213

The Geological Society of America

Special Paper 505
2014
What the dinosaur record says about extinction scenarios
J. David Archibald*
Department of Biology, San Diego State University, San Diego, California 92182-4614, USA
ABSTRACT
The record of dinosaurs over the last 10 m.y. of the Cretaceous, as well as sur-
rounding the Cretaceous-Paleogene boundary, helps to dene extinction scenarios.
Although Late Cretaceous dinosaur fossils occur on all present-day continents,
only in North America do we nd a terrestrial vertebrate fossil record spanning the
Cretaceous-Paleogene boundary, although promising work may yield comparable
records in South America, India, China, and Europe. For the present then, the North
American record represents the proxy for our knowledge of dinosaur extinction. Over
the last 10 m.y. of the Cretaceous (late Campanian to late Maastrichtian) in the north-
ern part of the western interior of North America, the number of nonavian dinosaur
species dropped from 49 to 25, almost a 50% reduction, even though a 16% greater
extent of fossil-bearing exposures record the last dinosaurs in the latest Cretaceous
in the western interior. Important, but less-well-exposed, nonavian-dinosaurbearing
units suggest this drop occurred around, or at least commenced by, the Campanian-
Maastrichtian boundary. These losses began during climatic uctuations, occurring
during and possibly in part caused by the last major regressive cycle of the Creta-
ceous, which also reduced the expanse of the low coastal plains inhabited by nonavian
dinosaurs. The pulse of Deccan Trap emplacement that began some time later in the
latest Cretaceous was also likely a major driver of climatic change. As for the dinosaur
record near the Cretaceous-Paleogene boundary, even the best-known records from
North America remain enigmatic and open to interpretation. Newer studies suggest
some decline in at least relative abundance approaching the Cretaceous- Paleogene
boundary, but the cause (or causes) for the nal extinction (if it was the case) of non-
avian dinosaurs remains unresolved, although the Chicxulub impact undoubtedly
played a major role.
*darchibald@mail.sdsu.edu
Archibald, J.D., 2014, What the dinosaur record says about extinction scenarios, in Keller, G., and Kerr, A.C., eds., Volcanism, Impacts, and Mass Extinctions:
Causes and Effects: Geological Society of America Special Paper 505, p. 213224, doi:10.1130/2014.2505(10). For permission to copy, contact editing@geosociety.
org. 2014 The Geological Society of America. All rights reserved.
INTRODUCTION
The best-preserved and most-studied sequences of Late Cre-
taceous and Paleocene fossil vertebrates occur in western North
America. From the latest Cretaceous of western North America
alone, over 100 vertebrate species are known, representing all
major vertebrate clades. Although nonavian dinosaurs repre-
sent at most only 20% of this taxonomic diversity, interest in the
possible causes and patterns of their extinction far outweighs
the interest for the extinction of other species, vertebrate or not
(Archibald, 1996). The study of dinosaur extinction is certainly
not the most conducive for general understanding of the processes
on August 25, 2014 specialpapers.gsapubs.org Downloaded from
214 Archibald
of extinction, quite simply because dinosaur population sizes and
the environments in which they lived did not greatly favor pres-
ervation in the fossil record, with some very notable exceptions.
Unfortunately, there are no such notable exceptions for any sites
near the end of the Cretaceous.
Can any evidence be offered relating to the question of what
the dinosaur record says about extinction scenarios, which the
organizers of this volume posed as a topic to be addressed. The
answer is a qualied yes. Specically, the two major issues
to be explored are: (1) What is the nonavian dinosaur record
throughout the later Cretaceous, approximately the last 10 m.y.?
(2) What is the nonavian dinosaur record like near the end of
the Cretaceous, say the last half-million years leading up to the
Cretaceous-Paleogene boundary? The emphasis here is by neces-
sity on these records and less on the possible extinction scenarios,
quite simply because it is easier to examine what the nonavian
dinosaur record is rather than why it is this way. The questions
of why are examined in the instances in which the patterns of
change in nonavian dinosaur taxonomic diversity seem to address
extinction scenarios.
THE LAST TEN MILLION YEARS
Most exposures in Dinosaur Provincial Park, Alberta, Can-
ada, range from 76 to 74 m.y. old, representing the later part of
the late Campanian portion of the Late Cretaceous (Currie and
Koppelhus, 2005). In western North America, terrestrial, geo-
logical sections based on vertebrate fossils of this age also are
known as the Judithian after the Judith River Formation, rst rec-
ognized in Montana, a partial equivalent of the Dinosaur Park
Formation in Canada.
Ornithischian and saurischian dinosaurs both occur com-
monly in the late Campanian (Judithian)aged sediments of
Dinosaur Park. Saurischians are limited to theropods, the group
including both birds and the mostly large to extremely large
ground-dwelling, bipedal, mostly carnivorous dinosaurs. Sauro-
pod saurischians do not make it this far north in the Late Cre-
taceous, being known only as far north as southern Wyoming.
