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

aInstitute of Paleontology, Museum of Natural History, Humboldt-University Berlin, Invalidenstr. 43, 10115 Berlin, Germany bSmithsonian Institution, Department of Zoology, National Museum of Natural History, W-329, MRC-163, Washington, DC 20560, USA

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.2004.05.025 196 W. Kiessling, R.C. Baron-Szabo / Palaeogeography, Palaeoclimatology, Palaeoecology 214 (2004) 195–223

1. Introduction compatible with the impact scenario at the K–T boundary. The mass extinction at the Cretaceous–Tertiary boundary (K–T, officially Cretaceous–Paleogene) still provokes many questions considering the rea- 2. Database and methods sons, magnitude, speed, selectivity and geographic patterns of extinction and subsequent recovery. The 2.1. Database Chicxulub impact is currently considered the most likely culprit for the mass extinction, but apart from Data in this paper are part of a comprehensive other potential causes, such as Deccan Trap volcan- geographic database on the K–T boundary (KTbase), ism (McLean, 1985; Courtillot, 1999), the impact- which currently comprises geological, mineralogical kill effect remains ambiguous in many instances. and paleontological information from 490 K–T Extinctions in some clades were severe, while others boundary sites (Kiessling and Claeys, 2001; Claeys show almost no taxonomic change at the K–T et al., 2002). A K–T boundary site is defined as an boundary. The most dramatic examples were area (100 km2 or less) where at least Maastrichtian reported from the marine plankton, where calcareous and/or Danian sediments are preserved. Closely nannoplankton and planktonic foraminifers faced spaced sections in the same depositional environment extremely high extinction (Gartner, 1996; Arenillas are combined for the summary data (e.g., sedimentol- et al., 2002), while radiolarians and dinoflagellates ogy and geochemistry) but paleontological data are only show some ecological response but almost no partitioned as finely as possible from published extinction (Hollis, 1997; Brinkhuis et al., 1998). sources. For the particular purpose of this study, the Among invertebrate macrofossils, detailed studies concept for K–T boundary sites has been widened to have revealed a similar though less extreme dispar- include also sites with only late Campanian or late ity: bivalves including the rudists exhibit a 63% Paleocene sediments preserved. This was done to generic extinction (Raup and Jablonski, 1993), compare the extinction and recovery at different whereas the extinctions in sea urchins was shown temporal scales. to have been not more than 36% (Smith and Jeffery, Data in KTbase were largely extracted from the 1998). Clearly, additional datasets have to be published literature but personal observations and analyzed at global scales to extract general patterns unpublished data are also included. Paleontological of the end-Cretaceous mass extinction and to get a data are stored in faunal lists representing individual better idea of the underlying cause(s). fossil collections. Each taxon reported in a collection This study documents global geographic patterns constitutes a taxonomic occurrence in the database. of extinction and recovery in scleractinian corals in Abundance data are available for less than one third of the Maastrichtian and Paleocene. We emphasize five the taxonomic occurrences. We thus used the number key points: of occurrences as an approximation of recorded abundances and applied this criterion for resampling. 1. the intensity of the end-Cretaceous coral extinc- All taxonomic data and some 80% of the occur- tions and the pace of recovery; rence data have been taxonomically revised at the 2. the ecological selectivity of coral extinctions; species-level (Baron-Szabo, 2002;B.-S.workin 3. the impact of coral extinctions on the global reef progress) and stratigraphic assignments were updated ecosystem; for the purpose of this study. The occurrence table 4. the geographic patterns of coral extinction and with updated taxonomic information is available as an recovery; electronic appendix (electronic appendix see Back- 5. a comparison of the results with other benthic ground Data Set). Only taxonomic data resolved at invertebrate groups. least to the genus level are represented in KTbase. Species-level taxonomic information is stored in the Although we do not focus on potential causes of the database whenever possible. Coral data from 228 late coral extinctions, we also discuss if our results are 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 460 460 valid species). The main reason for the low species. The low ratio of occurrences to species occurrence/diversity ratio is probably the way coral richness indicates that (1) corals in this time interval data are usually reported in the literature. Most coral are underexplored and the true biodiversity is far from collections stem from whole outcrops or local areas being completely known (e.g., very high), (2) coral and usually combine several beds or even lithostrati- occurrences are usually reported on rather large spatial graphic members. This limits the applicability of and temporal scales, and/or (3) in spite of our resampling techniques such as rarefaction. taxonomic revisions the corals may still suffer over- Sepkoski’s compendium of the stratigraphic ranges splitting. Our revisions have reduced the taxonomic of marine animal genera (Sepkoski, 2002) was used to noise considerably by removing many subjective determine long-term diversity dynamics of scleracti- synonyms (632 nominal species were parsed into nian corals and to compare two independent global 198 W. Kiessling, R.C. Baron-Szabo / Palaeogeography, Palaeoclimatology, Palaeoecology 214 (2004) 195–223 datasets at the K–T boundary. Sepkoski’s data were a homogeneous taxonomic concept and specimen- largely derived from the secondary literature and give based revisions more important than phylogenetic a comprehensive but probably biased view of evolu- analyses when diversity dynamics are analyzed at the tionary dynamics across the K–T boundary, due to the species and genus level. lack of taxonomic expertise in the compendium. Previous comparisons have demonstrated that 2.3. Stratigraphic framework although many data in the compendium are inaccu- rate, large-scale diversity patterns are little affected A thorough revision of all latest Cretaceous and because the errors are randomly distributed (Adrain Paleocene coral collections was required due to the and Westrop, 2000). Our comparison intended to often obsolete biostratigraphic data in older mono- provide a further test of the effect of errors in the graphs, revised chronostratigraphic assignments of compendium. Late Cretaceous foraminiferal zones (Robaszynski and Caron, 1995; Arz and Molina, 2002), the now 2.2. Taxonomic framework formal definition of the Campanian–Maastrichtian boundary (Odin, 2001; Odin and Lamaurelle, 2001) The large morphological variation of colonial and new Strontium isotope data from key localities corals traditionally has caused problems in their (Swinburne et al., 1992; Steuber et al., 2002). Several identification. Not only the overall shape of corals coral collections formerly thought to be of Maas- can vary substantially but also the corallite structure trichtian age (Alloiteau, 1952b; de la Revilla and can change within a colony (Veron, 1995). As Quintero, 1966; Tchechmedjieva, 1986) have been exemplified and discussed by Baron-Szabo (2002), shown to fall into the Campanian, whereas others, our supraspecific taxonomy is mostly based on formerly imprecisely dated as Campanian or early macrostructural characters, which for modern corals Maastrichtian, are now dated as late Maastrichtian have been shown to better accord with molecular data (Mitchell, 2002; Steuber et al., 2002). A reliable than the often used micro-skeletal characters (Veron et revision of stratigraphic assignments was possible for al., 1996). We are aware of pitfalls in this approach about 80% of our coral collections. but currently see this as the best way to integrate the Because coral-bearing deposits are rarely dated taxonomy of ancient corals where molecular data are precisely, and because the global approach has to not available and population dynamics cannot be include some problematic stratigraphic assignments, assessed in detail. Even the molecular approach we limit our discussion to standard stages and currently results in contrasting hypotheses on coral substages. More specific stratigraphic assignments evolution (Romano and Cairns, 2000). Phylogenetic are occasionally available but have been subsumed systematics based on skeletal characters may solve into those larger intervals. Our database separates late issues in some taxa (Cairns, 1997, 2001) but are Campanian (uc), unresolved late Campanian–early difficult to apply in colonial forms. Veron (1995) Maastrichtian (cm), early Maastrichtian (lm), unre- suggested that hybridization and homoplasy is so solved Maastrichtian (m), bmiddleQ to late Maastrich- common in corals that their dreticulate evolutionT will tian (mum), late Maastrichtian (um), early to middle always lead to equivocal cladograms. Moreover, the Danian (ld), late Danian (ud=Montian), unresolved scleractinian skeleton itself is sometimes seen as just a Danian (d), Selandian (mp), unresolved Paleocene (p) grade of organization without phylogenetic signifi- and Thanetian (up) ages. The numeric distribution of cance (Stanley, 2003). occurrences and reported species richness in seven Judging from previous comparisons between phy- stratigraphic intervals is summarized in Fig. 2. For the logenetic and typological approaches to analyzing analyses of extinction and recovery the data are usually long-term faunal change in corals (Johnson, 1998), we partitioned into four intervals: the first combines all are confident that results from our typological data from the late Campanian to the late Maastrichtian approach will not significantly differ from more (CM), the second is limited to occurrences of sophisticated phylogenetic analyses (but see Smith confirmed Maastrichtian age (M), the third comprises and Patterson, 1988 for a different view). We consider 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 species in those clades may host photosymbionts. (Pa). A comparison between CM and Pa has the Haimesastraea, although showing morphological advantage that sample sizes are maximized and the characteristics of z-like corals, has also been grouped time intervals are of similar duration (10 My), while with az-corals because of its commonness in high the comparison between M and D data provides a finer latitudes and siliciclastic environments (personal stratigraphic resolution. observations). It could be argued that genera of mixed trophic mode could as well be assigned to both the az- 2.4. Ecological categories like and z-like categories. However, the philosophy behind our approach is to separate taxa with optional Corals are traditionally categorized as either photosymbiosis from those which depend on photo- hermatypic (reef building) or ahermatypic (non reef symbiosis for survival. building), which is often treated as being equivalent to In comparison to bivalves, gastropods and echi- symbiont bearing (zooxanthellate) versus non-sym- noids, the categorization of other ecological traits is biont bearing (azooxanthellate). This view has been limited in fossil corals. Colony size has some relation critically assessed by Rosen and Turnsˇek (1989) and to physico-chemical variables but the response varies Rosen (2000), who suggested the terms z-like for significantly between clades (Veron, 1995) and thus forms resembling modern zooxanthellate corals and has no predictable relationship with ecological factors. az-like for forms that are more similar to modern Similar problems apply to coral shape. In addition to azooxanthellate corals. Although there is always some photosymbiosis, we have categorized colonial versus uncertainty with the ecological categorization of solitary growth, maximum adult size and colony extinct taxa, the lists of criteria compiled in Wilson integration. Colony integration was measured in four and Rosen (1998) and Rosen (2000) define a ordinal intervals: low (dendroid, phaceloid), medium homogenous protocol, thereby minimizing subjective (cerioid, plocoid), high (thamnasterioid) and very high errors. We categorized ecology at the species level, (meandroid). but for our analyses, categories have been assigned to genera. All genera containing one or more az-species 2.5. Assessment of data were assigned to the az-like category for this purpose. Genera such as Oculina and Cladocora are thus All too often, paleontological data are presented as generally treated as az-corals even though some extant such without any indication of statistical errors. 200 W. Kiessling, R.C. Baron-Szabo / Palaeogeography, Palaeoclimatology, Palaeoecology 214 (2004) 195–223

Because coral data are limited in comparison to other analyze extinction and recovery of corals based on macroinvertebrates such as bivalves and gastropods, multiple sources and discuss ecological selectivity and an indication of statistical confidence limits is geographic patterns. Their basic results were that (1) imperative to judge the significance of reported total generic extinctions were around 60% (see also percentage data. We followed Raup (1991) to Rosen in MacLeod et al., 1997); (2) z-like corals were indicate binomial errors on percentage values. These more strongly affected by the end-Cretaceous extinc- binomial errors reflect uncertainties of the percentage tions than az-like corals; (3) az-like corals became values depending on sample size (N)andthe relatively more common in the Paleocene; (4) biogeo- percentage values themselves. We report binomial graphic differentiation decreased in the Paleocene; errors as 95% confidence intervals in the text. In and (5) Paleocene coral reef development was nearly comparison to standard errors, this has the advantage independent of the previous extinction and cannot be that significant differences can immediately be viewed as a recovery phenomenon. recognized. The limited number of taxa implies fairly While these analyses provided an in-depth review, large confidence intervals implying considerable they have some shortcomings. Firstly, and most uncertainties on reported rates and differences. importantly, the taxonomic data were extracted from Statistical relationships between diversity dynamics the published literature without revisions. Secondly, and other attributes were generally tested with the analyses of extinction patterns were carried out on nonparametric statistics. Nonparametric statistics are rather large temporal scales (three intervals in the preferred because they do not assume that data are Cretaceous, one in the Paleocene) and some strati- normally distributed. Test results are reported in the graphic assignments were incorrect. Four out of text by the probability of randomness ( P). twelve of the bMaastrichtianQ localities indicated by Rosen and Turnsˇek (1989; Table 1) are now dated as late Campanian or older. Thirdly, the data were 3. Corals at the K–T boundary—state of the art analyzed from selected regions, rather than globally. Other original papers exploring corals at the K–T Only a handful of papers have systematically boundary were devoted to paleogeographic distribu- explored the fate of scleractinian corals around the tions of selected taxa (Beauvais and Beauvais, 1974) K–T boundary. Most noteworthy are the papers of and the fate of coral families (Barta-Calmus, 1984). Rosen and Turnsˇek (1989) and Rosen (2000) which All studies agree that the end-Cretaceous mass

Table 1 Extinction metrics of corals at the KT boundary Late Campanian–Maastrichtian data KTbase Error N Sepkoskia Error N Total (with singletons) 39.0 7.7 154 43.2 8.9 118 z-like (with singletons) 45.0 9.8 100 49.2 12.5 61 az-like (with singletons) 27.8 11.9 54 36.8 12.5 57 Total (without singletons) 34.3 7.8 143 38.0 9.2 108 z-like (without singletons) 43.9 9.8 98 47.5 12.7 59 az-like (without singletons) 13.3 9.9 45 26.5 16.5 49

Maastrichtian data only KTbase Error N Total (with singletons) 34.3 7.8 143 z-like (with singletons) 39.6 10.0 91 az-like (with singletons) 25.0 11.8 52 Total (without singletons) 28.8 7.7 132 z-like (without singletons) 39.6 10.0 89 az-like (without singletons) 9.3 8.7 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) 195–223 201 extinction affected scleractinian corals severely, but at climbed sharply in the late Oligocene and Neogene the same time call for more detailed studies. (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 4. Long-term ecological and evolutionary pattern coral reef production in the Paleocene contrasts earlier of scleractinian corals statements saying that the Paleocene was a time of depressed reef building (Bryan, 1991; Vecsei and 4.1. Late Cretaceous–Paleogene reefs Moussavian, 1997). The sedimentary environments of our coral collec- Looking at the fossil record of reefs, corals appear tions parallel the trend in reefs. Some 23% of all late to have benefited from the end-Cretaceous mass Campanian to Paleocene coral occurrences in KTbase extinction. Late Cretaceous shallow tropical shelves are reported from reefs (see Flu¨gel and Kiessling, were often dominated by rudists. These unusual 2002a for definition). Only 16F3% of the late bivalves were abundant nearly everywhere on carbo- Campanian and Maastrichtian (CM) coral occurrences nate platforms, sometimes even forming reefal struc- are from reefs, but significantly more Paleocene coral tures (Johnson et al., 2002; but see Gili et al., 1995, occurrences (36F5%) are known from reefal settings. for a different view). Corals instead, while also These data contrast with the partitioning of az-like and common, rarely achieved rock-forming abundance in z-like corals in both intervals. While z-like corals the Late Cretaceous, but in the Paleocene, corals constitute 59F4% in assemblages of CM, only dominate the global reef factory by far (Kiessling et 37F5% of all reported Paleocene occurrences are z- al., 1999). There is little evidence that rudists directly like (Fig. 1). These global data agree with the outcompeted corals (Gili et al., 1995; Skelton et al., geographically more restricted analysis of Rosen and 1997; Go¨tz, 2003). The relative scarcity of latest Turnsˇek (1989). The discrepancy between z-like coral Cretaceous coral reefs has been attributed to the and reef distributions has two reasons. Firstly, several absence of encrusters (Moussavian, 1992) and oce- az-like corals are involved in Paleocene reef building anographic changes (Scott, 1995). Although coral (e.g., Bernecker and Weidlich, 1990) but played no reefs remain rare for most of the Paleogene in terms of such role in the latest Cretaceous, and secondly, the absolute recorded numbers (Kiessling, 2002), their fewer Paleocene z-like taxa were relatively more globally preserved volume peaked in the Paleocene, common in reefal associations than the late Campa- dropped dramatically in the earliest Eocene and then nian–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. 