Palaeogeography, Palaeoclimatology, Palaeoecology 214 (2004) 195 – 223
www.elsevier.com/locate/palaeo
Extinction and recovery patterns of scleractinian corals at the
Cretaceous-Tertiary boundary
Wolfgang Kiesslinga,*, Rosemarie C. Baron-Szabob
a
Institute of Paleontology, Museum of Natural History, Humboldt-University Berlin, Invalidenstr. 43, 10115 Berlin, Germany
Smithsonian Institution, Department of Zoology, National Museum of Natural History, W-329, MRC-163, Washington, DC 20560, USA
b
Received 1 July 2003; received in revised form 20 April 2004; accepted 20 May 2004
Abstract
The extinction and recovery of scleractinian corals at the Cretaceous–Tertiary (K–T) boundary was analyzed based on a
global database of taxonomically revised late Campanian to Paleocene coral collections. In contrast to earlier statements, our
results indicate that extinction rates of corals were only moderate in comparison to other marine invertebrates. We have
calculated a 30% extinction rate for Maastrichtian coral genera occurring in more than one stratigraphic stage and more than one
geographic region. Reverse rarefaction suggests that some 45% of all coral species became extinct. Photosymbiotic
(zooxanthellate) corals were significantly more affected by the extinction than azooxanthellate corals; colonial forms were hit
harder than solitary forms, and among colonial forms an elevated integration of corallites raised extinction risk. Abundance, as
measured by the number of taxonomic occurrences, had apparently no influence on survivorship, but a wide geographic
distribution significantly reduced extinction risk. As in bivalves and echinoids neither species richness within genera nor larval
type had an effect on survivorship. An indistinct latitudinal gradient is visible in the extinction, but this is exclusively due to a
higher proportion of extinction-resistant azooxanthellate corals in higher-latitude assemblages. No significant geographic
hotspot could be recognized, neither in overall extinction rates nor in the extinction of endemic clades.
More clades than previously recognized passed through the K–T boundary only to become extinct within the Danian. These
failed survivors were apparently limited to regions outside the Americas. Recovery as defined by the proportional increase of
newly evolved genera, was more rapid for zooxanthellate corals than previously assumed and less uniform geographically than
the extinction. Although newly evolved Danian azooxanthellate genera were significantly more common than new
zooxanthellate genera, the difference nearly disappeared by the late Paleocene suggesting a more rapid recovery of
zooxanthellate corals in comparison to previous analyses. New Paleocene genera were apparently concentrated in low latitudes,
suggesting that the tropics formed a source of evolutionary novelty in the recovery phase.
D 2004 Elsevier B.V. All rights reserved.
Keywords: Cretaceous–Tertiary boundary; Mass extinction; Corals; Evolution; Reefs
* Corresponding author.
E-mail address: [email protected] (W. Kiessling).
0031-0182/$ - see front matter D 2004 Elsevier B.V. All rights reserved.
doi:10.1016/j.palaeo.2004.05.025
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W. Kiessling, R.C. Baron-Szabo / Palaeogeography, Palaeoclimatology, Palaeoecology 214 (2004) 195–223
1. Introduction
The mass extinction at the Cretaceous–Tertiary
boundary (K–T, officially Cretaceous–Paleogene)
still provokes many questions considering the reasons, magnitude, speed, selectivity and geographic
patterns of extinction and subsequent recovery. The
Chicxulub impact is currently considered the most
likely culprit for the mass extinction, but apart from
other potential causes, such as Deccan Trap volcanism (McLean, 1985; Courtillot, 1999), the impactkill effect remains ambiguous in many instances.
Extinctions in some clades were severe, while others
show almost no taxonomic change at the K–T
boundary. The most dramatic examples were
reported from the marine plankton, where calcareous
nannoplankton and planktonic foraminifers faced
extremely high extinction (Gartner, 1996; Arenillas
et al., 2002), while radiolarians and dinoflagellates
only show some ecological response but almost no
extinction (Hollis, 1997; Brinkhuis et al., 1998).
Among invertebrate macrofossils, detailed studies
have revealed a similar though less extreme disparity: bivalves including the rudists exhibit a 63%
generic extinction (Raup and Jablonski, 1993),
whereas the extinctions in sea urchins was shown
to have been not more than 36% (Smith and Jeffery,
1998). Clearly, additional datasets have to be
analyzed at global scales to extract general patterns
of the end-Cretaceous mass extinction and to get a
better idea of the underlying cause(s).
This study documents global geographic patterns
of extinction and recovery in scleractinian corals in
the Maastrichtian and Paleocene. We emphasize five
key points:
1.
2.
3.
4.
5.
the intensity of the end-Cretaceous coral extinctions and the pace of recovery;
the ecological selectivity of coral extinctions;
the impact of coral extinctions on the global reef
ecosystem;
the geographic patterns of coral extinction and
recovery;
a comparison of the results with other benthic
invertebrate groups.
Although we do not focus on potential causes of the
coral extinctions, we also discuss if our results are
compatible with the impact scenario at the K–T
boundary.
2. Database and methods
2.1. Database
Data in this paper are part of a comprehensive
geographic database on the K–T boundary (KTbase),
which currently comprises geological, mineralogical
and paleontological information from 490 K–T
boundary sites (Kiessling and Claeys, 2001; Claeys
et al., 2002). A K–T boundary site is defined as an
area (100 km2 or less) where at least Maastrichtian
and/or Danian sediments are preserved. Closely
spaced sections in the same depositional environment
are combined for the summary data (e.g., sedimentology and geochemistry) but paleontological data are
partitioned as finely as possible from published
sources. For the particular purpose of this study, the
concept for K–T boundary sites has been widened to
include also sites with only late Campanian or late
Paleocene sediments preserved. This was done to
compare the extinction and recovery at different
temporal scales.
Data in KTbase were largely extracted from the
published literature but personal observations and
unpublished data are also included. Paleontological
data are stored in faunal lists representing individual
fossil collections. Each taxon reported in a collection
constitutes a taxonomic occurrence in the database.
Abundance data are available for less than one third of
the taxonomic occurrences. We thus used the number
of occurrences as an approximation of recorded
abundances and applied this criterion for resampling.
All taxonomic data and some 80% of the occurrence data have been taxonomically revised at the
species-level (Baron-Szabo, 2002; B.-S. work in
progress) and stratigraphic assignments were updated
for the purpose of this study. The occurrence table
with updated taxonomic information is available as an
electronic appendix (electronic appendix see Background Data Set). Only taxonomic data resolved at
least to the genus level are represented in KTbase.
Species-level taxonomic information is stored in the
database whenever possible. Coral data from 228 late
Campanian to Paleocene collections (Fig. 1) comprise
W. Kiessling, R.C. Baron-Szabo / Palaeogeography, Palaeoclimatology, Palaeoecology 214 (2004) 195–223
197
Fig. 1. Global distribution of coral-bearing K–T-boundary sites (with at least genus-level taxonomic information) (triangles) and number of
occurrences per 308 grids (pies—only shown for grids with more then five occurrences) plotted on a 65 million years paleogeographic
reconstruction (modified from Golonka, 2002). Size of pies is proportional to the number of occurrences in each grid. Brick pattern:z-like (=taxa
that are likely to have hosted zooxanthellae) coral occurrences; black fill:az-like (=taxa that are likely to not have hosted zooxanthellae) coral
occurrences.
1235 taxonomic occurrences, 187 genera and 460
species. The low ratio of occurrences to species
richness indicates that (1) corals in this time interval
are underexplored and the true biodiversity is far from
being completely known (e.g., very high), (2) coral
occurrences are usually reported on rather large spatial
and temporal scales, and/or (3) in spite of our
taxonomic revisions the corals may still suffer oversplitting. Our revisions have reduced the taxonomic
noise considerably by removing many subjective
synonyms (632 nominal species were parsed into
460 valid species). The main reason for the low
occurrence/diversity ratio is probably the way coral
data are usually reported in the literature. Most coral
collections stem from whole outcrops or local areas
and usually combine several beds or even lithostratigraphic members. This limits the applicability of
resampling techniques such as rarefaction.
Sepkoski’s compendium of the stratigraphic ranges
of marine animal genera (Sepkoski, 2002) was used to
determine long-term diversity dynamics of scleractinian corals and to compare two independent global
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datasets at the K–T boundary. Sepkoski’s data were
largely derived from the secondary literature and give
a comprehensive but probably biased view of evolutionary dynamics across the K–T boundary, due to the
lack of taxonomic expertise in the compendium.
Previous comparisons have demonstrated that
although many data in the compendium are inaccurate, large-scale diversity patterns are little affected
because the errors are randomly distributed (Adrain
and Westrop, 2000). Our comparison intended to
provide a further test of the effect of errors in the
compendium.
2.2. Taxonomic framework
The large morphological variation of colonial
corals traditionally has caused problems in their
identification. Not only the overall shape of corals
can vary substantially but also the corallite structure
can change within a colony (Veron, 1995). As
exemplified and discussed by Baron-Szabo (2002),
our supraspecific taxonomy is mostly based on
macrostructural characters, which for modern corals
have been shown to better accord with molecular data
than the often used micro-skeletal characters (Veron et
al., 1996). We are aware of pitfalls in this approach
but currently see this as the best way to integrate the
taxonomy of ancient corals where molecular data are
not available and population dynamics cannot be
assessed in detail. Even the molecular approach
currently results in contrasting hypotheses on coral
evolution (Romano and Cairns, 2000). Phylogenetic
systematics based on skeletal characters may solve
issues in some taxa (Cairns, 1997, 2001) but are
difficult to apply in colonial forms. Veron (1995)
suggested that hybridization and homoplasy is so
common in corals that their dreticulate evolutionT will
always lead to equivocal cladograms. Moreover, the
scleractinian skeleton itself is sometimes seen as just a
grade of organization without phylogenetic significance (Stanley, 2003).