Thus, the Dinosaur Park nonavian dinosaur fauna can best be
grouped as ornithischians and theropods, in which ornithischi-
ans outnumber theropods by four to one. Among ornithischians,
~50% are duckbilled hadrosaurs, 25% are horned ceratopsians,
and the remaining 25% mostly represent armored ankylosaurs
and bone-headed pachycephalosaurs. In the park, the ornithischi-
ans comprise three genera of ankylosaurs, each with one species,
three species of the pachycephalosaur genus Stegoceras, six spe-
cies of ceratopsians belonging to four different genera, and eight
hadrosaurs arrayed in six different genera. Hypsilophodontidae
(a paraphyletic group including basal neornithischians and basal
ornithopods) is known only from isolated teeth in the park and
thus cannot be clearly assigned to a genus or species (Table 1;
references in Currie and Koppelhus, 2005; Archibald, 2011).
Theropods make up less than 10% of recovered fossils but
are taxonomically quite diverse. Of the 14 families (39 genera)
of dinosaurs recognized in the park, 42% of these families (40%
of the genera) are theropods (excluding birds). Thus, what thero-
pods lack in numbers of specimens some 75 m.y. ago (Ma), they
make up in diversity (Currie, 2005). This pattern of low speci-
men abundance but higher taxonomic diversity is echoed some
10 m.y. later, just before nonavian dinosaur extinction.
It would be best to possess good samples of nonavian dino-
saur taxa at half-million-year intervals, but this is unfortunately
not the case in North America. To have comparably well-sampled
dinosaur faunas, we must jump to the latest Cretaceous, or the
late Maastrichtian, which in North America is also called the Lan-
cian, the terrestrial vertebrate equivalent of this age, named after
the type Lance Formation of eastern Wyoming. The taxonomi-
cally most diverse Lancian-aged nonavian dinosaur fauna, with
18 species, is perhaps known from the type Lance Formation.
Its upper limit is essentially the Cretaceous-Paleogene boundary
at roughly 66 Ma. The base of the Lancian at 69 Ma is less well
constrained (see Cifelli et al., 2004; Wilson, 2005).
Comparing the Dinosaur Park and Lance Formations, the
former represents about 2 m.y. and the latter between 2 and 3 m.y.,
respectively. Being terrestrial, largely uvially derived units, we
must keep in mind that although both formations span the times
indicated, they do not preserve events through the entirety of
these interval, but rather represent time-averaged packets. Most
terrestrially derived fossils are from such time-averaged occur-
rences. Both formations represent similar depositional histo-
ries, preserving oodplain and coastal uvial deposits draining
easterly into the Pierre or Western Interior Seaway. The western
continent of Laramidia lay to the west, and the eastern continent
of Appalachia lay to the east of this seaway (Archibald, 1996),
transgressing and regressing some four times during the last
35 m.y. of the Late Cretaceous (Fig. 1). The last incarnation of
this epicontinental sea at its greatest width might have reached
1000 km wide and up to ve times as long. Such seaways inun-
dated almost all continents during the Late Cretaceous, equiva-
lent to an area probably approximately the size of modern Africa.
Nothing comparable in size exists today, although Hudson Bay,
the North Sea, the Baltic Sea, and the Red Sea all have been pro-
posed as smaller examples.
In a comparison of the nonavian dinosaur faunas, the Lance
Formation, although quite rich, does not match the older Dinosaur
Park Formation nonavian dinosaur fauna: Dinosaur Park has 39
species, whereas the Lance Formation has 18 species. This means
a drop of over 50% species abundance during the last 10 m.y.
of the Cretaceous in North America (Table 1). Additional Judi-
thian- and Lancian-aged nonavian dinosaur faunas in western
North America add further support in showing this decline is real.
Other major Judithian-aged formations in the northern western
interior are the Judith River and Two Medicine Formations in
central Montana; the other major formation of Lancian age is the
Hell Creek Formation of eastern Montana and western North and
South Dakota. Adding these additional records to augment the
Dinosaur Park and Lance Formations, we nd a loss of dinosaur
species of almost 50%, still a considerable decline (Table 1).
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What the dinosaur record says about extinction scenarios 215

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216 Archibald
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on August 25, 2014 specialpapers.gsapubs.org Downloaded from
What the dinosaur record says about extinction scenarios 217
Some have argued that the Dinosaur Park Formation and
Lance Formation are not ecologically equivalent but have offered
no data to support this claim, going so far as to call such an assess-
ment absurd (Fastovsky et al., 2005). Others have even attempted
without success to statistically discount this very real decline
(Fastovsky et al., 2005; Wang and Dodson, 2006; Lloyd et al.,
2008). In agreement with the results presented here, a study by
Barrett et al. (2009) compiled genus-richness estimates for three
Mesozoic dinosaur clades, noting a marked decline in dinosaur
generic richness in the nal stages of the Cretaceous, which indi-
cates that the clade decreased in diversity for several million years
Figure 1. (A) North American biogeography during the latter part of the
Cretaceous showing the major western continent of Laramidia and the
eastern continent of Appalachia separated by the epicontinental Pierre
Seaway. (B) The same region near the Cretaceous-Paleogene boundary
showing the Brazos River Embayment. (C) The same region near the
mid-Paleocene showing the Cannonball Sea (after Archibald, 2011).
before its nal extinction at the Cretaceous-Paleogene boundary.