202 W. Kiessling, R.C. Baron-Szabo / Palaeogeography, Palaeoclimatology, Palaeoecology 214 (2004) 195–223 probably related to the evolutionary radiation of only event without a significant drop in reefal encrusting algae (Steneck, 1983; Aguirre et al., carbonate production at the stage level (Flu¨gel and 2000) rather than to greater reef-building potential Kiessling, 2002b). This could either mean that coral of the corals themselves. Crustose coralline algae are reefs were little affected by the extinctions, or coral very important for reef cementation and ease the larval reefs recovered more rapidly than at other extinction settlement of benthic organisms (Fabricius and intervals. Although data are too limited to provide a De’ath, 2001). Nearly all well-known Paleocene reef final judgment, the observation that the first known occurrences are indeed characterized by rich flora of Paleocene shallow-water coral reefs are of mid Danian coralline algae (Babic et al., 1976; Babic and Zupanic, (P1b) age (Tragelehn, 1996; Montenat et al., 2002; 1981; Bryan, 1991; Schuster, 1996; Tragelehn, 1996; Baron-Szabo et al., 2004) points to an actual gap in Montenat et al., 2002). their record but at the same time suggests a rapid Among the Big Five Phanerozoic mass extinctions, recovery. Kiessling and Claeys (2001) observed a the K–T boundary is unusual in that it represents the 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) 195–223 203 coral-algal reefs are known from the Danian of that probably does not hold true for mass extinction Europe, North Africa, and South America, the oldest intervals as will be discussed below. Paleocene coral reefs in the Gulf Coast area are of Standing diversity (Fig. 4A) of corals exhibits a Thanetian (late Paleocene) age. Their hypothesis that discontinuous rise towards the Recent, a trend usually enhanced ecosystem devastation in the Gulf of observed in large compilations of taxon-ranges Mexico region due to the proximity of the Chicxulub (Benton, 1995; Jablonski et al., 2003). Interesting impact crater was responsible for this pattern is features of the coral diversity curve lie in the Late intriguing but remains untested. Jurassic (Oxfordian–Kimmeridgian) diversity peak, the Late Cretaceous (Turonian–Maastrichtian) diver- 4.2. Diversity dynamics of Mesozoic–Cenozoic sity plateau, and in the slow rise of generic richness scleractinians during most of the Paleogene (Danian–Rupelian). The latter two observations are relevant to this paper and Long-term diversity dynamics of scleractinian indicate (1) that the scarcity of Late Cretaceous Coral corals were analyzed using the compilation of Reefs was not mirrored by a depression of coral Sepkoski (2002) at a stage-level stratigraphic reso- diversity and (2) the end-Cretaceous rudist extinction lution. The analysis was limited to taxa known from was not directly related to coral diversity dynamics. 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, 5. Overview of Coral-Bearing K–T regions which will be discussed below, problems with stratigraphic ranges are significant, not only for 5.1. Antarctica, New Zealand and South America Sepkoski’s compendium but also for stratigraphic ranges of corals in general (Baron-Szabo, 2002). Maastrichtian and Paleocene coral faunas from the Imprecise age assignments of coral occurrences are James Ross Basin of Antarctica are well known inherent to their habitat preferences (shallow water (Filkorn, 1994; Stolarski, 1996). The detailed work of with few reliable stratigraphic markers) and thereby Filkorn (1994) has revealed a surprising biodiversity obscure the inferred evolutionary dynamics. Of the in the Maastrichtian as well as the Paleocene. The 871 scleractinian genera listed in Sepkoski’s compi- faunas are predominantly solitary and exclusively az- lation, 405 have imprecise assignments of first or last like. One possible exception is Cladocora gracilis occurrences. We have excluded all those genera from (d’Orbigny) (=Cladocora antarctica Filkorn). the analysis being aware that this method tends to Low diversity coral faunas have also been recorded underestimate standing diversity (the number of from New Zealand (Squires, 1958; Stilwell, 1997). K– genera inferred to have co-existed in a time interval) T corals are poorly explored in South America, and exaggerates extinction and origination rates. although they are relatively common in the Roca Similarly, genera that are only reported from one Formation of Argentina. Our preliminary observations stage (singletons) were also excluded from the indicate moderate diversities with about equal pro- analysis, which tends to reduce both standing portions of az- and z-like corals in northern Patagonia, diversity and turnover rates. Although singletons with a trend towards az-dominance towards the south. may yield a biological signal, it is more likely that Corals were apparently more common in the Danian they are mostly due to limited research or preserva- than in the Maastrichtian. The recent discovery of a tional problems (Foote, 1997, 2000). The resulting Danian coral reef in La Pampa, Argentina (Baron- graph is based on only 335 coral genera and one Szabo et al., 2004) and ongoing field work in the area should note that confidence intervals, although not will provide additional data in the near future. indicated, are quite large. Values were normalized by the duration of the stage (time scale of Golonka and 5.2. Caribbean and North America Kiessling, 2002), which makes the implicit assump- tion that taxonomic turnover was continuous Rich coral-rudist associations of Jamaica have long throughout the stratigraphic interval, an assumption attracted paleontologists focusing on the K–T extinc- 204 W. Kiessling, R.C. Baron-Szabo / Palaeogeography, Palaeoclimatology, Palaeoecology 214 (2004) 195–223 tions (Coates, 1977). Maastrichtian corals occur in a associations are dominated by az-like corals. Rich mixed volcaniclastic-carbonate platform (Mitchell, Campanian–Maastrichtian faunas are known from 2002). The richest coral collections stem from the Madagascar (Alloiteau, 1958) and consist of about Titanosarcolites limestone, which is now dated as late equal proportions of az-like and z-like corals. A Maastrichtian in most parts (Steuber et al., 2002). The wealth of new data has allowed for a revision of taxonomic work by one of the authors (B.-S., work in Alloiteau’s stratigraphic assignments (Bignot et al., progress) on the large collections of Coates, Kauffman 1998; Rogers et al., 2000; Abramovich et al., 2003). and Jackson (1966–1972) has unraveled what is by far the greatest Maastrichtian coral diversity in the 5.4. Alpine belt and southern Europe western Hemisphere. A recent review of Mitchell (2002) has provided details on vertical patterns of Maastrichtian carbonate platforms, rich in corals coral occurrences. Other K–T corals have been and rudists, cover much of the southern European described from the Danian of Puerto Rico (Berryhill shallow shelves but taxonomic data on well-dated et al., 1960) and the Late Cretaceous of Cuba (Wells, localities are scarce (Polsˇak, 1985; Parente, 1994). 1941). Although apparently rarer, data on Paleocene coral Maastrichtian and Paleocene coral faunas are well localities are much better and have revealed a known from the Gulf Coast region (USA) but contain surprising diversity (Drobne et al., 1988; Moussavian almost exclusively az-like corals. Older studies and Vecsei, 1995; Vecsei and Moussavian, 1997; describe a fairly high diversity (Stephenson, 1917; Turnsˇek and Drobne, 1998). Vaughan, 1920; Wells, 1933), but according to our In the Alps, too, the Paleocene record of corals is revisions the corals were excessively oversplit. much richer than the Maastrichtian record. Reefal Maastrichtian assemblages from Mexico contain more carbonates with corals and algae are known from the z-like corals with a moderate to high species-richness bmiddleQ Danian onwards (Lein, 1982; Tragelehn, (Myers, 1968; Filkorn, 2003; Schafhauser et al., 1996). Non-reefal Paleocene corals assemblages are 2003). Paleocene corals are obviously rare in the also widespread (Ku¨hn, 1930; Ku¨hn and Traub, region (Vaughan, 1900) and the oldest corals reported 1967). Paleocene coral-bearing reefal limestones are from reefal assemblages are of Thanetian age (Bryan, known from the Carpathians of Slovakia (Samuel et 1991; Stemann in Bryan et al., 1997). al., 1972). Unfortunately, very little taxonomic infor- mation on corals is available from all Alpine 5.3. Africa Paleocene reefs, but unpublished data of B.