Judging from previous comparisons between phylogenetic and typological approaches to analyzing
long-term faunal change in corals (Johnson, 1998), we
are confident that results from our typological
approach will not significantly differ from more
sophisticated phylogenetic analyses (but see Smith
and Patterson, 1988 for a different view). We consider
a homogeneous taxonomic concept and specimenbased revisions more important than phylogenetic
analyses when diversity dynamics are analyzed at the
species and genus level.
2.3. Stratigraphic framework
A thorough revision of all latest Cretaceous and
Paleocene coral collections was required due to the
often obsolete biostratigraphic data in older monographs, revised chronostratigraphic assignments of
Late Cretaceous foraminiferal zones (Robaszynski
and Caron, 1995; Arz and Molina, 2002), the now
formal definition of the Campanian–Maastrichtian
boundary (Odin, 2001; Odin and Lamaurelle, 2001)
and new Strontium isotope data from key localities
(Swinburne et al., 1992; Steuber et al., 2002). Several
coral collections formerly thought to be of Maastrichtian age (Alloiteau, 1952b; de la Revilla and
Quintero, 1966; Tchechmedjieva, 1986) have been
shown to fall into the Campanian, whereas others,
formerly imprecisely dated as Campanian or early
Maastrichtian, are now dated as late Maastrichtian
(Mitchell, 2002; Steuber et al., 2002). A reliable
revision of stratigraphic assignments was possible for
about 80% of our coral collections.
Because coral-bearing deposits are rarely dated
precisely, and because the global approach has to
include some problematic stratigraphic assignments,
we limit our discussion to standard stages and
substages. More specific stratigraphic assignments
are occasionally available but have been subsumed
into those larger intervals. Our database separates late
Campanian (uc), unresolved late Campanian–early
Maastrichtian (cm), early Maastrichtian (lm), unresolved Maastrichtian (m), bmiddleQ to late Maastrichtian (mum), late Maastrichtian (um), early to middle
Danian (ld), late Danian (ud=Montian), unresolved
Danian (d), Selandian (mp), unresolved Paleocene (p)
and Thanetian (up) ages. The numeric distribution of
occurrences and reported species richness in seven
stratigraphic intervals is summarized in Fig. 2. For the
analyses of extinction and recovery the data are usually
partitioned into four intervals: the first combines all
data from the late Campanian to the late Maastrichtian
(CM), the second is limited to occurrences of
confirmed Maastrichtian age (M), the third comprises
all confirmed Danian occurrences (D including the
W. Kiessling, R.C. Baron-Szabo / Palaeogeography, Palaeoclimatology, Palaeoecology 214 (2004) 195–223
199
Fig. 2. Numeric distribution of coral occurrences and recorded species richness in seven stratigraphical intervals in KTbase.
Montian) and the fourth sums data from the Paleocene
(Pa). A comparison between CM and Pa has the
advantage that sample sizes are maximized and the
time intervals are of similar duration (10 My), while
the comparison between M and D data provides a finer
stratigraphic resolution.
2.4. Ecological categories
Corals are traditionally categorized as either
hermatypic (reef building) or ahermatypic (non reef
building), which is often treated as being equivalent to
symbiont bearing (zooxanthellate) versus non-symbiont bearing (azooxanthellate). This view has been
critically assessed by Rosen and Turnšek (1989) and
Rosen (2000), who suggested the terms z-like for
forms resembling modern zooxanthellate corals and
az-like for forms that are more similar to modern
azooxanthellate corals. Although there is always some
uncertainty with the ecological categorization of
extinct taxa, the lists of criteria compiled in Wilson
and Rosen (1998) and Rosen (2000) define a
homogenous protocol, thereby minimizing subjective
errors. We categorized ecology at the species level,
but for our analyses, categories have been assigned to
genera. All genera containing one or more az-species
were assigned to the az-like category for this purpose.
Genera such as Oculina and Cladocora are thus
generally treated as az-corals even though some extant
species in those clades may host photosymbionts.
Haimesastraea, although showing morphological
characteristics of z-like corals, has also been grouped
with az-corals because of its commonness in high
latitudes and siliciclastic environments (personal
observations). It could be argued that genera of mixed
trophic mode could as well be assigned to both the azlike and z-like categories. However, the philosophy
behind our approach is to separate taxa with optional
photosymbiosis from those which depend on photosymbiosis for survival.
In comparison to bivalves, gastropods and echinoids, the categorization of other ecological traits is
limited in fossil corals. Colony size has some relation
to physico-chemical variables but the response varies
significantly between clades (Veron, 1995) and thus
has no predictable relationship with ecological factors.
Similar problems apply to coral shape. In addition to
photosymbiosis, we have categorized colonial versus
solitary growth, maximum adult size and colony
integration. Colony integration was measured in four
ordinal intervals: low (dendroid, phaceloid), medium
(cerioid, plocoid), high (thamnasterioid) and very high
(meandroid).
2.5. Assessment of data
All too often, paleontological data are presented as
such without any indication of statistical errors.
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analyze extinction and recovery of corals based on
multiple sources and discuss ecological selectivity and
geographic patterns. Their basic results were that (1)
total generic extinctions were around 60% (see also
Rosen in MacLeod et al., 1997); (2) z-like corals were
more strongly affected by the end-Cretaceous extinctions than az-like corals; (3) az-like corals became
relatively more common in the Paleocene; (4) biogeographic differentiation decreased in the Paleocene;
and (5) Paleocene coral reef development was nearly
independent of the previous extinction and cannot be
viewed as a recovery phenomenon.
While these analyses provided an in-depth review,
they have some shortcomings. Firstly, and most
importantly, the taxonomic data were extracted from
the published literature without revisions. Secondly,
the analyses of extinction patterns were carried out on
rather large temporal scales (three intervals in the
Cretaceous, one in the Paleocene) and some stratigraphic assignments were incorrect. Four out of
twelve of the bMaastrichtianQ localities indicated by
Rosen and Turnšek (1989; Table 1) are now dated as
late Campanian or older. Thirdly, the data were
analyzed from selected regions, rather than globally.
Other original papers exploring corals at the K–T
boundary were devoted to paleogeographic distributions of selected taxa (Beauvais and Beauvais, 1974)
and the fate of coral families (Barta-Calmus, 1984).
All studies agree that the end-Cretaceous mass
Because coral data are limited in comparison to other
macroinvertebrates such as bivalves and gastropods,
an indication of statistical confidence limits is
imperative to judge the significance of reported
percentage data. We followed Raup (1991) to
indicate binomial errors on percentage values. These
binomial errors reflect uncertainties of the percentage
values depending on sample size (N) and the
percentage values themselves. We report binomial
errors as 95% confidence intervals in the text. In
comparison to standard errors, this has the advantage
that significant differences can immediately be
recognized. The limited number of taxa implies fairly
large confidence intervals implying considerable
uncertainties on reported rates and differences.
Statistical relationships between diversity dynamics
and other attributes were generally tested with
nonparametric statistics. Nonparametric statistics are
preferred because they do not assume that data are
normally distributed. Test results are reported in the
text by the probability of randomness ( P).
3. Corals at the K–T boundary—state of the art
Only a handful of papers have systematically
explored the fate of scleractinian corals around the
K–T boundary. Most noteworthy are the papers of
Rosen and Turnšek (1989) and Rosen (2000) which
Table 1
Extinction metrics of corals at the KT boundary
Late Campanian–Maastrichtian data
KTbase
Error
N
Sepkoskia
Error
N
Total (with singletons)
z-like (with singletons)
az-like (with singletons)
Total (without singletons)
z-like (without singletons)
az-like (without singletons)
39.0
45.0
27.8
34.3
43.9
13.3
7.7
9.8
11.9
7.8
9.8
9.9
154
100
54
143
98
45
43.2
49.2
36.8
38.0
47.5
26.5
8.9
12.5
12.5
9.2
12.7
16.5
118
61
57
108
59
49
Maastrichtian data only
KTbase
Error
N
Total (with singletons)
z-like (with singletons)
az-like (with singletons)
Total (without singletons)
z-like (without singletons)
az-like (without singletons)
34.3
39.6
25.0
28.8
39.6
9.3
7.8
10.0
11.8
7.7
10.0
8.7
143
91
52
132
89
43
All values except N (number of genera) in percent. Errors represent 95% confidence intervals.
a
Analyses based on data in Sepkoski (2002). Results excluding cm and uc data are the same and therefore not shown.
W. Kiessling, R.C. Baron-Szabo / Palaeogeography, Palaeoclimatology, Palaeoecology 214 (2004) 195–223
extinction affected scleractinian corals severely, but at
the same time call for more detailed studies.
4. Long-term ecological and evolutionary pattern
of scleractinian corals
4.1. Late Cretaceous–Paleogene reefs
Looking at the fossil record of reefs, corals appear
to have benefited from the end-Cretaceous mass
extinction. Late Cretaceous shallow tropical shelves
were often dominated by rudists. These unusual
bivalves were abundant nearly everywhere on carbonate platforms, sometimes even forming reefal structures (Johnson et al., 2002; but see Gili et al., 1995,
for a different view). Corals instead, while also
common, rarely achieved rock-forming abundance in
the Late Cretaceous, but in the Paleocene, corals
dominate the global reef factory by far (Kiessling et
al., 1999). There is little evidence that rudists directly
outcompeted corals (Gili et al., 1995; Skelton et al.,
1997; Götz, 2003). The relative scarcity of latest
Cretaceous coral reefs has been attributed to the
absence of encrusters (Moussavian, 1992) and oceanographic changes (Scott, 1995). Although coral
reefs remain rare for most of the Paleogene in terms of
absolute recorded numbers (Kiessling, 2002), their
globally preserved volume peaked in the Paleocene,
dropped dramatically in the earliest Eocene and then
201
climbed sharply in the late Oligocene and Neogene
(Fig. 3). While these observations concur with Rosen
(2000) that the Late Cretaceous was a time of very
limited coral reef development, the relatively massive
coral reef production in the Paleocene contrasts earlier
statements saying that the Paleocene was a time of
depressed reef building (Bryan, 1991; Vecsei and
Moussavian, 1997).