The authors of this study show that many uctuations in dino-
saur diversity through the Mesozoic found in previous studies
are likely not real but are artifacts of fossil preservation. These
authors further found that when biases in fossil preservation are
taken into account, several genuine diversity signals are present;
the most intriguing of these, according to these authors, shows
that both ornithischian and theropod dinosaur lineages began to
decline several million years prior to the Cretaceous-Paleogene
extinction event, again in accord with the results discussed here.
In 2012, Brusatte et al. (2012) calculated what they termed
the morphological disparity or anatomical variability found in
seven major dinosaur groups during the latest Cretaceous. They
found both geographic and clade-specic heterogeneity. The
morphological disparity declined for large-bodied herbivores
(ceratopsids and hadrosauroids) and some North American taxa
during the nal two stages of the Cretaceous. At the same time,
carnivorous dinosaurs, mid-sized herbivores, and some Asian
taxa did not decrease in morphological disparity. These authors
argued that the pattern of Late Cretaceous dinosaur evolution was
complex and that there was no universal biodiversity trend. As
with earlier studies, Brusatte et al. (2012) found that some dino-
saur groups endured long-term declines in morphological vari-
ability before their nal extinction at the end of the Cretaceous.
The data presented here as well as by Barrett et al. (2009) and
Brusatte et al. (2012) indicate that the Late Cretaceous vertebrate
fossil record in western North America is quite good and by all
measures is reliable in providing proportional changes, if not abso-
lute changes, between Judithian and Lancian, and Lancian and Cre-
taceous-Paleogene boundary sites. Nevertheless, Fastovsky et al.
(2005, e75) repeated one of the common canards against compari-
son of the Dinosaur Park and Lance Formation nonavian dinosaur
faunas: There is no reason to suppose that the richest formations of
each time interval are spatially equivalent or preserve faunas with
equal delity. If by a possible lack of spatial equivalence of the
rock units yielding these dinosaurs bones, they mean the two are not
equivalent in areal exposure, they are correct, but certainly not in the
way they implied. The argument here basically is that if you have
more exposure, everything else being equal, you should also have
more species of dinosaurs preserved. Following their reasoning, if
the much greater taxonomic richness of the Dinosaur Park Forma-
tion could be attributed to a greater geologic exposure, these authors
might have a case. Such is in fact, not the case.
Eberth (2005, p. 58) noted that the bedrock exposures in
the [Dinosaur Provincial] Park comprise Canadas largest area of
badlands, covering approximately 75 km
2
. The Dinosaur Park
on August 25, 2014 specialpapers.gsapubs.org Downloaded from
218 Archibald
Formation comprises a large portion of this area, and as noted
above, has produced 39 species of nonavian dinosaur. More
broadly, the Dinosaur Park Formation in Alberta and Saskatch-
ewan plus the equivalently aged Judith River and Two Medicine
Formations to the south in Montana are mapped as covering
something like 48,000 km
2
(Archibald, 2011). Doing this, an
additional 10 non avian dinosaur taxa can be added the 39 already
known in and near Dinosaur Provincial Park, giving a total of 49
late Campanian (Judithian) nonavian dinosaurs from these three
formations. By comparison, the late Maastrichtian (Lancian)-
aged Lance Formation of Wyoming and the Hell Creek Forma-
tion of eastern Montana and western North and South Dakota are
mapped as covering at least 57,000 km
2
; yet at most ve addi-
tional dinosaur species can be added, yielding perhaps 25 spe-
cies of nonavian dinosaurs for this interval (Archibald, 2011).
Clearly, the late Campanian (Judithian) nonavian-dinosaurbear-
ing formations in western North America have less exposure than
late Maastrichtian (Lancian) nonavian dinosaur-bearing forma-
tions in western North America (Fig. 2). Although the late Cam-
panian faunas are known from 16% fewer exposures than those
faunas from the late Maastrichtian faunas, the former have pro-
duced almost double the number of species. Barrett et al. (2009)
echoed such a percentage drop in dinosaur diversity.
The second claim by Fastovsky et al. (2005), that the dino-
saur faunas of the Judithian and Lancian are not preserved with
equal delity, cannot be justied other than possibly by statistical
Figure 2. The mapped area of the late Campanian (ca. 75 Ma) Dino-
saur Park Formation and its equivalents (48,000 km
2
) compared to
the mapped areas of late Maastrichtian (ca. 66 Ma) Lance and Hell
Creek Formations (with 57,000 km
2
; after Archibald, 2011).
machinations. The issue of whether the Dinosaur Park and age-
equivalent formations and the Lance and age-equivalent forma-
tions are equally rich in the number of identiable specimens that
each has yielded cannot be addressed as denitively as the ques-
tion concerning spatial equivalence. There are no complete lists
of the identiable specimens that have been removed over the
past 100 yr and more from these units. Many museums around
the world have nonavian dinosaurs from especially the Lance and
Hell Creek Formations. It is fair to say that both sets of formations
have yielded countless nonavian dinosaur specimens, thus it is up
to naysayers such as Fastovsky and his colleagues to demonstrate
that there is a signicant preservational bias between these two
sets of formations. Barring this, the argument of preservational
bias between the Dinosaur Park and Lance Formations must be
treated as a red herring.