-S. (work in progress) could be used and the bmorphotypesQ The best-explored K–T coral faunas are known figured in Tragelehn (1996) could mostly be identified from Egypt (Quaas, 1902; Wanner, 1902; Hassan and to the species level. Salama, 1969; Schuster, 1996). Maastrichtian corals With few exceptions, the coral faunas from the were apparently rare, and exclusively az-like corals Pyrenees (northern Spain and southern France) con- have been described, which all passed the K–T tribute little to the K–T discussion. Nearly all faunas boundary (Quaas, 1902; Tantawy et al., 2001). Rich previously described as Maastrichtian are now dated Paleocene faunas consist of about equal proportions of as late Campanian or older (Arde`vol et al., 2000) and az-like and z-like corals, the latter forming coral reefs data of Paleocene corals are limited (Alloiteau and (Schuster, 1996). In Somalia, Maastrichtian and Tissier, 1958). Paleocene coral-bearing sediments are known, but only the late Paleocene–early Eocene Auradu Lime- 5.5. Northern Europe and Greenland stone is well described taxonomically (Gregory, 1900; Carbone et al., 1993). K–T corals from West Africa Coral faunas have been well studied in several are only known from older publications (Alloiteau, regions including Denmark, the Limburg region 1952a; Barta-Calmus, 1969). Taxonomic revisions (Belgium, southern Netherlands, westernmost Ger- were possible but the stratigraphic assignments could many), central France, northern Germany, eastern not be updated. Both Maastrichtian and Paleocene England, and Poland. In the whole region az-like W. Kiessling, R.C. Baron-Szabo / Palaeogeography, Palaeoclimatology, Palaeoecology 214 (2004) 195–223 205 corals prevail. However, the late Maastrichtian of the Paleocene corals from Arabia, although some algal Maastricht area (southern Limburg) contains a sur- reefs have been described (Ra´cz, 1979). prising number of z-like taxa (Leloux, 1999), as does K–T corals from India and Pakistan are only the Danian reef of Vigny (Meyer, 1987). For the known from old reports (Stoliczka, 1873; Duncan, Maastricht region, corals from the upper Meerssen 1880; Noetling, 1897). Although we could revise their Member (IVf-6) were considered to be Cretaceous taxonomy from the published material, the strati- and all corals from the Geulhem Member were graphic data were sometimes difficult to assign to included in the Danian. We have excluded data from modern chronostratigraphy. From the corals reported the topmost Meerssen bed IVf-7, which is Danian at by Stoliczka (1873), the recent lithostratigraphic the type locality but is not readily recognized in other correlations of Sundaram et al. (2001) were used to localities (Leloux, 1999). assign ages. All corals reported by Noetling (1897) The Vigny reef (Montenat et al., 2002) seems to be were treated as Maastrichtian. The Duncan (1880) a key locality for the evaluation of coral extinctions, faunas were stratigraphically partitioned according to because all corals in this reef were said to belong to the stratigraphic assignments of Eames (1968) and Cretaceous genera (Meyer, 1987). A number of genera Adams (1970). Himalayan coral data are scarce. were previously thought to have disappeared from the Faunal lists are available for undifferentiated Campa- fossil record at the end of the Cretaceous or even nian–Maastrichtian and Montian strata (Liao and Xia, earlier and representatives of those genera are often 1994; Lo¨ser and Liao, 2001). not reported from other Danian localities. Because only a few corals were sketched by Meyer and considering the often-incorrect identifications, we 6. Evaluation of global turnover rates were reluctant to utilize the taxonomic data as reported. We have only included figured taxa in our 6.1. General considerations database, where reliable identifications were possible from the drawings. In this section we assess our database to discuss the Maastrichtian and especially Paleocene corals are magnitude of extinction and compare our results to widespread in the Ukraine and the European part of published and unpublished analyses. Because we Russia (Kuzmicheva, 1987). Nearly all of them are az- focus on extinctions related to the K–T boundary, like, as are the corals in boundary sections of we emphasize the dataset with confirmed Maastrich- Denmark and southern Sweden (Floris, 1979). Fairly tian age (M). Last occurrences of taxa in the late rich az-coral assemblages are known from K–T Campanian and unresolved late Campanian to early sections in the Nuussuaq area of West Greenland Maastrichtian, however, have to be considered (Floris, 1972). With new biostratigraphic data avail- because corals are relatively rare in comparison to able (Nøhr-Hansen and Dam, 1997), these assemb- other invertebrate groups such as bivalves and gastro- lages could be reliably assigned stratigraphically. pods. This implies a strong Signor–Lipps effect (Signor and Lipps, 1982) meaning that the last 5.6. Arabia, Pakistan, India and Tibet reported occurrence of a taxon is unlikely to represent its evolutionary last occurrence. A coral genus that Several coral-bearing sections are known from the has last been sampled in the late Campanian may Oman Mountains but are best known from the border therefore well have existed until the K–T boundary or region of Oman and the United Arab Emirates even above. (Metwally, 1996; Baron-Szabo, 2000). A surprising Results on extinction rates differ considerably diversity of corals has been reported from these depending on how singletons are treated. Singletons, deposits, mostly dated as bmiddleQ to late Maastrich- defined as taxa occurring in only one stratigraphic tian (Smith et al., 1995). A moderately diverse late interval, are continuously discussed in the macro- Campanian to early Maastrichtian coral fauna has also evolutionary literature (e.g., Foote, 2000). In most been reported from Saudi Arabia (Abed and El- recent studies on long-term diversity dynamics, Asa’ad, 1981). We know of no descriptions of singletons are excluded from the analyses because 206 W. Kiessling, R.C. Baron-Szabo / Palaeogeography, Palaeoclimatology, Palaeoecology 214 (2004) 195–223 they have been shown to create artificial patterns (see is lower when only Maastrichtian (M) data are section on diversity dynamics). This is because considered (Table 1). Because the data without single- singletons are thought to be mostly due to mono- tons are more reliable, our best approximation of the graphic effects or exceptional preservation. However, generic extinction of corals at the K–T boundary is singletons may also carry an evolutionary signal between 29% (M data) and 34% (CM data). To better representing short-lived clades. Separating those two constrain the generic extinction intensity related to the hypotheses is difficult and has to be done on a case- K–T boundary, we have carried out a simple test of by-case basis. Here we define singletons as genera confidence intervals on reported stratigraphic ranges which were both apparently restricted to one strati- of the 11 non-singleton genera that are last reported graphic stage and have been described from just one from late Campanian or late Campanian–early Maas- region. This concept is less rigorous than the unique- trichtian localities. We have transferred the usual occurrence concept sometimes applied to microfossil metrics reported in meters of section (Marshall, assemblages (Buzas and Culver, 1998) but is more 1990, 1998) to chronostratigraphic ages and calculated confined than the exclusion of taxa known only from the stratigraphic confidence intervals of those genera one time interval regardless of geographic distribution based on the reported range in millions of years and the (Foote, 2000). number of stratigraphic horizons in which the genera Results on taxonomic turnover may also be were reported using the compilation of Baron-Szabo influenced by peculiar collections with little similarity (2002). The 50% confidence intervals of three of the to others. We have identified two regions in our ten genera thought to become extinct in the early database, which may bias some of the results: the Maastrichtian or earlier reach the K–T boundary Maastrichtian of the Antarctic Peninsula and the (Negoporites, Placohelia, Stephanosmilia). We have Danian of Vigny (France). While the potential bias included a random selection of two genera in the K–T of the Antarctic Peninsula is largely balanced by the victims category (according to Marshall’s method half exclusion of singletons (most of the unique genus of the taxa whose 50% error bars reach the boundary occurrences are also stratigraphic singletons), the should be included in the victims category). Thereby Vigny fauna poses larger problems. Although all we achieve 30F8% as our best approximation of end- problematic occurrences have been excluded from our Cretaceous scleractinian coral extinctions at the genus dataset, the fauna still contains two survivors (Bra- level. The results excluding data from Antarctica and chyphyllia and Synastrea), which have not been Vigny (ÀAV data) are slightly higher. Overall non- reported from other Paleocene localities. Therefore, singleton