The sedimentary environments of our coral collections parallel the trend in reefs. Some 23% of all late
Campanian to Paleocene coral occurrences in KTbase
are reported from reefs (see Flügel and Kiessling,
2002a for definition). Only 16F3% of the late
Campanian and Maastrichtian (CM) coral occurrences
are from reefs, but significantly more Paleocene coral
occurrences (36F5%) are known from reefal settings.
These data contrast with the partitioning of az-like and
z-like corals in both intervals. While z-like corals
constitute 59F4% in assemblages of CM, only
37F5% of all reported Paleocene occurrences are zlike (Fig. 1). These global data agree with the
geographically more restricted analysis of Rosen and
Turnšek (1989). The discrepancy between z-like coral
and reef distributions has two reasons. Firstly, several
az-like corals are involved in Paleocene reef building
(e.g., Bernecker and Weidlich, 1990) but played no
such role in the latest Cretaceous, and secondly, the
fewer Paleocene z-like taxa were relatively more
common in reefal associations than the late Campanian–Maastrichtian ones. The latter observation is
Fig. 3. Global preserved reef volume constructed by corals in the Mesozoic and Cenozoic plotted at a stage-level stratigraphic resolution.
Calculated with the PaleoReef database following Kiessling et al. (2000). Note logarithmic scale. Tr—Triassic, J—Jurassic, K—Cretaceous,
Pg—Paleogene, Ng—Neogene.
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probably related to the evolutionary radiation of
encrusting algae (Steneck, 1983; Aguirre et al.,
2000) rather than to greater reef-building potential
of the corals themselves. Crustose coralline algae are
very important for reef cementation and ease the larval
settlement of benthic organisms (Fabricius and
De’ath, 2001). Nearly all well-known Paleocene reef
occurrences are indeed characterized by rich flora of
coralline algae (Babic et al., 1976; Babic and Zupanic,
1981; Bryan, 1991; Schuster, 1996; Tragelehn, 1996;
Montenat et al., 2002).
Among the Big Five Phanerozoic mass extinctions,
the K–T boundary is unusual in that it represents the
only event without a significant drop in reefal
carbonate production at the stage level (Flügel and
Kiessling, 2002b). This could either mean that coral
reefs were little affected by the extinctions, or coral
reefs recovered more rapidly than at other extinction
intervals. Although data are too limited to provide a
final judgment, the observation that the first known
Paleocene shallow-water coral reefs are of mid Danian
(P1b) age (Tragelehn, 1996; Montenat et al., 2002;
Baron-Szabo et al., 2004) points to an actual gap in
their record but at the same time suggests a rapid
recovery. Kiessling and Claeys (2001) observed a
peculiar geographic pattern in reef recovery. While
Fig. 4. Diversity dynamics of Mesozoic–Cenozoic well-known scleractinian coral genera plotted at a stage-level stratigraphic resolution. (A)
Standing diversity. (B) Per genus extinction and origination rates. Genera occurring in only one stage (singletons) were excluded from this
analysis as were genera whose first and last occurrence is not resolved to the stage-level. Note that this method tends to underestimate standing
diversity while extinction and origination rates are exaggerated. Based on data from Sepkoski (2002). Rhae=Rhaetian; Plie=Pliensbachian;
Kimm=Kimmeridgian; Ceno=Cenomanian; Maa=Maastrichtian.
W. Kiessling, R.C. Baron-Szabo / Palaeogeography, Palaeoclimatology, Palaeoecology 214 (2004) 195–223
coral-algal reefs are known from the Danian of
Europe, North Africa, and South America, the oldest
Paleocene coral reefs in the Gulf Coast area are of
Thanetian (late Paleocene) age. Their hypothesis that
enhanced ecosystem devastation in the Gulf of
Mexico region due to the proximity of the Chicxulub
impact crater was responsible for this pattern is
intriguing but remains untested.
4.2. Diversity dynamics of Mesozoic–Cenozoic
scleractinians
Long-term diversity dynamics of scleractinian
corals were analyzed using the compilation of
Sepkoski (2002) at a stage-level stratigraphic resolution. The analysis was limited to taxa known from
at least two separate geological stages and taxa
whose first and last occurrences can be detailed to
the stage-level. In addition to taxonomic problems,
which will be discussed below, problems with
stratigraphic ranges are significant, not only for
Sepkoski’s compendium but also for stratigraphic
ranges of corals in general (Baron-Szabo, 2002).
Imprecise age assignments of coral occurrences are
inherent to their habitat preferences (shallow water
with few reliable stratigraphic markers) and thereby
obscure the inferred evolutionary dynamics. Of the
871 scleractinian genera listed in Sepkoski’s compilation, 405 have imprecise assignments of first or last
occurrences. We have excluded all those genera from
the analysis being aware that this method tends to
underestimate standing diversity (the number of
genera inferred to have co-existed in a time interval)
and exaggerates extinction and origination rates.
Similarly, genera that are only reported from one
stage (singletons) were also excluded from the
analysis, which tends to reduce both standing
diversity and turnover rates. Although singletons
may yield a biological signal, it is more likely that
they are mostly due to limited research or preservational problems (Foote, 1997, 2000). The resulting
graph is based on only 335 coral genera and one
should note that confidence intervals, although not
indicated, are quite large. Values were normalized by
the duration of the stage (time scale of Golonka and
Kiessling, 2002), which makes the implicit assumption that taxonomic turnover was continuous
throughout the stratigraphic interval, an assumption
203
that probably does not hold true for mass extinction
intervals as will be discussed below.
Standing diversity (Fig. 4A) of corals exhibits a
discontinuous rise towards the Recent, a trend usually
observed in large compilations of taxon-ranges
(Benton, 1995; Jablonski et al., 2003). Interesting
features of the coral diversity curve lie in the Late
Jurassic (Oxfordian–Kimmeridgian) diversity peak,
the Late Cretaceous (Turonian–Maastrichtian) diversity plateau, and in the slow rise of generic richness
during most of the Paleogene (Danian–Rupelian). The
latter two observations are relevant to this paper and
indicate (1) that the scarcity of Late Cretaceous Coral
Reefs was not mirrored by a depression of coral
diversity and (2) the end-Cretaceous rudist extinction
was not directly related to coral diversity dynamics.
5. Overview of Coral-Bearing K–T regions
5.1. Antarctica, New Zealand and South America
Maastrichtian and Paleocene coral faunas from the
James Ross Basin of Antarctica are well known
(Filkorn, 1994; Stolarski, 1996). The detailed work of
Filkorn (1994) has revealed a surprising biodiversity
in the Maastrichtian as well as the Paleocene. The
faunas are predominantly solitary and exclusively azlike. One possible exception is Cladocora gracilis
(d’Orbigny) (=Cladocora antarctica Filkorn).
Low diversity coral faunas have also been recorded
from New Zealand (Squires, 1958; Stilwell, 1997). K–
T corals are poorly explored in South America,
although they are relatively common in the Roca
Formation of Argentina. Our preliminary observations
indicate moderate diversities with about equal proportions of az- and z-like corals in northern Patagonia,
with a trend towards az-dominance towards the south.
Corals were apparently more common in the Danian
than in the Maastrichtian. The recent discovery of a
Danian coral reef in La Pampa, Argentina (BaronSzabo et al., 2004) and ongoing field work in the area
will provide additional data in the near future.
5.2. Caribbean and North America
Rich coral-rudist associations of Jamaica have long
attracted paleontologists focusing on the K–T extinc-
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tions (Coates, 1977). Maastrichtian corals occur in a
mixed volcaniclastic-carbonate platform (Mitchell,
2002). The richest coral collections stem from the
Titanosarcolites limestone, which is now dated as late
Maastrichtian in most parts (Steuber et al., 2002). The
taxonomic work by one of the authors (B.-S., work in
progress) on the large collections of Coates, Kauffman
and Jackson (1966–1972) has unraveled what is by far
the greatest Maastrichtian coral diversity in the
western Hemisphere. A recent review of Mitchell
(2002) has provided details on vertical patterns of
coral occurrences. Other K–T corals have been
described from the Danian of Puerto Rico (Berryhill
et al., 1960) and the Late Cretaceous of Cuba (Wells,
1941).
Maastrichtian and Paleocene coral faunas are well
known from the Gulf Coast region (USA) but contain
almost exclusively az-like corals. Older studies
describe a fairly high diversity (Stephenson, 1917;
Vaughan, 1920; Wells, 1933), but according to our
revisions the corals were excessively oversplit.
Maastrichtian assemblages from Mexico contain more
z-like corals with a moderate to high species-richness
(Myers, 1968; Filkorn, 2003; Schafhauser et al.,
2003). Paleocene corals are obviously rare in the
region (Vaughan, 1900) and the oldest corals reported
from reefal assemblages are of Thanetian age (Bryan,
1991; Stemann in Bryan et al., 1997).