Yet another clear pattern of change in the last 10 m.y. of the
Cretaceous shows that the decline in dinosaur species abundance
is most certainly real. This concerns the patterns of change over
the last 10 m.y. for more common hadrosaurids and ceratopsians
compared to the far rarer theropods. If the Lance nonavian dino-
saur fauna were more poorly sampled than the Dinosaur Park
fauna, it would show rst in a depauperate nonavian theropod
fauna. The rarer theropods should be the rst to disappear from
the record, if it is a matter of sampling. This is decidedly not the
case. For Dinosaur Park, nonavian theropods comprise ~44% of
the species abundance and account for only 10% of the specimens.
By the time of the Lance Formation, whereas nonavian theropods
are equally rare as specimens, if anything, they may have rela-
tively increased taxonomically, ranging from 44% to 52% of the
nonavian dinosaur species abundance. Their numbers of species,
however, declined between 32% and 53% compared to those of
theropods from the Dinosaur Park faunas (Table 1). This is further
strong evidence indicating that comparative sampling is not an
issue. The decline of nonavian theropods in the last 10 m.y. of the
Cretaceous in western North America is most certainly real.
The most-striking pattern of decline concerns the most com-
mon nonavian dinosaurs in the faunas from the Dinosaur Park
and LanceHell Creek Formations, namely, the hadrosaurids and
ceratopsians. Recall that for the Dinosaur Park fauna, among
the ornithischians, hadrosaurs (50%) and ceratopsians (25%)
account for 75% of the specimens. For LanceHell Creek For-
mations, this percentage is, if anything, higher, yet hadrosaurids
declined taxonomically by 80% and the ceratopsians declined
by 40% between these two faunas. Even with this precipitous
taxonomic decline, by end of the Cretaceous in western North
America, hadrosaurids and ceratopsians remain very common
as specimens. Unequivocally, hadrosaurids and ceratopsians
declined in taxonomic diversity in the western interior of North
America during the last 10 m.y. of the Cretaceous. These patterns
of change demonstrate that it was the most common hadrosaurids
and, to a lesser extent, ceratopsians rather than the rarer theropod
taxa that suffered the greatest taxonomic decline during the last
10 m.y. of the Cretaceous in western North America. No statisti-
cal special pleading can explain away these patterns of decline.
on August 25, 2014 specialpapers.gsapubs.org Downloaded from
What the dinosaur record says about extinction scenarios 219
Determining the causes of these taxonomic declines is harder
to do. A possibility noted earlier concerns the fact that the deposi-
tion of the Lance and Hell Creek Formations occurred as part of
the last and most massive regression of the Pierre Seaway leading
to near the Cretaceous-Paleogene boundary. The fossil vertebrate
evidence available for the four transgressions and regressions in
North America during the 35 m.y. of the Late Cretaceous sug-
gests that during times of transgressions, freshwater vertebrates
suffered but land-dwelling species, including nonavian dinosaurs,
fared better, even showing a taxonomic increase. The opposite
occurred during regressions, and, at best, land-dwelling species
show taxonomic stasis, such as for the Dinosaur Park Formation,
or decline during the much greater regression that occurred when
the LanceHell Creek vertebrate faunas lived (Archibald, 1996).
Some parallels can be drawn between the changes that
occurred between the Dinosaur Park and LanceHell Creek
nonavian dinosaur faunas and differences we see in extant mam-
malian biotas. The single LanceHell Creek genus of hadrosau-
rid, Edmontosaurus, is arguably represented by more identi-
able remains than any of the seven hadrosaurids from Dinosaur
Park. Although the number of hadrosaurid species dramatically
declined in the last 10 m.y. of the Cretaceous, the one or two spe-
cies that replaced them were quite common. This is similar to the
pattern of African savannah ungulates compared to ungulates of
pre-European human invasion of North America. In Africa, there
are as many as 60 medium (above 25 kg) to large grazers and
browsers (elephants, rhinoceroses, zebra, pigs, hippopotamuses,
deer, giraffes, antelope, sheep, goats, cattle-like bovids; Halten-
north and Diller, 1992), whereas in North America, there are only
12 such mammals (peccaries, deer, elk, moose, caribou, sheep,
goats, bison, pronghorns; Whitaker, 1980). In Africa, some large
migrating mammals such as zebras and wildebeest have prodi-
gious numbers, but other herbivorous species are not uncommon.
