Our results are almost an order of magnitude above
Wang and Dodson’s estimations (i.e. between 213 and 246
species for the whole Maastrichtian stage), a result of the
over-simplification of the palaeogeographical framework by
these authors. Fastovsky et al. [2004] concluded that “both
South America and Asia have much to contribute to the
question of late Cretaceous dinosaur diversity and that the
key to better understanding Mesozoic dinosaur richness is
not collecting more dinosaurs but obtaining better temporal
control on the ones we already have”. A better temporal
control is indeed extremely important but new data from
outside Northern America are even more crucial to help to
define palaeobioprovinces of the Cretaceous-Tertiary
transition and their degree of endemism, which will allow
more precise estimates of the real dinosaur biodiversity.
Work on Maastrichtian marine invertebrates [cf. a sum-
mary in Jablonski, 1997] suggests that they suffered a bru-
tal extinction at the Cretaceous-Tertiary boundary, partially
APPENDIX 1: LIST OF TAXA
Data compiled from Weishampel et al. [2004] have no reference added
Late Maastrichtian
Western North America (42 species)
Coelurosauria Inc. Sed.: Ricardoestesia gilmorei, cf. R. isosceles;
Ornithomimidae: Ornithomimus velox; Oviraptorosauria: Caenagnathus
sp., Chirostenotes elegans; C. pergracilis; Alvarezsauridae indet.
555
obscured in the fossil record by a Signor-Lipps effect. As
the last observed occurrence of a species rarely coincides
with its real extinction, the sampled fossil distributional
pattern after a sudden extinction will tend to look like that
of a gradual extinction. The same hypothesis may be ap-
plied to continental vertebrates, taking into account that
their fossil record is much worse than that for marine ani-
mals. Given that many of the Late Maastrichtian dinosaur
species were sampled at a single locality or formation, it is
possible to use the mean duration of dinosaur genera to es-
tablish their probable distribution. The total duration of the
Maastrichtian stage is estimated at 5.1 m.y. [70.6-65.5 Ma]
according to Gradstein et al. [2008]. The Late Maastrichtian
begins with the CF4 planktic foraminiferal zone 68.3 Ma
ago [cf. Li et al., 1999], thus the total duration of the Late
Maastrichtian substage is of about 2.8 million years. Given
that the mean duration of a dinosaurian genus was calcu-
lated by Dodson [1990] to be of 7.7 m.y., the presence of a
dinosaur species in the short Late Maastrichtian time inter-
val strongly suggests that it experienced the crucial Creta-
ceous-Tertiary events. Thus, the Late Maastrichtian dinosaur
stratigraphical distribution is consistent with the termina-
tion of most of the lineages (i.e. between 628 and 1078 spe-
cies) at the Cretaceous-Tertiary boundary. In the present
state of knowledge, the hypothesis of a sudden impact-re-
lated extinction of non-avian dinosaurs at the Cretaceous-
Tertiary boundary is supported by our data: speculations
about a random extinction of a handful of dinosaur species
are no more tenable as it is a large and diversified group of
land vertebrates which disappeared at the Cretaceous-Ter-
tiary boundary.
Beyond demonstrating that hundreds or thousands of di-
nosaur species went extinct at the Cretaceous-Tertiary
boundary, our results also show that the global diversity of di-
nosaur during the Mesozoic (1850 genera for Wang and
Dodson [2006]; 3400 genera for Russell [1995]) has been
largely underestimated as the palaeobiogeographical context
has not been enough taken into account. The use of spe-
cies-area relationships at various geographic levels (regional,
continental, global) may provide new insights on Mesozoic
biodiversity and help to recognize long term ecological trends.
Acknowledgements. – This work has been inspired by Jean-Claude Rage’s
papers on the palaeobiogeography of Mesozoic and Cenozoic vertebrates. I
thank Eric Buffetaut and Jean-Sébastien Steyer for inviting me to contri-
bute to this volume honouring Jean-Claude’s stimulating work. Pascal Go-
defroit and Julien Claude provided useful reviews which greatly improved
this paper. Special thanks to Julien Claude for his invaluable assistance in
the realization of figure 1.
(LONGRICH & CURRIE, 2009b); Tyrannosauridae: Tyrannosaurus rex,
Nanotyrannus lancensis, Albertosaurus sarcophagus, Aublysodon
mirandus; Dromaeosauridae: Dromaeosaurus albertensis, cf. Saurorni-
tholestes langstoni; Troodontidae: Troodon formosus; Saltasauridae:
Alamosaurus sanjuanensis; Euornithopoda: Thescelosaurus neglectus,
Parksosaurus warreni, Bugenasaura infernalis; Hadrosauridae:
Edmontosaurus regalis, E. annectens, E. sa-skatchewanensis,
Parasaurolophus walkeri, Anatotitan copei; Nodosauridae:
Edmontonia cf. rugosidens, E. longiceps, Glyptodontopelta mimus;
Bull. Soc. géol. Fr., 2012, no 6
556 LE LOEUFF J.
Ankylosauridae: Ankylosaurus magniventris; Pachycephalosauridae:
Pachycephalosaurus wyomingensis, Stegoceras edmontonense, Stygi-
moloch spinifer, Sphaerotholus buchholtzae, S. goodwini, Dracorex
hogwartsia (BAKKER et al., 2006); Neoceratopsia: Leptoceratops
gracilis, Montanaceratops sp.; Ceratopsidae: Triceratops horridus,
Torosaurus latus, T. cf. utahensis (HUNT AND LEHMAN, 2008);
Pentaceratops sp., Diceratops hatcheri, Eotriceratops xerinsularis
(WU et al., 2007), Chasmosaurinae indet. [Farke and Williamson,
2006].
Hell Creek Formation, Montana (70000 km2), 20 species
Hell Creek Formation, Montana-Dakotas (140000 km2), 28 species
Lance Formation, USA (330000 km 2), 30 species
Late Maastrichtian formations, Montana (100.000 km 2), 25 species.
Asia (22 species)
Nanxiong basin: Theropoda indet. [Russell et al., 1993];
Tyrannosauridae: Tarbosaurus sp.; Therizinosauridae: Nanshiun-
gosaurus brevispinus; Nemegtosauridae: Nemegtosaurus pachi;
Hadrosauridae Microhadrosaurus nanshiungensis (DONG, 1979);
Ornithopoda indet. [Russell et al., 1993].
Amur River [Godefroit et al., 2008] (P. Godefroit, pers. comm):
Theropoda indet. ; Sauropoda indet. ; Hadrosauridae: Olorotitan
arharensis ; Kerberosaurus manakini; Amurosaurus riabinini ;
Charonosaurus jiayinensis; Sahaliyania elunchunorum ; Wulaga-
saurus dongi; unnamed hadrosaurid. Ankylosauridae indet.
[Tumanova et al., 2004].
Kakanaut [Godefroit et al., 2009]: Dromaeosauridae indet.;
Tyrannosauridae indet.; Troodontidae indet.; Hadrosauridae indet.;
Ornithopoda indet.; Neoceratopsia indet.; Ankylosauria indet..
Eastern North America (3 species)
Theropoda indet.: Dryptosaurus aquilunguis; Ornithomimidae:
Coelosaurus antiquus; Hadrosauridae: “Hadrosaurus” minor
Europe (17 species)
Netherlands/Germany: Theropoda indet.: Betasuchus bredai;
Hadrosauridae: 3 unnamed species [Jagt et al., 2003].
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