5.3. Africa
The best-explored K–T coral faunas are known
from Egypt (Quaas, 1902; Wanner, 1902; Hassan and
Salama, 1969; Schuster, 1996). Maastrichtian corals
were apparently rare, and exclusively az-like corals
have been described, which all passed the K–T
boundary (Quaas, 1902; Tantawy et al., 2001). Rich
Paleocene faunas consist of about equal proportions of
az-like and z-like corals, the latter forming coral reefs
(Schuster, 1996). In Somalia, Maastrichtian and
Paleocene coral-bearing sediments are known, but
only the late Paleocene–early Eocene Auradu Limestone is well described taxonomically (Gregory, 1900;
Carbone et al., 1993). K–T corals from West Africa
are only known from older publications (Alloiteau,
1952a; Barta-Calmus, 1969). Taxonomic revisions
were possible but the stratigraphic assignments could
not be updated. Both Maastrichtian and Paleocene
associations are dominated by az-like corals. Rich
Campanian–Maastrichtian faunas are known from
Madagascar (Alloiteau, 1958) and consist of about
equal proportions of az-like and z-like corals. A
wealth of new data has allowed for a revision of
Alloiteau’s stratigraphic assignments (Bignot et al.,
1998; Rogers et al., 2000; Abramovich et al., 2003).
5.4. Alpine belt and southern Europe
Maastrichtian carbonate platforms, rich in corals
and rudists, cover much of the southern European
shallow shelves but taxonomic data on well-dated
localities are scarce (Polšak, 1985; Parente, 1994).
Although apparently rarer, data on Paleocene coral
localities are much better and have revealed a
surprising diversity (Drobne et al., 1988; Moussavian
and Vecsei, 1995; Vecsei and Moussavian, 1997;
Turnšek and Drobne, 1998).
In the Alps, too, the Paleocene record of corals is
much richer than the Maastrichtian record. Reefal
carbonates with corals and algae are known from the
bmiddleQ Danian onwards (Lein, 1982; Tragelehn,
1996). Non-reefal Paleocene corals assemblages are
also widespread (Kühn, 1930; Kühn and Traub,
1967). Paleocene coral-bearing reefal limestones are
known from the Carpathians of Slovakia (Samuel et
al., 1972). Unfortunately, very little taxonomic information on corals is available from all Alpine
Paleocene reefs, but unpublished data of B.-S. (work
in progress) could be used and the bmorphotypesQ
figured in Tragelehn (1996) could mostly be identified
to the species level.
With few exceptions, the coral faunas from the
Pyrenees (northern Spain and southern France) contribute little to the K–T discussion. Nearly all faunas
previously described as Maastrichtian are now dated
as late Campanian or older (Ardèvol et al., 2000) and
data of Paleocene corals are limited (Alloiteau and
Tissier, 1958).
5.5. Northern Europe and Greenland
Coral faunas have been well studied in several
regions including Denmark, the Limburg region
(Belgium, southern Netherlands, westernmost Germany), central France, northern Germany, eastern
England, and Poland. In the whole region az-like
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corals prevail. However, the late Maastrichtian of the
Maastricht area (southern Limburg) contains a surprising number of z-like taxa (Leloux, 1999), as does
the Danian reef of Vigny (Meyer, 1987). For the
Maastricht region, corals from the upper Meerssen
Member (IVf-6) were considered to be Cretaceous
and all corals from the Geulhem Member were
included in the Danian. We have excluded data from
the topmost Meerssen bed IVf-7, which is Danian at
the type locality but is not readily recognized in other
localities (Leloux, 1999).
The Vigny reef (Montenat et al., 2002) seems to be
a key locality for the evaluation of coral extinctions,
because all corals in this reef were said to belong to
Cretaceous genera (Meyer, 1987). A number of genera
were previously thought to have disappeared from the
fossil record at the end of the Cretaceous or even
earlier and representatives of those genera are often
not reported from other Danian localities. Because
only a few corals were sketched by Meyer and
considering the often-incorrect identifications, we
were reluctant to utilize the taxonomic data as
reported. We have only included figured taxa in our
database, where reliable identifications were possible
from the drawings.
Maastrichtian and especially Paleocene corals are
widespread in the Ukraine and the European part of
Russia (Kuzmicheva, 1987). Nearly all of them are azlike, as are the corals in boundary sections of
Denmark and southern Sweden (Floris, 1979). Fairly
rich az-coral assemblages are known from K–T
sections in the Nuussuaq area of West Greenland
(Floris, 1972). With new biostratigraphic data available (Nøhr-Hansen and Dam, 1997), these assemblages could be reliably assigned stratigraphically.
5.6. Arabia, Pakistan, India and Tibet
Several coral-bearing sections are known from the
Oman Mountains but are best known from the border
region of Oman and the United Arab Emirates
(Metwally, 1996; Baron-Szabo, 2000). A surprising
diversity of corals has been reported from these
deposits, mostly dated as bmiddleQ to late Maastrichtian (Smith et al., 1995). A moderately diverse late
Campanian to early Maastrichtian coral fauna has also
been reported from Saudi Arabia (Abed and ElAsa’ad, 1981). We know of no descriptions of
205
Paleocene corals from Arabia, although some algal
reefs have been described (Rácz, 1979).
K–T corals from India and Pakistan are only
known from old reports (Stoliczka, 1873; Duncan,
1880; Noetling, 1897). Although we could revise their
taxonomy from the published material, the stratigraphic data were sometimes difficult to assign to
modern chronostratigraphy. From the corals reported
by Stoliczka (1873), the recent lithostratigraphic
correlations of Sundaram et al. (2001) were used to
assign ages. All corals reported by Noetling (1897)
were treated as Maastrichtian. The Duncan (1880)
faunas were stratigraphically partitioned according to
the stratigraphic assignments of Eames (1968) and
Adams (1970). Himalayan coral data are scarce.
Faunal lists are available for undifferentiated Campanian–Maastrichtian and Montian strata (Liao and Xia,
1994; Löser and Liao, 2001).
6. Evaluation of global turnover rates
6.1. General considerations
In this section we assess our database to discuss the
magnitude of extinction and compare our results to
published and unpublished analyses. Because we
focus on extinctions related to the K–T boundary,
we emphasize the dataset with confirmed Maastrichtian age (M). Last occurrences of taxa in the late
Campanian and unresolved late Campanian to early
Maastrichtian, however, have to be considered
because corals are relatively rare in comparison to
other invertebrate groups such as bivalves and gastropods. This implies a strong Signor–Lipps effect
(Signor and Lipps, 1982) meaning that the last
reported occurrence of a taxon is unlikely to represent
its evolutionary last occurrence. A coral genus that
has last been sampled in the late Campanian may
therefore well have existed until the K–T boundary or
even above.
Results on extinction rates differ considerably
depending on how singletons are treated. Singletons,
defined as taxa occurring in only one stratigraphic
interval, are continuously discussed in the macroevolutionary literature (e.g., Foote, 2000). In most
recent studies on long-term diversity dynamics,
singletons are excluded from the analyses because
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they have been shown to create artificial patterns (see
section on diversity dynamics). This is because
singletons are thought to be mostly due to monographic effects or exceptional preservation. However,
singletons may also carry an evolutionary signal
representing short-lived clades. Separating those two
hypotheses is difficult and has to be done on a caseby-case basis. Here we define singletons as genera
which were both apparently restricted to one stratigraphic stage and have been described from just one
region. This concept is less rigorous than the uniqueoccurrence concept sometimes applied to microfossil
assemblages (Buzas and Culver, 1998) but is more
confined than the exclusion of taxa known only from
one time interval regardless of geographic distribution
(Foote, 2000).
Results on taxonomic turnover may also be
influenced by peculiar collections with little similarity
to others. We have identified two regions in our
database, which may bias some of the results: the
Maastrichtian of the Antarctic Peninsula and the
Danian of Vigny (France). While the potential bias
of the Antarctic Peninsula is largely balanced by the
exclusion of singletons (most of the unique genus
occurrences are also stratigraphic singletons), the
Vigny fauna poses larger problems. Although all
problematic occurrences have been excluded from our
dataset, the fauna still contains two survivors (Brachyphyllia and Synastrea), which have not been
reported from other Paleocene localities. Therefore,
we have tested the sensitivity of our results to those
faunas by running all analyses with the complete
exclusion of Antarctica and Vigny (-AV data) and
report all noticeable differences.
6.2. Extinction intensities
The approach most widely used to evaluate
extinction intensities is based on the evaluation of
genus ranges. The total generic extinction rate of corals
is 39F8% when we simply count all late Campanian–
Maastrichtian (CM) coral genera (including Lazarus
genera) and extract those that have no record in the
Cenozoic (Table 1). This number is lower but not
significantly different from the result based on
Sepkoski’s compendium (Sepkoski, 2002), which
gives 43F9%. The difference remains approximately
the same when we exclude singletons. The extinction
is lower when only Maastrichtian (M) data are
considered (Table 1). Because the data without singletons are more reliable, our best approximation of the
generic extinction of corals at the K–T boundary is
between 29% (M data) and 34% (CM data). To better
constrain the generic extinction intensity related to the
K–T boundary, we have carried out a simple test of
confidence intervals on reported stratigraphic ranges
of the 11 non-singleton genera that are last reported
from late Campanian or late Campanian–early Maastrichtian localities. We have transferred the usual
metrics reported in meters of section (Marshall,
1990, 1998) to chronostratigraphic ages and calculated
the stratigraphic confidence intervals of those genera
based on the reported range in millions of years and the
number of stratigraphic horizons in which the genera
were reported using the compilation of Baron-Szabo
(2002). The 50% confidence intervals of three of the
ten genera thought to become extinct in the early
Maastrichtian or earlier reach the K–T boundary
(Negoporites, Placohelia, Stephanosmilia). We have
included a random selection of two genera in the K–T
victims category (according to Marshall’s method half
of the taxa whose 50% error bars reach the boundary
should be included in the victims category). Thereby
we achieve 30F8% as our best approximation of endCretaceous scleractinian coral extinctions at the genus
level. The results excluding data from Antarctica and
Vigny ( AV data) are slightly higher. Overall nonsingleton extinction rates are 31% for the M data and
37% for the CM data.