The migratory populations of wildebeest in the Serengeti-Mara
have been measured as high as 1.3 million individuals, and single
concentrations can range from 10,000 to 20,000 animals but are
usually smaller (Kingdon, 1982). Whereas in North America, the
bison was far and away the most numerous large grazing mam-
mal when Europeans arrived some 500 yr ago. In the fteenth
century, 60 million bison roamed North America in massive
migrations. Hornaday (1886) estimated in one area that a migrat-
ing herd reached over four million animals! By 1900, only 1000
animals remained, but this has now grown to over 30,000 indi-
viduals (Whitaker, 1980). North America had a large biomass of
large ungulates, but it was dominated by a single species. It is
possible that a trend toward lower diversity, but still with a large
biomass, transpired for large herbivorous nonavian dinosaurs in
the waning 10 m.y. of the Cretaceous in North America.
This likely scenario does not explain why the diversity of the
carnivorous and scavenging theropods did not decline as dramati-
cally over the same 10 m.y. Although the numbers of potential
prey species clearly declined, as suggested in the comparison of
Lancian hadrosaurs and North American bison, likely the total
numbers of individuals remained similar over the same 10 m.y.
Another possibility is that because the vast majority of these
theropods are ostrich size or smaller, with a diversity of diets, a
decline in the larger hadrosaurid and ceratopsian diversity would
not have affected these smaller theropods as dramatically.
Given that the decline in dinosaur taxonomic diversity
between the late Campanian (Judithian) and the late Maastrich-
tian (Lancian) is real, when did these declines occur and why?
These questions may be addressed as more intermediate-aged
nonavian-dinosaurbearing sites are studied. One such study by
Eberth et al. (2013) examined the dinosaur biostratigraphy of
the Edmonton Group in Alberta, Canada, in considerable detail,
encompassing beds ranging in age from ca. 72.5 to 65.5 Ma, or
the latest Campanian into the early Paleogene. The upper por-
tion, the Scollard Formation, includes the Cretaceous-Paleogene
boundary, with the late Maastrichtian (Lancian) portion including
the same dinosaur species known further to the south in the Lance
and Hell Creek Formations. Below this, there is the Horseshoe
Canyon Formation, which spans the Campanian- Maastrichtian
(Edmontonian) boundary, including the best-known nonavian
dinosaur fauna of this age in the northern part of present-day
North America. The Eberth et al. (2013) study included occur-
rences and biostratigraphic distributions of the nonavian dino-
saurs from the Horseshoe Canyon and Scollard Formations,
which record the regressive phase of the Bearpaw transgressive-
regressive (T-R) cycle and the Fox Hills T-R cycle. Comparing
the areas of the Horseshoe Canyon Formation outcrops shown in
their gure 1 to those of the older and younger beds shown here
in Figure 1, it is clear the Horseshoe Canyon Formation repre-
sents considerably less exposure yet has produced 23 genera of
nonavian dinosaurs. The middle two columns in Table 1 list these
genera. As seen in this table, the taxonomic diversity of nonavian
dinosaurs from the Horseshoe Canyon Formation rivals that of
the younger LanceHell Creek Formations.
Two factors, however, suggest that nonavian dinosaurs
are underrepresented from this intermediate interval. First, as
noted, the Horseshoe Canyon Formation exposures, although
extensive, are far less so than the earlier and later nonavian-
dinosaur producing exposures in northern Laramidia, likely in
part yielding this lower taxonomic diversity. Second, when taxa
are compared across the three time intervals shown in Table 1,
one nds that there may be as many as six genera found both
in the older Dinosaur Park and the younger LanceHell Creek
Formations not known from the intermediate Horseshoe Canyon
Formation. This follows the same pattern found for all of North
America by Archibald and MacLeod (2007). If these Lazarus
taxa are added to those known from the Horseshoe Canyon For-
mation, this interval would have upwards of 29 nonavian dino-
saur genera, surpassing that of the much more greatly exposed
late Maastrichtian LanceHell Creek Formations, but still con-
siderably less than the late Campanian Dinosaur Park Formation.
Although the dip in dinosaurian taxonomic diversity in the
latest Campanian and early Maastrichtian is almost certainly in
part artifactual, caused by the nature of the fossil record, some
of it is certainly real, suggesting that whatever the cause(s) of
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220 Archibald
the decline between the late Campanian and late Maastrich-
tian, much of the decline may have begun or occurred by the
Campanian-Maastrichtian transition. Eberth et al. (2013) noted
that there is no evidence of signicant climatic changes in the
older, but relatively shorter expanse of the Dinosaur Park Forma-
tion, whereas each of the members of the Horseshoe Canyon For-
mation, being as long as the entire duration of the Dinosaur Park
Formation, shows considerable climatic change. Compared to the
Dinosaur Park Formation ornithischians, those in the Horseshoe
Canyon Formation show longer-term morphological stasis and
major climate-changedriven nonavian dinosaur migration into
and out of the area. In the Horseshoe Canyon Formation interval,
the authors detail climatic shifts involving a shift from warm and
wet to cool and seasonally dry and back again, and nally to a
warm and seasonally dry climate in the overlying latest Creta-
ceous Scollard Formation.