extinction rates are 31% for the M data and we have tested the sensitivity of our results to those 37% for the CM data. faunas by running all analyses with the complete Species-level extinction rates are usually not exclusion of Antarctica and Vigny (-AV data) and provided in global surveys owing to the strong noise report all noticeable differences. introduced by taxonomic dchauvinotypyT (Rosen, 1988). In spite of our taxonomic revisions, the 6.2. Extinction intensities apparent species level extinction rate of corals at the K–T boundary is 63F6% (M data). This value is still The approach most widely used to evaluate arguably high, perhaps due to sampling problems in extinction intensities is based on the evaluation of the Paleocene. With reverse rarefaction (Raup, 1979) genus ranges. The total generic extinction rate of corals we achieve 45F5% species extinctions. We used the is 39F8% when we simply count all late Campanian– number of species in our recorded Cretaceous coral Maastrichtian (CM) coral genera (including Lazarus genera as a basis for the taxon-size distribution, which genera) and extract those that have no record in the seems to be more reliable than the estimate based on Cenozoic (Table 1). This number is lower but not modern echinoids as done by Raup (1979). However, significantly different from the result based on reverse rarefaction may generally underestimate spe- Sepkoski’s compendium (Sepkoski, 2002), which cies loss (Jablonski, 1995) and thus the true species gives 43F9%. The difference remains approximately level extinction rate may have been somewhat higher the same when we exclude singletons. The extinction than 45%. W. Kiessling, R.C. Baron-Szabo / Palaeogeography, Palaeoclimatology, Palaeoecology 214 (2004) 195–223 207

Our family level extinction rate is slightly lower facies trends, such as Jamaica (Mitchell, 2002), than reported in earlier summaries (Barta-Calmus, Slovenia (Turnsˇek, 1994), and southern Spain (Go¨tz, 1984). Five out of 43 (12%) late Campanian– 2003), corals disappear even earlier than rudists. If, Maastrichtian coral families have no Cenozoic record like the rudists, scleractinian corals had experienced and only four out of 42 families (10%) from M complete extinction at the K–T boundary, a gradual localities (including ghost ranges) have no younger extinction would surely have been inferred for the record (Cladophylliidae, Isastreidae, Microsolenidae, corals. Pachyphylliidae). All these families had only one or two genera in the Maastrichtian. The one family with 6.4. Survivorship and abundance a reported last occurrence in the late Campanian, the monogeneric Negoporitidae, is generally rare in the Evaluations of extinction risks are based on both Cretaceous and only reported by one occurrence in the the late Campanian–Maastrichtian (CM) and the late Campanian of Spain (Go¨tz, 2003—described as Maastrichtian (M) occurrence matrices. The number Actinaraea sp.). of taxonomic occurrences was used as a rough proxy for abundance. Although this measure is crude in 6.3. Gradual or abrupt extinction? comparison to counts of specimens (not available for all collections), the close relationship between the two The current resolution of the data does not permit metrics (Buzas et al., 1982; Alroy, 2000; He and a conclusive assertion on the rapidity of the coral Gaston, 2000; unpublished data of W.K.) supports the extinctions. Nevertheless, our data allow some state- feasibility of this approach. ments about the sometimes quoted gradual decline of The assessment of relationships between survivor- corals prior to the K–T boundary (Moussavian, ship and the number of occurrences produces 1992). Rosen (in MacLeod et al., 1997) has already equivocal results. When Lazarus taxa are included noted that newer data tend to show that coral (zero occurrences in CM, but known from earlier extinctions have been more concentrated in the later and later intervals), we achieve a significant relation- Maastrichtian than previously realized. Our database ship between the probability of survival and confirms this: when all CM coral occurrences are abundance ( P=0.021; Mann–Whitney U-test) for analyzed, the total extinction of non-singleton coral the CM dataset. However, the M dataset produces genera is only 5.5% higher than for M corals. Even no significant dependency ( P=0.143). While the without considering stratigraphic confidence inter- latter observation is in line with recent analyses on vals, the late Campanian–early Maastrichtian back- bivalves saying that survivorship and abundance are ground extinction rate appears to be unusually low, decoupled at the K–T boundary (Lockwood, 2003), which agrees well with the low post-Cenomanian/ the former test implies that rare genera were indeed pre-Maastrichtian extinction rates from the analysis more prone to extinction than abundant genera. of Sepkoski’s dataset (Fig. 4B). From this and from Because the data excluding Antarctica and Vigny the long stratigraphic confidence intervals on the last (ÀAV data) confirm that there is no significant reported occurrence of coral genera we infer that difference in survival probabilities ( P=0.216 and coral extinctions may well have been concentrated in P=0.654, respectively), we follow Lockwood (2003) the late Maastrichtian. Because rich coral faunas with and conclude that abundance was not an insurance many victim genera are known even from latest against extinction. Maastrichtian localities, such as the Maastricht area Species-rich genera are not less likely to become (Leloux, 1999) and Jamaica (Mitchell, 2002 and extinct than non-singleton genera with few species, unpublished data of B.-S. with new stratigraphic data both in the CM ( P=0.109) and M ( P=0.167) data in Steuber et al., 2002), the extinctions may even (same results with ÀAV data). Previous analyses of have been concentrated at the K–T boundary itself. large species-level datasets have similarly resulted in The decline in sampling (reported coral occurren- no significant differences in extinction risk between ces), however, appears to be gradual in the late species-poor and species-rich genera (Jablonski, Maastrichtian. In several sections with shallowing 1986a; Smith and Jeffery, 1998). 208 W. Kiessling, R.C. Baron-Szabo / Palaeogeography, Palaeoclimatology, Palaeoecology 214 (2004) 195–223

6.5. Ecological selectivity of extinctions extinction risk was higher for corals with high corallite integration (meandroid, thamnasterioid) than Rosen and Turnsˇek (1989) made a strong case of for less integrated colonies (CM data, P=0.014; M selective extinctions of z-like corals at the K–T data, P=0.024; also significant for the ÀAV dataset). boundary and our data confirm their basic result. When, in addition to our four corallite integration The raw data (including the singletons), however, categories, we include solitary forms as having 0 although showing higher extinction rates of z-like corallite integration, the dependence becomes even corals in comparison to az-like taxa (Table 1) produce more significant ( Pb0.001). no significant differences. The same applies to the Larval strategy is another possible selection crite- analysis based on Sepkoski’s compendium. However, rion discussed in the literature. Previous analyses have significant differences emerge when the analysis is shown that no significant correlation between larval limited to non-singleton genera (Fig. 5; P=0.001 for ecology (e.g., planktotrophic versus lecitotrophic) and both CM and M; Mann–Whitney U). The difference is extinction risk exists at the K–T boundary (Valentine due the commonness of Maastrichtian az-like single- and Jablonski, 1986; Smith and Jeffery, 1998). Too ton genera, which are mostly concentrated in the little is known about larval ecology of ancient sclerac- James Ross Basin of Antarctica (Filkorn, 1994). tinians to test this hypothesis. From the few genera (13) When we filter out the AV data, significant differences that we can assign to larval feeding mode (all survivors, are even achieved if singletons are included. The assignment based on Edinger and Risk, 1995), nearly result supports the view that the end-Cretaceous equal proportions are brooders (lecitotrophic larvae) or extinction was selective against photosymbiosis or broadcasters (planktotrophic larvae). This suggests that some ecological attribute associated with photosym- larval feeding mode did not act as a selective criterion biosis (Rosen and Turnsˇek, 1989). at the K–T boundary. The difference of non-singleton extinction rates is The mean skeleton size of solitary coral genera had also significant between colonial and solitary genera. no significant influence on survivorship ( P=0.69) and However, because coloniality and ecological mode are the mean size of Maastrichtian and Paleocene genera closely linked in corals (in our dataset 84% of az-like was nearly identical. This suggests that the extinction taxa are solitary and 95% of z-like taxa are colonial) it did not select against large-sized genera. However, this is difficult to separate both factors. For colonial corals 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 demon- strates the importance of keeping dz-coralsT and dreef coralsT as separate concepts (Rosen, 2000).