Species-level extinction rates are usually not
provided in global surveys owing to the strong noise
introduced by taxonomic dchauvinotypyT (Rosen,
1988). In spite of our taxonomic revisions, the
apparent species level extinction rate of corals at the
K–T boundary is 63F6% (M data). This value is still
arguably high, perhaps due to sampling problems in
the Paleocene. With reverse rarefaction (Raup, 1979)
we achieve 45F5% species extinctions. We used the
number of species in our recorded Cretaceous coral
genera as a basis for the taxon-size distribution, which
seems to be more reliable than the estimate based on
modern echinoids as done by Raup (1979). However,
reverse rarefaction may generally underestimate species loss (Jablonski, 1995) and thus the true species
level extinction rate may have been somewhat higher
than 45%.
W. Kiessling, R.C. Baron-Szabo / Palaeogeography, Palaeoclimatology, Palaeoecology 214 (2004) 195–223
Our family level extinction rate is slightly lower
than reported in earlier summaries (Barta-Calmus,
1984). Five out of 43 (12%) late Campanian–
Maastrichtian coral families have no Cenozoic record
and only four out of 42 families (10%) from M
localities (including ghost ranges) have no younger
record (Cladophylliidae, Isastreidae, Microsolenidae,
Pachyphylliidae). All these families had only one or
two genera in the Maastrichtian. The one family with
a reported last occurrence in the late Campanian, the
monogeneric Negoporitidae, is generally rare in the
Cretaceous and only reported by one occurrence in the
late Campanian of Spain (Götz, 2003—described as
Actinaraea sp.).
6.3. Gradual or abrupt extinction?
The current resolution of the data does not permit
a conclusive assertion on the rapidity of the coral
extinctions. Nevertheless, our data allow some statements about the sometimes quoted gradual decline of
corals prior to the K–T boundary (Moussavian,
1992). Rosen (in MacLeod et al., 1997) has already
noted that newer data tend to show that coral
extinctions have been more concentrated in the later
Maastrichtian than previously realized. Our database
confirms this: when all CM coral occurrences are
analyzed, the total extinction of non-singleton coral
genera is only 5.5% higher than for M corals. Even
without considering stratigraphic confidence intervals, the late Campanian–early Maastrichtian background extinction rate appears to be unusually low,
which agrees well with the low post-Cenomanian/
pre-Maastrichtian extinction rates from the analysis
of Sepkoski’s dataset (Fig. 4B). From this and from
the long stratigraphic confidence intervals on the last
reported occurrence of coral genera we infer that
coral extinctions may well have been concentrated in
the late Maastrichtian. Because rich coral faunas with
many victim genera are known even from latest
Maastrichtian localities, such as the Maastricht area
(Leloux, 1999) and Jamaica (Mitchell, 2002 and
unpublished data of B.-S. with new stratigraphic data
in Steuber et al., 2002), the extinctions may even
have been concentrated at the K–T boundary itself.
The decline in sampling (reported coral occurrences), however, appears to be gradual in the late
Maastrichtian. In several sections with shallowing
207
facies trends, such as Jamaica (Mitchell, 2002),
Slovenia (Turnšek, 1994), and southern Spain (Götz,
2003), corals disappear even earlier than rudists. If,
like the rudists, scleractinian corals had experienced
complete extinction at the K–T boundary, a gradual
extinction would surely have been inferred for the
corals.
6.4. Survivorship and abundance
Evaluations of extinction risks are based on both
the late Campanian–Maastrichtian (CM) and the
Maastrichtian (M) occurrence matrices. The number
of taxonomic occurrences was used as a rough proxy
for abundance. Although this measure is crude in
comparison to counts of specimens (not available for
all collections), the close relationship between the two
metrics (Buzas et al., 1982; Alroy, 2000; He and
Gaston, 2000; unpublished data of W.K.) supports the
feasibility of this approach.
The assessment of relationships between survivorship and the number of occurrences produces
equivocal results. When Lazarus taxa are included
(zero occurrences in CM, but known from earlier
and later intervals), we achieve a significant relationship between the probability of survival and
abundance ( P=0.021; Mann–Whitney U-test) for
the CM dataset. However, the M dataset produces
no significant dependency ( P=0.143). While the
latter observation is in line with recent analyses on
bivalves saying that survivorship and abundance are
decoupled at the K–T boundary (Lockwood, 2003),
the former test implies that rare genera were indeed
more prone to extinction than abundant genera.
Because the data excluding Antarctica and Vigny
( AV data) confirm that there is no significant
difference in survival probabilities ( P=0.216 and
P=0.654, respectively), we follow Lockwood (2003)
and conclude that abundance was not an insurance
against extinction.
Species-rich genera are not less likely to become
extinct than non-singleton genera with few species,
both in the CM ( P=0.109) and M ( P=0.167) data
(same results with AV data). Previous analyses of
large species-level datasets have similarly resulted in
no significant differences in extinction risk between
species-poor and species-rich genera (Jablonski,
1986a; Smith and Jeffery, 1998).
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6.5. Ecological selectivity of extinctions
Rosen and Turnšek (1989) made a strong case of
selective extinctions of z-like corals at the K–T
boundary and our data confirm their basic result.
The raw data (including the singletons), however,
although showing higher extinction rates of z-like
corals in comparison to az-like taxa (Table 1) produce
no significant differences. The same applies to the
analysis based on Sepkoski’s compendium. However,
significant differences emerge when the analysis is
limited to non-singleton genera (Fig. 5; P=0.001 for
both CM and M; Mann–Whitney U). The difference is
due the commonness of Maastrichtian az-like singleton genera, which are mostly concentrated in the
James Ross Basin of Antarctica (Filkorn, 1994).
When we filter out the AV data, significant differences
are even achieved if singletons are included. The
result supports the view that the end-Cretaceous
extinction was selective against photosymbiosis or
some ecological attribute associated with photosymbiosis (Rosen and Turnšek, 1989).
The difference of non-singleton extinction rates is
also significant between colonial and solitary genera.
However, because coloniality and ecological mode are
closely linked in corals (in our dataset 84% of az-like
taxa are solitary and 95% of z-like taxa are colonial) it
is difficult to separate both factors. For colonial corals
extinction risk was higher for corals with high
corallite integration (meandroid, thamnasterioid) than
for less integrated colonies (CM data, P=0.014; M
data, P=0.024; also significant for the AV dataset).
When, in addition to our four corallite integration
categories, we include solitary forms as having 0
corallite integration, the dependence becomes even
more significant ( Pb0.001).
Larval strategy is another possible selection criterion discussed in the literature. Previous analyses have
shown that no significant correlation between larval
ecology (e.g., planktotrophic versus lecitotrophic) and
extinction risk exists at the K–T boundary (Valentine
and Jablonski, 1986; Smith and Jeffery, 1998). Too
little is known about larval ecology of ancient scleractinians to test this hypothesis. From the few genera (13)
that we can assign to larval feeding mode (all survivors,
assignment based on Edinger and Risk, 1995), nearly
equal proportions are brooders (lecitotrophic larvae) or
broadcasters (planktotrophic larvae). This suggests that
larval feeding mode did not act as a selective criterion
at the K–T boundary.
The mean skeleton size of solitary coral genera had
no significant influence on survivorship ( P=0.69) and
the mean size of Maastrichtian and Paleocene genera
was nearly identical. This suggests that the extinction
did not select against large-sized genera. However, this
result should be viewed with caution because sizes
were compiled on a species and not specimen basis and
the dataset is limited. Comparing the sizes of colonial
corals is hampered by often incomplete preservation.
Extinction rates did also not vary significantly
between habitats. Although there was a tendency for
greater extinction rates of corals inhabiting reefs
(32F11%) the difference to extinction rates in coastal
environments (24F11%), carbonate platforms
(30F9%), and outer shelf environments (29F8%) is
far from being significant. This indirectly demonstrates the importance of keeping dz-coralsT and dreef
coralsT as separate concepts (Rosen, 2000).
6.6. Taxonomic structure of extinctions
Fig. 5. Ecological selectivity of coral extinctions at the K–T
boundary. Extinctions rates depend strongly on feeding mode with
presumably zooxanthellae bearing corals (z-like) being much more
affected than azooxanthellate corals (az-like).
Extinctions are unequally distributed among higher
coral taxa (Fig. 6). Non-singleton generic extinctions
vary significantly between coral suborders ( P=0.002;
Kruskal-Wallis H). Although the differences are
partially due to the unequal partitioning of az-like
W. Kiessling, R.C. Baron-Szabo / Palaeogeography, Palaeoclimatology, Palaeoecology 214 (2004) 195–223
209
Fig. 6. Distribution of genus extinctions in coral suborders. Suborders with mainly az-like genera are little affected, but extinction probability
also varies significantly among z-like groups (e.g., Astrocoeniina and Microsolenina).
genera in suborders (the Caryophylliina and Dendrophylliina with unusually low extinction rates are
exclusively az-like), extinction intensity also varies
significantly between suborders with mostly z-like
genera (e.g., Astrocoeniina and Microsolenina). This
corroborates former results indicating that extinctions
are correlated with taxonomic groups (Smith and
Jeffery, 1998).
6.7. Dead Clade Walking (DCW)
The concept of clades surviving mass extinctions
but becoming extinct soon after has long been of
interest to the scientific community, but only recently
a statistical test was applied, which verified that this is
a general phenomenon after mass extinction events
(Jablonski, 2002). The hypothesis of Dead Clade
Walking (DCW) predicts that significantly more
clades become extinct in the stage after a mass
extinction than during background extinction times.