Some of these regional climatic changes appear to have been
tectonically driven, but some changes may also have been more
global in scale. What the results of these global changes might
have been is not yet clear, but the longer-term pattern of regres-
sion commencing in earnest ca. 73 Ma, with a much shorter inter-
val of transgression during the latest Campanian into the Maas-
trichtian (Eberth et al., 2013), might have been in part responsible
for these climatic uctuations. This regression also diminished
the expanse of low coastal plains that the nonavian dinosaurs
inhabited, as well as provided new avenues of intercontinental
migrations (Archibald, 1996, 2011). Still later, ca. 67.4 Ma, the
rst but smallest of the three pulses of Deccan volcanism may
have also contributed to climatic uctuations, with the second,
much larger pulse near the Cretaceous-Paleogene boundary con-
tributing even further to climatic change. A smaller third pulse
occurred in early Paleogene (Keller et al., 2012).
APPROACHING THE CRETACEOUS-PALEOGENE
BOUNDARY
Although there is little question that at least in North Amer-
ica, nonavian dinosaur species abundance declined by at least
50% over the last 10 m.y. of the Cretaceous, the record becomes
more problematic as one approaches the Cretaceous-Paleogene
boundary. In order to examine this boundary interval, we must
utilize the latest Cretaceous Hell Creek Formation and the over-
lying early Paleocene Tullock Formation of eastern Montana as
well as parts of western North and South Dakota. This is because
the records of all vertebrates, not just nonavian dinosaurs, have
been more thoroughly studied in these areas, the Cretaceous-
Paleogene boundary is better known, and at least in eastern Mon-
tana, there is a reasonably good vertebrate record in the overlying
Paleocene beds.
Several studies have attempted to determine the diversity
of nonavian dinosaurs in the Hell Creek Formation. One study
(Sheehan et al., 1991) examined what happened to familial-level
diversity of nonavian dinosaurs in a three-part partitioning of the
Hell Creek Formation, arguing that the pattern was commensu-
rate with sudden extinction. Unfortunately, one could not detect
changes below the family level, which was a level too coarse to
detect if there had been any decline within the families (Hurlbert
and Archibald, 1995).
In some recent studies, Pearson et al. (2001; see also Pearson
et al., 2002) identied over 10,000 specimens of very small to
very large vertebrates of 61 vertebrate taxa from a 100 m sec-
tion of Hell Creek Formation exposed mostly in southwestern
North Dakota. One of their major conclusions seems well justi-
ed, namely, that there is no evidence for a decline in overall
vertebrate diversity through the vertical extent of the formation.
Problems arise with their second conclusion; there is no evidence
for a decline of nonavian dinosaur diversity in the 3 m below
the Cretaceous-Paleogene boundary. There is scarce evidence for
any vertebrate fossils in at least the last 1.77 m of their section
(Fig. 3). The single specimen they identied that constitutes this
record is identied as an indeterminate ceratopsid. Of the over
10,000 specimens they collected, there are only three nonavian
dinosaurs identied between 2 and 3 m below the Cretaceous-
Paleogene boundary. These consist of an indeterminate cera-
topsid, an indeterminate hadrosaurine, and the theropod tooth
taxon Richardoestesia. Moving down section, according to their
gures, one encounters ve nonavian dinosaur taxa just below
3 m and eight taxa just below 8 m. Does this mean that, counter
to their arguments, in fact we see a decline from eight to ve to
three to one and nally to zero by 1.77 m below the Cretaceous-
Paleogene boundary. I think not.
The left side of Figure 3 is after their gure 2, whereas the
right side shows their record for nonavian dinosaurs recast as
a spindle or minaret-shaped diagram. In the spindle diagram,
numbers show the nonavian dinosaur taxa either collected at that
level or implied by their lines connecting occurrences below and
above. If this record is to be believed, there is relative taxonomic
stability until about the 4 m mark, when numbers start to drop,
reaching zero before the Cretaceous-Paleogene boundary. This
record would seem to indicate a gradual disappearance of dino-
saurs in these sections. The authors argue that this is artifactual
and that in reality this dinosaur record is commensurate with a
catastrophic extinction. This argument mostly derives from the
so-called Signor-Lipps effect, which argues that because the
fossil record is never complete, we should not expect to have
found the rst or the last fossil records of any particular species
or higher group of organisms. Invoking the Signor-Lipps effect
means, however, that one can claim that catastrophic, gradual, or
some other pattern of extinction is commensurate with the fossil
record. The authors note that possibly the gap near the top of
the Hell Creek Formation is a function of depositional conditions
associated with the transitions between formations rather than
reecting true terminal Cretaceous faunal diversity, an assess-
ment with which I agree.
If the single indeterminate ceratopsid noted above is dis-
counted, the 1.77 m becomes a 2.37 m fossil gap in this data
set. This gap has also sometimes been referred to as the 3 m or
10 ft gap. Some 30 yr ago, this notoriously unfossiliferous gap
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What the dinosaur record says about extinction scenarios 221
was rst thought possibly to represent the level at which dino-
saur extinction occurred. This gap occurs below the Cretaceous-
Paleogene boundary, based on the much more easily and
better-sampled plant pollen record and the iridium-enriched level
thought to mark the Chicxulub impact. Even before the advent
of the impact theory of extinction, it was realized that although
not completely barren, it is a generally unfossiliferous zone for
vertebrate fossils. Thus, for over 30 yr, no one seriously argued
that the base of this gap represented the level of nal extinction
for nonavian dinosaurs.