6.6. Taxonomic structure of extinctions

Extinctions are unequally distributed among higher coral taxa (Fig. 6). Non-singleton generic extinctions Fig. 5. Ecological selectivity of coral extinctions at the K–T boundary. Extinctions rates depend strongly on feeding mode with vary significantly between coral suborders ( P=0.002; presumably zooxanthellae bearing corals (z-like) being much more Kruskal-Wallis H). Although the differences are affected than azooxanthellate corals (az-like). 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 Dendro- boundary became extinct in the Paleocene; 12 have phylliina with unusually low extinction rates are their last occurrence in the Danian. Thus the Danian exclusively az-like), extinction intensity also varies has 13% DCWs at the genus level. This is a significantly between suborders with mostly z-like conservative estimate, because most taxa of the genera (e.g., Astrocoeniina and Microsolenina). This apparently DCW-rich coral fauna from Vigny (Meyer, corroborates former results indicating that extinctions 1987) were not included in our database (see Section are correlated with taxonomic groups (Smith and 5.5). Removing the two remaining DCW genera only Jeffery, 1998). known from Vigny still makes 11% DCW s. str., a value significantly higher than the 3% recorded in 6.7. Dead Clade Walking (DCW) Sepkoski’s compendium. Our data do not allow testing hypotheses of the The concept of clades surviving mass extinctions cause of the DCW phenomenon. However, the ecology but becoming extinct soon after has long been of of coral DCWs gives some indication. Although the interest to the scientific community, but only recently number of z-like genera in the Danian is nearly equal a statistical test was applied, which verified that this is to the number of az-like genera (59/57) the dichotomy a general phenomenon after mass extinction events in the ecological traits of DCW corals is as great as for (Jablonski, 2002). The hypothesis of Dead Clade the K–T extinction. Z-like DCW genera are more Walking (DCW) predicts that significantly more common than az-like ones (9 and 3 genera, respec- clades become extinct in the stage after a mass tively). This suggests that either a similar ecological extinction than during background extinction times. selectivity has acted on the survival fauna as is evident Our coral data indicate that the DCW phenomenon for the end-Cretaceous extinction, or that stochastic is more profound for K–T corals than previously processes preferentially extinguished the more dam- recognized. Nineteen coral genera surviving the K–T aged z-like corals in the Danian (bottleneck effect). 210 W. Kiessling, R.C. Baron-Szabo / Palaeogeography, Palaeoclimatology, Palaeoecology 214 (2004) 195–223

6.8. Changes in standing diversity 6.9. Appreciation of biases

Due to delayed recovery, the K–T boundary is Recent analyses of large datasets suggested that not only characterized by significantly elevated most if not all mass extinction events recognized in extinction rates but also by a pronounced decline the fossil record are exaggerated and can largely be in standing diversity. Both the biodiversity based on explained by heterogeneities of the rock record (Peters through-ranging genera and the raw pattern of and Foote, 2002). Although there are pitfalls in the recorded species and genera indicate that corals approach of Peters and Foote (2002) (Foote, 2003; exhibit a marked decline of standing diversity. Crampton et al., 2003), the issue has to be discussed. However, these raw data have to be normalized The difference in the number of sampled corals for the strong differences between the Maastrichtian between the Maastrichtian and Paleocene is indeed and Danian fossil record. Because advanced resam- profound. When recorded and inferred standing pling methods for analyses based on through generic diversities are compared, the simple com- ranging-taxa (Alroy, 2000) are not applicable (not pleteness measure, which represents the ratio of taxa sufficient temporal bins available), we used simple actually recorded in an interval to taxa inferred to rarefaction to account for differences in sampling. have been present (Fara, 2001), declines from 94% in The result indicates that late Maastrichtian coral the late Campanian–Maastrichtian (CM), 88% in the diversity was indeed significantly higher than early/ Maastrichtian (M), 86% in the whole Paleocene (Pa) middle Danian diversity (Fig. 7). However, this to 71% in the Danian (D). This decline in the quality difference is not evident when whole stages are of the fossil record results in a severe Lazarus effect compared (Maastrichtian and Danian). If the same (Jablonski, 1986b). Many surviving genera are limited number of taxonomic occurrences had been recorded in CM but not in Pa (15 out of 94), six recorded from the Maastrichtian as is present in the survivors are known from Pa but not from CM and Danian, the number of recorded genera is likely to three are neither recorded in CM nor in Pa. When the be nearly identical. Thus the drop of standing extinction rate is calculated without considering these diversity is unlikely to have persisted for an Lazarus taxa, the non-singleton genus extinction rate extended period after the K–T boundary, which is rises to 48F8% (CM). However, when the percentage in contrast to previous analyses (Rosen and Turnsˇek, of Paleocene Lazarus genera is subtracted from the 1989). 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) 195–223 211 tion that the same percentage of genera is not recorded Table 2 in the Paleocene but did survive the K–T boundary Origination metrics of corals in the Paleocene event), the end-Cretaceous extinction rate would drop Danian KTbase Error N Sepkoskia Error N to 29F7% (CM) or 24F7% (M). This is still Total (with 21.0 7.3 119 23.0 8.8 87 significantly above the mean extinction rate for singletons) scleractinian coral genera (12F4% per stage calcu- z-like (with 9.8 7.5 61 3.1 6.0 32 singletons) lated from Sepkoski, 2002) and suggests that the K–T az-like (with 32.8 12.1 58 34.5 12.6 55 boundary represents a true mass extinction event for singletons) scleractinian corals. Total (without 20.3 7.3 118 18.3 8.4 82 singletons) 6.10. Origination rates z-like (without 9.8 7.5 61 3.1 6.0 32 singletons) az-like (without 31.6 12.1 57 28.0 12.4 50 The evolution of new genera within the Paleocene singletons) was apparently gradual, but concentrated in the Danian where 21% of the genera are new. The overall Total Paleocene KTbase Error N Sepkoskia Error N origination rates derived from our database are almost Total (with 27.7 7.7 130 35.6 9.2 104 identical to those derived from Sepkoski’s compen- singletons) dium (Sepkoski, 2002). A significant difference, z-like (with 20.3 9.5 69 6.1 8.1 33 singletons) however, is noted for the origination rate of z-like az-like (with 36.1 12.1 61 49.3 11.6 71 corals (Table 2). For the whole Paleocene we note singletons) 20F9% newly evolved z-like genera, while only Total (without 26.6 7.7 128 25.6 9.0 90 3F6% (0–9%) new genera are derived from the singletons) compendium. Similar to the compendium data, our z-like (without 20.3 9.5 69 3.1 6.0 32 singletons) recorded origination rates in the Danian are signifi- az-like (without 33.9 12.1 59 37.9 12.5 58 cantly higher for az-like genera than for z-like genera. singletons) However, the difference is not significant when the All values except N in percent. Errors represent 95% confidence whole Paleocene is considered. intervals. Another important observation is the low number a Analyses based on data in Sepkoski (2002). of singleton taxa both in the Danian alone and the entire Paleocene. The one Danian stage singleton than the extinctions. Origination rates in the Danian (Faksephyllia) is not even a singleton according to were apparently independent of coloniality and newly our definition above, because it is recorded from a evolved genera occurred as often as surviving genera number of regions (Greenland, Denmark, Austria, and the solitary newcomers were not significantly Kazachstan). This contrasts to the eleven Maastrich- different in mean size from survivors. tian singleton genera (Table 1). Thus there is no evidence for an expansion of short-lived clades in 6.11. Comparison with other databases the Danian. The origination of new Paleocene clades was less As reported above, our extinction and recovery heterogeneous among coral suborders than in the data are often similar with data derived from Sepkoski extinction, and the overall dependency is not signifi- (2002). The overlap of confidence intervals between cant ( P=0.172; Kruskal Wallis test). The Astrocoe- analyses based on our and Sepkoski’s compilation is niina and Caryophylliina have the relatively strongest remarkable because most of the data in Sepkoski’s enrichment of new genera, while these groups had compendium can be shown to be incorrect. The most suffered unusually low extinctions at the end of the important source of error is introduced by incorrect Cretaceous (Fig. 6). Apart from the