Our coral data indicate that the DCW phenomenon
is more profound for K–T corals than previously
recognized. Nineteen coral genera surviving the K–T
boundary became extinct in the Paleocene; 12 have
their last occurrence in the Danian. Thus the Danian
has 13% DCWs at the genus level. This is a
conservative estimate, because most taxa of the
apparently DCW-rich coral fauna from Vigny (Meyer,
1987) were not included in our database (see Section
5.5). Removing the two remaining DCW genera only
known from Vigny still makes 11% DCW s. str., a
value significantly higher than the 3% recorded in
Sepkoski’s compendium.
Our data do not allow testing hypotheses of the
cause of the DCW phenomenon. However, the ecology
of coral DCWs gives some indication. Although the
number of z-like genera in the Danian is nearly equal
to the number of az-like genera (59/57) the dichotomy
in the ecological traits of DCW corals is as great as for
the K–T extinction. Z-like DCW genera are more
common than az-like ones (9 and 3 genera, respectively). This suggests that either a similar ecological
selectivity has acted on the survival fauna as is evident
for the end-Cretaceous extinction, or that stochastic
processes preferentially extinguished the more damaged z-like corals in the Danian (bottleneck effect).
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6.8. Changes in standing diversity
6.9. Appreciation of biases
Due to delayed recovery, the K–T boundary is
not only characterized by significantly elevated
extinction rates but also by a pronounced decline
in standing diversity. Both the biodiversity based on
through-ranging genera and the raw pattern of
recorded species and genera indicate that corals
exhibit a marked decline of standing diversity.
However, these raw data have to be normalized
for the strong differences between the Maastrichtian
and Danian fossil record. Because advanced resampling methods for analyses based on through
ranging-taxa (Alroy, 2000) are not applicable (not
sufficient temporal bins available), we used simple
rarefaction to account for differences in sampling.
The result indicates that late Maastrichtian coral
diversity was indeed significantly higher than early/
middle Danian diversity (Fig. 7). However, this
difference is not evident when whole stages are
compared (Maastrichtian and Danian). If the same
limited number of taxonomic occurrences had been
recorded from the Maastrichtian as is present in the
Danian, the number of recorded genera is likely to
be nearly identical. Thus the drop of standing
diversity is unlikely to have persisted for an
extended period after the K–T boundary, which is
in contrast to previous analyses (Rosen and Turnšek,
1989).
Recent analyses of large datasets suggested that
most if not all mass extinction events recognized in
the fossil record are exaggerated and can largely be
explained by heterogeneities of the rock record (Peters
and Foote, 2002). Although there are pitfalls in the
approach of Peters and Foote (2002) (Foote, 2003;
Crampton et al., 2003), the issue has to be discussed.
The difference in the number of sampled corals
between the Maastrichtian and Paleocene is indeed
profound. When recorded and inferred standing
generic diversities are compared, the simple completeness measure, which represents the ratio of taxa
actually recorded in an interval to taxa inferred to
have been present (Fara, 2001), declines from 94% in
the late Campanian–Maastrichtian (CM), 88% in the
Maastrichtian (M), 86% in the whole Paleocene (Pa)
to 71% in the Danian (D). This decline in the quality
of the fossil record results in a severe Lazarus effect
(Jablonski, 1986b). Many surviving genera are
recorded in CM but not in Pa (15 out of 94), six
survivors are known from Pa but not from CM and
three are neither recorded in CM nor in Pa. When the
extinction rate is calculated without considering these
Lazarus taxa, the non-singleton genus extinction rate
rises to 48F8% (CM). However, when the percentage
of Paleocene Lazarus genera is subtracted from the
calculations of extinction rates (based on the assump-
Fig. 7. Comparison of global sample-standardized standing diversities for scleractinian corals in well resolved stratigraphic intervals based on
rarefaction analyses (95% confidence intervals are indicated). Coral diversity in the Upper Maastrichtian is significantly higher than in the
Lower/Middle Danian. Upper Campanian sample-standardized diversity is statistically indistinguishable from upper Maastrichtian or Lower/
Middle Danian ones.
W. Kiessling, R.C. Baron-Szabo / Palaeogeography, Palaeoclimatology, Palaeoecology 214 (2004) 195–223
tion that the same percentage of genera is not recorded
in the Paleocene but did survive the K–T boundary
event), the end-Cretaceous extinction rate would drop
to 29F7% (CM) or 24F7% (M). This is still
significantly above the mean extinction rate for
scleractinian coral genera (12F4% per stage calculated from Sepkoski, 2002) and suggests that the K–T
boundary represents a true mass extinction event for
scleractinian corals.
6.10. Origination rates
The evolution of new genera within the Paleocene
was apparently gradual, but concentrated in the
Danian where 21% of the genera are new. The overall
origination rates derived from our database are almost
identical to those derived from Sepkoski’s compendium (Sepkoski, 2002). A significant difference,
however, is noted for the origination rate of z-like
corals (Table 2). For the whole Paleocene we note
20F9% newly evolved z-like genera, while only
3F6% (0–9%) new genera are derived from the
compendium. Similar to the compendium data, our
recorded origination rates in the Danian are significantly higher for az-like genera than for z-like genera.
However, the difference is not significant when the
whole Paleocene is considered.
Another important observation is the low number
of singleton taxa both in the Danian alone and the
entire Paleocene. The one Danian stage singleton
(Faksephyllia) is not even a singleton according to
our definition above, because it is recorded from a
number of regions (Greenland, Denmark, Austria,
Kazachstan). This contrasts to the eleven Maastrichtian singleton genera (Table 1). Thus there is no
evidence for an expansion of short-lived clades in
the Danian.
The origination of new Paleocene clades was less
heterogeneous among coral suborders than in the
extinction, and the overall dependency is not significant ( P=0.172; Kruskal Wallis test). The Astrocoeniina and Caryophylliina have the relatively strongest
enrichment of new genera, while these groups had
suffered unusually low extinctions at the end of the
Cretaceous (Fig. 6). Apart from the significant
dependence of Danian originations from inferred
ecological mode (az-like versus z-like; P=0.003),
originations seem to have been even more random
211
Table 2
Origination metrics of corals in the Paleocene
N
Sepkoskia
7.3
119
23.0
8.8
87
9.8
7.5
61
3.1
6.0
32
32.8
12.1
58
34.5
12.6
55
20.3
7.3
118
18.3
8.4
82
9.8
7.5
61
3.1
6.0
32
31.6
12.1
57
28.0
12.4
50
Total Paleocene
KTbase
Error
N
Sepkoskia
Error
Total (with
singletons)
z-like (with
singletons)
az-like (with
singletons)
Total (without
singletons)
z-like (without
singletons)
az-like (without
singletons)
27.7
7.7
130
35.6
9.2
104
20.3
9.5
69
6.1
8.1
33
36.1
12.1
61
49.3
11.6
71
26.6
7.7
128
25.6
9.0
90
20.3
9.5
69
3.1
6.0
32
33.9
12.1
59
37.9
12.5
58
Danian
KTbase
Total (with
singletons)
z-like (with
singletons)
az-like (with
singletons)
Total (without
singletons)
z-like (without
singletons)
az-like (without
singletons)
21.0
Error
Error
N
N
All values except N in percent. Errors represent 95% confidence
intervals.
a
Analyses based on data in Sepkoski (2002).
than the extinctions. Origination rates in the Danian
were apparently independent of coloniality and newly
evolved genera occurred as often as surviving genera
and the solitary newcomers were not significantly
different in mean size from survivors.
6.11. Comparison with other databases
As reported above, our extinction and recovery
data are often similar with data derived from Sepkoski
(2002). The overlap of confidence intervals between
analyses based on our and Sepkoski’s compilation is
remarkable because most of the data in Sepkoski’s
compendium can be shown to be incorrect. The most
important source of error is introduced by incorrect
stratigraphic ranges (Fig. 8). Subjective synonyms,
although relatively common (second most common
after unrealized boundary genera), have a limited
effect, because about half the senior synonyms were
212
W. Kiessling, R.C. Baron-Szabo / Palaeogeography, Palaeoclimatology, Palaeoecology 214 (2004) 195–223
Fig. 8. Errors in stratigraphic ranges of coral genera in Sepkoski’s (2002) compilation and their effect on diversity dynamics across the K–T
boundary. The most striking difference is the large number of genera that were not recognized as being present in the K–T boundary interval.
The largest net effect on extinction rate is given by range-extensions of genera formerly thought to become extinct.
not listed as boundary genera in Sepkoski’s compendium. Maastrichtian standing diversity is mostly
affected by the number of genera that were not
considered as boundary genera (occurrence in, or
range through Maastrichtian–Paleocene) in Sepkoski
(2002). However, the net effect on extinction rates is
low because the new boundary genera are approximately equally partitioned between victims and
survivors. The net effect on extinction rates is largest
for the stratigraphic range errors in Sepkoski’s
compendium. Especially the number of previously
unrealized boundary genera affects calculated turnover rates. The same is true for originations in the
Paleocene. While the number of wrong originations is
almost exactly matched by the number of new
originations from erroneous boundary crossers, 18
new Paleocene originations (mostly z-like) are recognized from a downward extension of first occurrences.
Another recent detailed comparison of diversity
dynamics extracted from Sepkoski’s compendium
(trilobites) and those compiled from taxonomic experts, has resulted in nearly identical patterns (Adrain
and Westrop, 2000). Similar to our results Adrain and
Westrop (2000) noted that more than 50% of data in
Sepkoski (2002) are wrong due to taxonomic noise
and a false assignment of first and/or last occurrences.