Unfortunately, this fact was lost on the authors of a paper
in 2011 (Lyson et al., 2011) claiming to show that a chunk of a
ceratopsian brow horn found 13 cm below what they identied
as the Cretaceous-Paleogene boundary proved (p. 14) that a gap
devoid of non-avian dinosaur fossils does not exist and is incon-
sistent with the hypothesis that non-avian dinosaurs were extinct
prior to the K-T boundary impact event. Even taking the conclu-
sions of their eld evidence for this occurrence at face value, it
simply represents one more rare occurrence of a bone, dinosaur
or otherwise, in this quite unfossiliferous gap in fossil representa-
tion near the top of the Hell Creek Formation.
Not surprisingly, this very real gap creates considerable
problems in gauging what was happening not just to dinosaurs,
but all vertebrates during this nearly barren interval. Some, such
as Pearson et al. (2001) and Lyson et al. (2011), have tried to
argue this gap away, basically saying that because no decline
Figure 3. Isolated occurrences of taxa (dots)
are identied within clades connected by
vertical lines within the upper 15 m of the
Hell Creek Formation, exposed mostly in
southwestern North Dakota (after Pearson
et al., 2001). Spindle diagram to the right
shows repeating occurrences of dinosaurs
at the various levels and interpolates the
presence of a taxon if it was recovered
above and below that level (after Archibald,
2011). Note the decline from seven to zero
taxa in the last 4 m and the total lack of any
vertebrates in the uppermost 1.77 m. K
Cretaceous; TTertiary.
D
e
p
t
h

(
m
)
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222 Archibald
in vertebrate diversity can be detected, no such decline, espe-
cially for dinosaurs, occurred in the 3 m below the Cretaceous-
Paleogene boundary. Given that this 3 m (or 2.37 m) gap in the
records compiled by Pearson et al. (2001) is by far the largest
gap in the upper 12 m of the Hell Creek Formation, such a claim
is not warranted. There is no evidence supporting a catastrophic
(bang) or gradual (whimper) extinction in these last 2 to 3 m of
the Hell Creek Formation.
This unfossiliferous gap is vexing. Why is there such a gap
at all? The views vary, and frankly all explanations seem to have
some merit, but choosing one over another is problematic. One
suggestion is that a lack of channel deposits within this 3 m inter-
val may explain the absence of fossil localities. The vast major-
ity of both small and large fossil vertebrates in the Hell Creek
Formation are associated with uvial or riverlain deposits. These
can be the channels themselves, sand bars that breached the chan-
nels during ood events, or more rarely oodplain deposits on
the margins of the streams. Especially compared to the overly-
ing Paleocene Tullock (or Fort Union as it is referred to in the
Dakotas) Formation, the Hell Creek Formation tends to preserve
smaller streams or at least muddier streams in its upper reaches,
although very large channels certainly are not present. The upper
reaches of the Hell Creek Formation for the most part lack larger
channel deposits, although most of the sediment still appears to
be uvially related, and thus the rarity of channel deposits to
explain the absence of fossil localities in the last 3 m seems to
have merit.
Another suggestion is that as a result of the Chicxulub impact
at the Cretaceous-Paleogene boundary, the resulting global acid
rain fallout likely caused the destruction of microfossils imme-
diately underlying the impact layer (Bailey et al., 2005). At rst
an appealing idea, the survival of most aquatic vertebrate species
belies the idea of a very low pH acid rain. The possibility of the
dissolution of most teeth and bones in the upper 3 m of the Hell
Creek Formation may yet prove to be true, but not from acid rain.
Rather, the carbonaceous and lignitic coals that most often over-
lie the top of the Hell Creek Formation could have caused the
leaching near the top of the Hell Creek Formation. This admit-
tedly more prosaic explanation seems more likely to have been
the cause.
Finding and thoroughly exploring other terrestrial
Cretaceous-Paleogene sections will help to determine if this gap
is a regionally driven phenomenon or is more global in extent.
Finding a single bone in a single section, such as done by Lyson
et al. (2011), does not begin to address the issue or answer any
question about the rapidity of dinosaur extinction. Similarly,
although not concerned with the subject of this paper (i.e., non-
avian dinosaur extinction), two recent papers dealing with extinc-
tions of other vertebrate groups misrepresent the nature of the
vertebrate fossil record near and at the Cretaceous-Paleogene
boundary. One study shows the fossil record of lizards and snakes
extending exactly to the Cretaceous-Paleogene boundary (Lon-
grich et al., 2012). The second, and more egregious in its misrep-
resentation of the fossil record, shows the record for birds as sin-
gle dots immediately below the Cretaceous-Paleogene boundary
(Longrich et al., 2011). Although some species of snakes, lizards,
and birds may have become extinct at the Cretaceous-Paleogene
boundary, to my knowledge, there is no record of any of these
taxa within 2 to 3 m of the Cretaceous-Paleogene boundary. The
problem with all of these studies is the impression of a sudden
extinction, when in fact such evidence is completely lacking.