significant stratigraphic ranges (Fig. 8). Subjective synonyms, dependence of Danian originations from inferred although relatively common (second most common ecological mode (az-like versus z-like; P=0.003), after unrealized boundary genera), have a limited originations seem to have been even more random 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 compen- and a false assignment of first and/or last occurrences. dium. Maastrichtian standing diversity is mostly However, our data show a systematic tendency affected by the number of genera that were not towards lower extinction rates, a higher number of considered as boundary genera (occurrence in, or DCWs and a greater proportion of newly evolved z- range through Maastrichtian–Paleocene) in Sepkoski like genera as compared to Sepkoski’s compendium. (2002). However, the net effect on extinction rates is Although we agree with Adrain and Westrop (2000) low because the new boundary genera are approx- that the basic evolutionary patterns can be correctly imately equally partitioned between victims and derived from analyses of Sepkoski’s database, a survivors. The net effect on extinction rates is largest critical review is clearly necessary when analyzing for the stratigraphic range errors in Sepkoski’s mass extinctions. compendium. Especially the number of previously There is the great difference in extinction rates unrealized boundary genera affects calculated turn- between our database and the analysis of Rosen and over rates. The same is true for originations in the Turnsˇek (1989), who have indicated a 60% extinction Paleocene. While the number of wrong originations is at the genus level and 97% at the species level. almost exactly matched by the number of new Although we could confirm that non-singleton z-like originations from erroneous boundary crossers, 18 corals where significantly more affected than az-like new Paleocene originations (mostly z-like) are recog- corals, the dichotomy is manifested at 44% versus nized from a downward extension of first occurrences. 13% (CM data) rather than the 70% versus 40% given Another recent detailed comparison of diversity by Rosen and Turnsˇek. The great difference in results dynamics extracted from Sepkoski’s compendium is probably (1) due to the unrevised taxonomic dataset (trilobites) and those compiled from taxonomic ex- Rosen and Turnsˇek have used and (2) a result of the perts, has resulted in nearly identical patterns (Adrain large stratigraphic intervals they have considered and Westrop, 2000). Similar to our results Adrain and (Late Cretaceous and Paleocene). It is difficult to Westrop (2000) noted that more than 50% of data in judge which bias is greater. We see, however, that Sepkoski (2002) are wrong due to taxonomic noise taxonomic revisions are very important when evaluat- W. Kiessling, R.C. Baron-Szabo / Palaeogeography, Palaeoclimatology, Palaeoecology 214 (2004) 195–223 213 ing mass extinctions or diversity dynamics in general region size to the number of occurrences. We discuss (see also Jablonski et al., 2003). analyses based on intermediate (19 coral-bearing regions), coarse (10 regions) and very coarse (4 regions) geographic scales. The analysis based on an 7. Geographic patterns 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 Geographic extinction patterns of the late Campa- scales. Compared to similar studies using a 108 grid nian–Maastrichtian (CM) dataset agree well with those (Raup and Jablonski, 1993), our scale had to be coarser of the more restricted Maastrichtian dataset. To max- because coral data are much scarcer. We conducted imize sample size and statistical confidence we have analyses of extinction and recovery rates on variable therefore used the larger CM dataset for the discussion. geographic scales from basin to hemisphere, applying Extinction intensities are apparently not randomly both objective (grids) and subjective (biogeographic distributed geographically when stratigraphic single- regions) definitions of areas. While objective criteria tons are excluded. The highest extinction rates are for defining regions have the advantage of permitting a recorded in low paleolatitudes, and drop off towards neutral approach to geographic patterns, subjective higher palaeolatitudes. The pattern is best visible on criteria can provide a more natural definition of manually defined regions at the intermediate geo- geographic regions. Moreover, the filter of z10 genera graphic resolution (Fig. 9) but the same basic results per region is the more severe, the smaller the regions were achieved with other definitions of regions. are defined. This can partly be balanced by adjusting 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 this analysis, however, that North America was not a significant differences even at the coarsest geograph- hot spot in end-Cretaceous coral extinctions. ical scales. The general tendency towards lower ex- tinction rates in higher latitudes is likely to be an artifact 7.3. Extinction risk and geographic distribution of latitudinal variations in the composition of regional assemblages. Limiting the analysis to z-like corals Geographic ranges of CM corals were measured by produces no latitudinal cline in extinction rates (Fig. the number of geographic regions in which coral 10), nor any significant differences between geogra- genera are recorded. We have tested the relationship phic regions. This suggests that all patterns are strongly between survivorship and geographic distribution on controlled by the relative contribution of the extinction- different geographic scales. As in the previous resistant az-like corals to regional assemblages and analyses, stratigraphic singletons were excluded. latitude had no direct influence of extinction rates. There is a clear relationship between extinction Similar to previous analyses on bivalves (Raup and risk and geographic distribution, which is nearly Jablonski, 1993), we do not detect any hot spots in the independent of how the regions are defined. extinctions. Although the Mediterranean Tethys con- Survivorship is most clearly linked to geographic stantly exhibits slightly elevated extinctions in com- distribution in the 308 binning analysis ( P=0.002; parison to other regions, both for whole faunas and for Mann–Whitney U) but the relationship is also only z-like corals, the differences are small and far significant at the intermediate ( P=0.004) and from being statistically significant. Even on very large coarse geographic scales ( P=0.009). Even when geographic scales, non-singleton CM extinction rates including survivors without reported occurrences in are similar and do not deviate markedly from the the late Campanian–Maastrichtian (Lazarus genera, global mean: North America, 33F11%; Europe, number of regions=0), the relationship remains 35F10%; Africa and India, 30F11%. The same significant. These results strengthen previous anal- applies for endemics to regions. Even the raw data yses, which conclude that wide geographic distri- indicate little variation in extinction risk of endemics butions of clades enhance the probability of (North Africa, 50%; Europe, 50%; North America, survivorship at the K–T boundary (Jablonski and 40%) and statistical errors are so large that no Raup, 1995) and other mass extinction intervals conclusive statement is possible. It is evident from (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 because reported generic richness is low for all regions except Europe (large binomial errors) and the Although the locus of evolution of new genera in Paleocene coral record of Northern Europe is the Paleocene is unlikely to be captured by our basically limited to the Danian. However, the database approach, the distribution on new genera in apparent concentration of new Paleocene z-like regional assemblages may give some insight into genera in low latitudes (Fig. 12), may represent a evolutionary dynamics. Due to the Danian Lazarus biological signal, even though the latitudinal gradient effect (29% of all coral genera, inferred to be is not significant. Our data do not allow testing for present, are not recorded), the regional assemblages the concentration of invaders and bloom-taxa, which are generally richer in new genera than the global have been noted to be exceptional in Gulf Coast average (Table 2) and thus have to be interpreted bivalve faunas (Jablonski, 1998). cautiously. New genera in the Danian were nearly The previous observation of an increase of uniformly distributed (Fig. 11A). However, averaged cosmopolitanism of corals in the Paleocene (Rosen over the whole Paleocene, the percentage of Paleo- and Turnsˇek, 1989) cannot be confirmed by our data. cene genera relative to Cretaceous survivors is The mean number of geographic regions in which apparently higher in low latitudes (Fig. 11B), coral genera are recorded does not differ significantly although the maximum difference is noted between between the Maastrichtian and the Danian, nor does it North Africa (region 8, 35F14% new genera of total differ between