However, our data show a systematic tendency
towards lower extinction rates, a higher number of
DCWs and a greater proportion of newly evolved zlike genera as compared to Sepkoski’s compendium.
Although we agree with Adrain and Westrop (2000)
that the basic evolutionary patterns can be correctly
derived from analyses of Sepkoski’s database, a
critical review is clearly necessary when analyzing
mass extinctions.
There is the great difference in extinction rates
between our database and the analysis of Rosen and
Turnšek (1989), who have indicated a 60% extinction
at the genus level and 97% at the species level.
Although we could confirm that non-singleton z-like
corals where significantly more affected than az-like
corals, the dichotomy is manifested at 44% versus
13% (CM data) rather than the 70% versus 40% given
by Rosen and Turnšek. The great difference in results
is probably (1) due to the unrevised taxonomic dataset
Rosen and Turnšek have used and (2) a result of the
large stratigraphic intervals they have considered
(Late Cretaceous and Paleocene). It is difficult to
judge which bias is greater. We see, however, that
taxonomic revisions are very important when evaluat-
W. Kiessling, R.C. Baron-Szabo / Palaeogeography, Palaeoclimatology, Palaeoecology 214 (2004) 195–223
ing mass extinctions or diversity dynamics in general
(see also Jablonski et al., 2003).
7. Geographic patterns
213
region size to the number of occurrences. We discuss
analyses based on intermediate (19 coral-bearing
regions), coarse (10 regions) and very coarse (4
regions) geographic scales. The analysis based on an
objective definition of regions applies 308 latitude–
longitude grids and detects 21 coral-bearing regions.
7.1. Definitions of regions
7.2. Geographic patterns of extinctions
Patterns of assemblage composition, extinction and
recovery have been analyzed at different geographic
scales. Compared to similar studies using a 108 grid
(Raup and Jablonski, 1993), our scale had to be coarser
because coral data are much scarcer. We conducted
analyses of extinction and recovery rates on variable
geographic scales from basin to hemisphere, applying
both objective (grids) and subjective (biogeographic
regions) definitions of areas. While objective criteria
for defining regions have the advantage of permitting a
neutral approach to geographic patterns, subjective
criteria can provide a more natural definition of
geographic regions. Moreover, the filter of z10 genera
per region is the more severe, the smaller the regions
are defined. This can partly be balanced by adjusting
Geographic extinction patterns of the late Campanian–Maastrichtian (CM) dataset agree well with those
of the more restricted Maastrichtian dataset. To maximize sample size and statistical confidence we have
therefore used the larger CM dataset for the discussion.
Extinction intensities are apparently not randomly
distributed geographically when stratigraphic singletons are excluded. The highest extinction rates are
recorded in low paleolatitudes, and drop off towards
higher palaeolatitudes. The pattern is best visible on
manually defined regions at the intermediate geographic resolution (Fig. 9) but the same basic results
were achieved with other definitions of regions.
Sample sizes, even in broader defined regions, are
Fig. 9. Geographic pattern of end-Cretaceous coral extinctions. Extinction rates per biogeographic region (intermediate scale) for late
Campanian–Maastrichtian non-singleton coral collections plotted on the geographic mean of occurrences. Black: Percentage of extinct genera
recorded in the region. White: Percentage of genera recorded in the region known to be extant in the region or elsewhere after the K–T
boundary. Size of circles is proportional to the number of genera in each region. Regions with less than 10 genera have been filtered out.
214
W. Kiessling, R.C. Baron-Szabo / Palaeogeography, Palaeoclimatology, Palaeoecology 214 (2004) 195–223
too small and confidence intervals too large, to produce
significant differences even at the coarsest geographical scales. The general tendency towards lower extinction rates in higher latitudes is likely to be an artifact
of latitudinal variations in the composition of regional
assemblages. Limiting the analysis to z-like corals
produces no latitudinal cline in extinction rates (Fig.
10), nor any significant differences between geographic regions. This suggests that all patterns are strongly
controlled by the relative contribution of the extinctionresistant az-like corals to regional assemblages and
latitude had no direct influence of extinction rates.
Similar to previous analyses on bivalves (Raup and
Jablonski, 1993), we do not detect any hot spots in the
extinctions. Although the Mediterranean Tethys constantly exhibits slightly elevated extinctions in comparison to other regions, both for whole faunas and for
only z-like corals, the differences are small and far
from being statistically significant. Even on very large
geographic scales, non-singleton CM extinction rates
are similar and do not deviate markedly from the
global mean: North America, 33F11%; Europe,
35F10%; Africa and India, 30F11%. The same
applies for endemics to regions. Even the raw data
indicate little variation in extinction risk of endemics
(North Africa, 50%; Europe, 50%; North America,
40%) and statistical errors are so large that no
conclusive statement is possible. It is evident from
this analysis, however, that North America was not a
hot spot in end-Cretaceous coral extinctions.
7.3. Extinction risk and geographic distribution
Geographic ranges of CM corals were measured by
the number of geographic regions in which coral
genera are recorded. We have tested the relationship
between survivorship and geographic distribution on
different geographic scales. As in the previous
analyses, stratigraphic singletons were excluded.
There is a clear relationship between extinction
risk and geographic distribution, which is nearly
independent of how the regions are defined.
Survivorship is most clearly linked to geographic
distribution in the 308 binning analysis ( P=0.002;
Mann–Whitney U) but the relationship is also
significant at the intermediate ( P=0.004) and
coarse geographic scales ( P=0.009). Even when
including survivors without reported occurrences in
the late Campanian–Maastrichtian (Lazarus genera,
number of regions=0), the relationship remains
significant. These results strengthen previous analyses, which conclude that wide geographic distributions of clades enhance the probability of
survivorship at the K–T boundary (Jablonski and
Raup, 1995) and other mass extinction intervals
(Jablonski, 1995).
Fig. 10. Latitudinal pattern of extinction rates. Extinction rates for all corals (black diamonds) and for z-like corals (white squares) calculated for
208 latitudinal bands. Error bars demarcate 95% confidence intervals in each direction. Data only shown for bands with more than 10 genera in
each band. The apparent latitudinal cline in extinction rates is not seen in z-like corals suggesting that it is controlled by the relative abundance
of extinction-resistant az-like corals.
W. Kiessling, R.C. Baron-Szabo / Palaeogeography, Palaeoclimatology, Palaeoecology 214 (2004) 195–223
215
Fig. 11. Geographic pattern of the proportions of incoming genera in Danian (A) and Paleocene (B) regional (coarse scale) assemblages.
Apparent origination rates per region (coarse geographic resolution) are indicated. Black: Percentage of newly evolved genera in the region.
White: Percentage of surviving genera (known from here or elsewhere prior to the K–T boundary) in the region. Size of circles is proportional to
the number of Danian (A) and Paleocene (B) genera in each area. Areas with less than 10 genera have been filtered out. Percentage values in (A)
indicate the proportion of DCWs in the surviving genera in the region. Bold numbers indicate number of region.
216
W. Kiessling, R.C. Baron-Szabo / Palaeogeography, Palaeoclimatology, Palaeoecology 214 (2004) 195–223
7.4. Geographic patterns of recovery
Although the locus of evolution of new genera in
the Paleocene is unlikely to be captured by our
database approach, the distribution on new genera in
regional assemblages may give some insight into
evolutionary dynamics. Due to the Danian Lazarus
effect (29% of all coral genera, inferred to be
present, are not recorded), the regional assemblages
are generally richer in new genera than the global
average (Table 2) and thus have to be interpreted
cautiously. New genera in the Danian were nearly
uniformly distributed (Fig. 11A). However, averaged
over the whole Paleocene, the percentage of Paleocene genera relative to Cretaceous survivors is
apparently higher in low latitudes (Fig. 11B),
although the maximum difference is noted between
North Africa (region 8, 35F14% new genera of total
faunas) and the Gulf of Mexico (region 6, 19F15%),
which were both in the tropics in the Paleocene.
Latitudinal differences become greater when only zlike corals are analyzed. New z-like corals in the
Paleocene are much more common in North Africa
(45F22%) and Pakistan (40F21%) than in Northern
Europe (8F14%), while the Mediterranean Tethys
(14F13%) and the Gulf of Mexico region (14F18%)
have intermediate concentrations of new z-like
genera. This statement must also be qualified,
because reported generic richness is low for all
regions except Europe (large binomial errors) and the
Paleocene coral record of Northern Europe is
basically limited to the Danian. However, the
apparent concentration of new Paleocene z-like
genera in low latitudes (Fig. 12), may represent a
biological signal, even though the latitudinal gradient
is not significant. Our data do not allow testing for
the concentration of invaders and bloom-taxa, which
have been noted to be exceptional in Gulf Coast
bivalve faunas (Jablonski, 1998).
The previous observation of an increase of
cosmopolitanism of corals in the Paleocene (Rosen
and Turnšek, 1989) cannot be confirmed by our data.
The mean number of geographic regions in which
coral genera are recorded does not differ significantly
between the Maastrichtian and the Danian, nor does it
differ between the late Campanian–Maastrichtian and
the Paleocene.
Danian DCW genera are only recorded in northern
Europe (7 out of 33 surviving genera are DCWs),
southern Europe (5/30), North Africa (3/18) and India
(2/4). Other regions with a moderate or good Danian
record such as the Gulf Coast/Caribbean and South
America do not show any DCWs. Thus DCWs seem
to be concentrated in Eurasia and Africa. This implies
that the DCW phenomenon was indeed influenced by
spatial distribution (Jablonski, 2002), but it was not a
Fig. 12. Latitudinal pattern of Paleocene origination rates. Origination rates for all corals (black diamonds) and for z-like corals (white squares)
calculated for 208 latitudinal bands. Error bars demarcate 95% confidence intervals in each direction. Data only shown for bands with more than
10 genera in each category. A latitudinal cline of origination rates can be seen for z-like corals (but note large confidence intervals).