Another approach to examining what was occurring leading
up to and across the Cretaceous-Paleogene boundary has used
large samples of vertebrate microfossils amassed over almost
40 yr, mostly by crews often under the direction of W.A. Cle-
mens from the Museum of Paleontology at the University of
California, Berkeley, working in the late Maastrichtian (Lancian)
Hell Creek Formation and early Paleocene Tullock Member of
northeastern Montana. One more recent approach by Wilson
and colleagues (Hutchison et al., 2004; Wilson, 2005) placed
vertebrate localities into a temporal framework utilizing strati-
graphic position. Although their work did not address dinosaur
records, it is relevant to understanding possible broader climatic
changes that certainly would have affected dinosaurs. These
authors noted changes in the better-sampled amphibian, turtle,
and mammalian paleocommunities approaching the Cretaceous-
Paleogene boundary. They compared these data with those for
published paleooral and paleoclimatic data. Echoing the nd-
ings of Pearson and colleagues, Wilson and colleagues did not
nd any evidence of long-term changes within paleocommuni-
ties leading up to the Cretaceous-Paleogene boundary, although
they did nd some changes in relative abundances, taxonomic
composition, and body sizes that likely reect normal responses
to background levels of climate change. However, unlike Pearson
and colleagues, Wilson and colleagues did nd dramatic changes
in turtle and mammalian paleocommunities within the last
100 k.y. of the Cretaceous. These authors went on to write that
while their results are consistent with a sudden extinction, such
as from an extraterrestrial impact, the response was nonlinear,
so that long-term causes or multiple short-term causes cannot be
rejected. They further wrote that their assessments will improve
with increased temporal resolution of the chronostratigraphic
framework, improvement of the density of fossil sampling, the
incorporation of other fossil taxa such as plants and mollusks,
and comparisons with other study areas.
This work shows that during the last 2 m.y. of the Creta-
ceous, maximum diversities of turtles, mammals, and plants cor-
relate with the maximum latest Cretaceous warming trend, while
the drop in taxonomic richness of these three groups correlates
to a rapid drop in paleotemperature in the last 100,000 yr of the
Cretaceous of the same region (Hutchison et al., 2004; Wilson,
2005). If veried by further studies, these results clearly point
to factors driving climatic change well before an extraterrestrial
impact that would be expected to have affected dinosaur relative
abundances and/or taxonomic diversity in some fashion.
Finally, a more recent approach by Horner et al. (2011)
examined in considerable detail the stratigraphic distribution of
nonavian dinosaur skeletons and individual bones in the Hell
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What the dinosaur record says about extinction scenarios 223
Creek Formation of northeastern Montana. Whereas up to six
genera are known in the lower two thirds based on skeletons
and individual bones (Triceratops, Tyrannosaurus, Edmonto-
saurus, Thescelosaurus, Ornithomimus, Pachycephalosaurus,
and Ankylosaurus), only three genera (Triceratops, Tyranno-
saurus, Edmontosaurus) are known in the upper one third, and
this portion is dominated by a single taxonTriceratops. The
evidence then, if it shows anything, is that there was a decline in
relative abundance, if not dinosaur diversity, in the upper Hell
Creek Formation.
CONCLUSIONS
The lack of well-studied, fossiliferous Cretaceous- Paleogene
terrestrial sections on all continents except western North Ameri-
can means that the records from these North American locations
must for now act as proxy for our knowledge of dinosaur extinc-
tion. During the last 10 m.y. of the Cretaceous, in at least the
northern part of the western interior of North America, nonavian
dinosaur species dropped from 49 to 25, almost a 50% decline.
This is the case even though the latest Cretaceous vertebrate-
bearing beds are represented by a 16% greater areal extent than
beds some 10 m.y. earlier in the same region. This decrease in
nonavian dinosaur taxonomic diversity commenced near the
Campanian-Maastrichtian boundary and likely continued during
the greatest marine regression to occur in the past 250 m.y. and
later during phases of Deccan Trap emplacement. Resulting cli-
matic changes and loss of habitat likely contributed to this decline.
The nonavian dinosaur record near the Cretaceous-Paleogene
boundary, even for the best-known records from North America,
remains enigmatic and open to interpretation. Newer studies sug-
gest some decline in at least relative abundance of nonavian dino-
saurs as one approaches the Cretaceous-Paleogene boundary, but
the cause (or causes) for their nal extinction (if it was the case)
remains unresolved, although the Chicxulub impact undoubtedly
played a major role following upon the effects of longer-term
regression and volcanism.
ACKNOWLEDGMENTS
The organizers of the International Conference on Volcanism,
Impacts and Mass Extinctions: Causes and Effects Norman
MacLeod, Gerta Keller, and Andrew Kerrare thanked for
their kind invitation to participate and for arranging partial
nancial support. Alexander Averianov, Hans Dieter-Sues, and
Gerta Keller are thanked for reading and providing helpful
comments on the manuscript.
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