W. Kiessling, R.C. Baron-Szabo / Palaeogeography, Palaeoclimatology, Palaeoecology 214 (2004) 195–223
simple function of paleolatitude. Another noteworthy
feature is the fact that the majority of Danian DCWs
are just recorded from one region (7) and no DCW is
recorded from more than two regions. This confined
distribution of DCW genera substantiates the interpretation that the DCW phenomenon is probably
caused by bottleneck effects (Jablonski, 2002).
8. Discussion and conclusions
The currently best approximation of scleractinian
coral extinction rates at the K–T boundary is 30F8%
at the genus level. This is among the lowest extinction
rates thus far reported from benthic marine invertebrates at the K–T boundary. Even the echinoids,
although as thoroughly revised taxonomically as our
dataset (Smith and Jeffery, 2000) still exhibit a generic
extinction rate of 36% (Smith and Jeffery, 1998). The
low extinction rate recorded for corals is in line with
recent interpretations that the magnitude of mass
extinction events is probably strongly exaggerated
(Peters and Foote, 2002). However, balancing for
Lazarus effects caused by variations in the quality of
the fossil record, verifies that the K–T boundary
represented a true mass extinction event for scleractinian corals. The reason for the lower extinction rate in
comparison to previous analyses is only partially due
to the improving fossil record in the Paleocene:
several genera previously thought to become extinct
at the K–T boundary (e.g., Calamophylliopsis, Phyllocoenia, Synastrea), have now been recorded from
the Danian and changed from the victims into the
Dead Clade Walking category (Jablonski, 2002), that
is, they survived the K–T boundary but became
extinct soon after, probably due to bottleneck effects.
The greatest effect on extinction rates is given by the
specimen-based taxonomic revisions. While the net
effect of the improving Paleogene fossil record is
nearly balanced by the also improving Maastrichtian
record (with new victims discovered, which were
previously thought to have become extinct before),
the specimen-based comparisons of Cretaceous and
Cenozoic corals (B.-S.) have extended many stratigraphical ranges across the K–T boundary.
The most significant dependency of extinction
intensities in scleractinian controls is seen in their
ecological mode. Corals inferred to have hosted
217
zooxanthellate symbionts in their tissues (z-like
corals) are much stronger affected than corals inferred
to have lacked these symbionts (az-like corals). This
difference, previously noted by Rosen and Turnšek
(1989) on a geographically confined dataset, is now
confirmed at global scales. However, it is not justified
to speak of a breakdown of photosymbiosis at the K–
T boundary. Although az-like corals dominate Danian
assemblages, the extinction of z-like corals was not as
severe as previously thought and coral reefs, often
dominated by z-like corals, are more common than in
the Maastrichtian. A global z-like coral reef gap, if it
existed at all, is confined to the earliest Danian.
An additional ecological link to extinction risk is
indicated by morphological complexity, as measured
by coloniality and colony integration. Although we
can hardly envision a mechanism how a mass
extinction could select against coloniality in itself,
chances are that complexity (see McShea, 1996, for
definitions of complexity) played a role in the
selective extinction as has been previously suggested
(Rosen and Turnšek, 1989). The higher the colony
integration the greater was the chance of extinction at
the K–T boundary. There is some evidence that corals
with a high integration of corallites also have a high
physiological integration (Rosen, 1986; Coates and
Jackson, 1987). Colony integration could also be
related to physiological dependency on photosymbiosis, but current data are too limited for a conclusive
statement. A quantitative test of modern corals based
on the compilation of Stimson et al. (2002) did not
produce significant relationships between corallite
integration and zooxanthellate densities, which are
thought to be related to survival probability during
bleaching events (Stimson et al., 2002). Zooxanthellae
densities are usually low in meandroid as well as
solitary forms and have a maximum in plocoid and
cerioid forms. This might indicate that there was
indeed a general selection against physiological
complexity in the end-Cretaceous extinctions.
Widespread geographic distribution in the Maastrichtian formed an insurance against extinction at the
K–T boundary. Apart from this observation, geographic patterns of extinction and recovery are
indistinct, perhaps owing to the limited sample sizes.
Extinction rates are slightly elevated in low paleolatitudes, but this is exclusively due to the variable
proportions of extinction-resistant az-like genera in
218
W. Kiessling, R.C. Baron-Szabo / Palaeogeography, Palaeoclimatology, Palaeoecology 214 (2004) 195–223
regional assemblages. The nearly uniform extinction
pattern of z-like corals and endemic clades underlines
the idea that the end-Cretaceous extinctions were
caused by a global catastrophe rather than a combination of regional factors. Patterns of recovery are
somewhat more variable geographically, especially
for z-like corals. New Paleocene z-like clades were
proportionally more common in low latitudes than in
higher latitudes. If the regional proportions of new
genera have some relationship to geographic variations in evolutionary rates, we could conclude that the
tropics represented a source of evolutionary novelty in
the post-extinction phase. The pattern of the tropics as
a source of evolutionary novelty has also been
observed during background extinction times (Jablonski, 1993). Because the latitudinal cline is not
statistically significant, any far-reaching conclusion
has to remain open until more data become available,
especially from the Southern Hemisphere.
In spite of conjectured deviations in the mode of
coral evolution from the normal bDarwinianQ pattern
(Veron, 1995), there are striking similarities with other
benthic invertebrates in the macroevolutionary
response at the K–T boundary (Table 3). Because
now three major groups of benthic invertebrates
(bivalves, echinoids, corals) have been rigorously
analyzed at a global scale, we can start to extract
general rules for the K–T boundary. In line with
previous analyses we have found few significant
Table 3
Comparison of generic survivorship dependencies valid for different
groups of benthic invertebrates
Trait
Coralsa
Bivalvesb
Echinoidsc
Abundance
Feeding strategy
Coloniality
Larval strategy
Infaunal vs. epifaunal
Habitat
Size of solitary genera
Taxonomic structure
Number of species in genus
Geographic range
Endemic to America
no?
yes
yes
no?
N.A.
no
no
yes
no
yes
no
no
no?
N.A.
no
no
no
no
yes?
no
yes
no
no
yes
N.A.
no
no
no
?
yes
no
no
yes
N.A.—not applicable.
a
Scleractinians only (this study).
b
Raup and Jablonski (1993), Jablonski and Raup (1995),
Lockwood (2003).
c
Smith and Jeffery (1998).
dependencies of extinction risk and we follow
Jablonski (1986a) that the main reason may lie in
macroevolutionary particularities of mass extinctions.
However, feeding strategy clearly has a major
selective effect in the extinction of corals and
apparently in the other groups as well. While in
corals simple zooplankton feeding seems to have been
more advantageous to survival than a combination of
photosymbiosis and zooplankton feeding; deposit
feeding was apparently favored in bivalves (Rhodes
and Thayer, 1991; but see Jablonski and Raup, 1995
for alternative explanations); and omnivory or selective detritivory enhanced survivorship in echinoids
(Smith and Jeffery, 1998; Jeffery, 2001). Similarly,
extinctions were heterogeneously distributed among
higher taxonomic levels in all three groups, but
species richness in clades did not influence their
survivorship. Previous studies also agree that abundance per se had no significant effect on extinction
risk (Smith and Jeffery, 1998; Lockwood, 2003). The
results are less uniform regarding geographic distribution. Our results agree with Raup and Jablonski
(1993) in that survivorship was more likely when
clades where widely distributed geographically and
there was no geographic hotspot of extinctions. Smith
and Jeffery (1998), however, note the opposite in both
cases, identifying North America as the region with
the highest extinction risk. While this would be in line
with an impact scenario as a cause for the extinction
(enhanced effects proximal to the Chicxulub site), we
argue that the echinoid data alone are too weak to
confirm this.
Although our study provides important results on
macroevolutionary processes around the K–T boundary, we have no direct hint on the cause(s) of the
mass extinction. Our data show no evidence for long
term-climate change as a dominant trigger of the
extinction, because extinctions of z-like corals, which
are very sensitive to climate change today, have no
relationship with paleolatitude. Global cooling would
predict enhanced extinctions in high latitudes,
whereas global warming should have led to stronger
extinctions in the tropics. Ecological selectivity of the
extinction is compatible with incident light reduction
(dust and sulfate aerosols) caused by a bolide impact,
but several other processes could also explain the
pattern. Geographic links to the Chicxulub impact
can hardly be seen in our data; there was no
W. Kiessling, R.C. Baron-Szabo / Palaeogeography, Palaeoclimatology, Palaeoecology 214 (2004) 195–223
extinction hotspot in southern North America. The
apparently delayed recovery of coral reefs, the
absence of DCWs and the somewhat (but not
significantly) reduced recovery in the Caribbean
and North America, are the only hints that this
region was peculiar.
Acknowledgements
We are especially grateful to David Jablonski
(University of Chicago) and Brian Rosen (Natural
History Museum, London) for their constructive and
helpful reviews. Martin Aberhan (Museum of Natural
History, Berlin), David Lazarus (Museum of Natural
History, Berlin) and Dennis Opresko (Oak Ridge
National Laboratory) also provided valuable comments. Jacob Leloux and Jean-Michel Pacaud are
thanked for their help in organizing missing literature.
We thank the German Research Foundation (DFG)
for supporting the projects Ba 1830/3 and Ki 806/1.
This is Paleobiology Database publication #26.
Appendix A. Supplementary data
Supplementary data associated with this article can
be found, in the online version, at doi:10.1016/
j.palaeo.2004.05.025.
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