the late Campanian–Maastrichtian and faunas) and the Gulf of Mexico (region 6, 19F15%), the Paleocene. which were both in the tropics in the Paleocene. Danian DCW genera are only recorded in northern Latitudinal differences become greater when only z- Europe (7 out of 33 surviving genera are DCWs), like corals are analyzed. New z-like corals in the southern Europe (5/30), North Africa (3/18) and India Paleocene are much more common in North Africa (2/4). Other regions with a moderate or good Danian (45F22%) and Pakistan (40F21%) than in Northern record such as the Gulf Coast/Caribbean and South Europe (8F14%), while the Mediterranean Tethys America do not show any DCWs. Thus DCWs seem (14F13%) and the Gulf of Mexico region (14F18%) to be concentrated in Eurasia and Africa. This implies have intermediate concentrations of new z-like that the DCW phenomenon was indeed influenced by genera. This statement must also be qualified, 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) 195–223 217 simple function of paleolatitude. Another noteworthy zooxanthellate symbionts in their tissues (z-like feature is the fact that the majority of Danian DCWs corals) are much stronger affected than corals inferred are just recorded from one region (7) and no DCW is to have lacked these symbionts (az-like corals). This recorded from more than two regions. This confined difference, previously noted by Rosen and Turnsˇek distribution of DCW genera substantiates the inter- (1989) on a geographically confined dataset, is now pretation that the DCW phenomenon is probably confirmed at global scales. However, it is not justified caused by bottleneck effects (Jablonski, 2002). 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 8. Discussion and conclusions severe as previously thought and coral reefs, often dominated by z-like corals, are more common than in The currently best approximation of scleractinian the Maastrichtian. A global z-like coral reef gap, if it coral extinction rates at the K–T boundary is 30F8% existed at all, is confined to the earliest Danian. at the genus level. This is among the lowest extinction An additional ecological link to extinction risk is rates thus far reported from benthic marine inverte- indicated by morphological complexity, as measured brates at the K–T boundary. Even the echinoids, by coloniality and colony integration. Although we although as thoroughly revised taxonomically as our can hardly envision a mechanism how a mass dataset (Smith and Jeffery, 2000) still exhibit a generic extinction could select against coloniality in itself, extinction rate of 36% (Smith and Jeffery, 1998). The chances are that complexity (see McShea, 1996, for low extinction rate recorded for corals is in line with definitions of complexity) played a role in the recent interpretations that the magnitude of mass selective extinction as has been previously suggested extinction events is probably strongly exaggerated (Rosen and Turnsˇek, 1989). The higher the colony (Peters and Foote, 2002). However, balancing for integration the greater was the chance of extinction at Lazarus effects caused by variations in the quality of the K–T boundary. There is some evidence that corals the fossil record, verifies that the K–T boundary with a high integration of corallites also have a high represented a true mass extinction event for scleracti- physiological integration (Rosen, 1986; Coates and nian corals. The reason for the lower extinction rate in Jackson, 1987). Colony integration could also be comparison to previous analyses is only partially due related to physiological dependency on photosymbio- to the improving fossil record in the Paleocene: sis, but current data are too limited for a conclusive several genera previously thought to become extinct statement. A quantitative test of modern corals based at the K–T boundary (e.g., Calamophylliopsis, Phyl- on the compilation of Stimson et al. (2002) did not locoenia, Synastrea), have now been recorded from produce significant relationships between corallite the Danian and changed from the victims into the integration and zooxanthellate densities, which are Dead Clade Walking category (Jablonski, 2002), that thought to be related to survival probability during is, they survived the K–T boundary but became bleaching events (Stimson et al., 2002). Zooxanthellae extinct soon after, probably due to bottleneck effects. densities are usually low in meandroid as well as The greatest effect on extinction rates is given by the solitary forms and have a maximum in plocoid and specimen-based taxonomic revisions. While the net cerioid forms. This might indicate that there was effect of the improving Paleogene fossil record is indeed a general selection against physiological nearly balanced by the also improving Maastrichtian complexity in the end-Cretaceous extinctions. record (with new victims discovered, which were Widespread geographic distribution in the Maas- previously thought to have become extinct before), trichtian formed an insurance against extinction at the the specimen-based comparisons of Cretaceous and K–T boundary. Apart from this observation, geo- Cenozoic corals (B.-S.) have extended many strati- graphic patterns of extinction and recovery are graphical ranges across the K–T boundary. indistinct, perhaps owing to the limited sample sizes. The most significant dependency of extinction Extinction rates are slightly elevated in low paleo- intensities in scleractinian controls is seen in their latitudes, but this is exclusively due to the variable ecological mode. Corals inferred to have hosted 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 dependencies of extinction risk and we follow pattern of z-like corals and endemic clades underlines Jablonski (1986a) that the main reason may lie in the idea that the end-Cretaceous extinctions were macroevolutionary particularities of mass extinctions. caused by a global catastrophe rather than a combi- However, feeding strategy clearly has a major nation of regional factors. Patterns of recovery are selective effect in the extinction of corals and somewhat more variable geographically, especially apparently in the other groups as well. While in for z-like corals. New Paleocene z-like clades were corals simple zooplankton feeding seems to have been proportionally more common in low latitudes than in more advantageous to survival than a combination of higher latitudes. If the regional proportions of new photosymbiosis and zooplankton feeding; deposit genera have some relationship to geographic varia- feeding was apparently favored in bivalves (Rhodes tions in evolutionary rates, we could conclude that the and Thayer, 1991; but see Jablonski and Raup, 1995 tropics represented a source of evolutionary novelty in for alternative explanations); and omnivory or selec- the post-extinction phase. The pattern of the tropics as tive detritivory enhanced survivorship in echinoids a source of evolutionary novelty has also been (Smith and Jeffery, 1998; Jeffery, 2001). Similarly, observed during background extinction times (Jablon- extinctions were heterogeneously distributed among ski, 1993). Because the latitudinal cline is not higher taxonomic levels in all three groups, but statistically significant, any far-reaching conclusion species richness in clades did not influence their has to remain open until more data become available, survivorship. Previous studies also agree that abun- especially from the Southern Hemisphere. dance per se had no significant effect on extinction In spite of conjectured deviations in the mode of risk (Smith and Jeffery, 1998; Lockwood, 2003). The coral evolution from the normal bDarwinianQ pattern results are less uniform regarding geographic distri- (Veron, 1995), there are striking similarities with other bution. Our results agree with Raup and Jablonski benthic invertebrates in the macroevolutionary (1993) in that survivorship was more likely when response at the K–T boundary (Table 3). Because clades where widely distributed geographically and now three major groups of benthic invertebrates there was no geographic hotspot of extinctions. Smith (bivalves, echinoids, corals) have been rigorously and Jeffery (1998), however, note the opposite in both analyzed at a global scale, we can start to extract cases, identifying North America as the region with general rules for the K–T boundary. In line with the highest extinction risk. While this would be in line previous analyses we have found few significant with an impact scenario as a cause for the extinction (enhanced effects proximal to the Chicxulub site), we Table 3 argue that the echinoid data alone are too weak to Comparison of generic survivorship dependencies valid for different confirm this. groups of benthic invertebrates Although our study provides important results on Trait Coralsa Bivalvesb Echinoidsc macroevolutionary processes around the K–T boun- Abundance no? no no dary, we have no direct hint on the cause(s) of the Feeding strategy yes no? yes mass extinction. Our data show no evidence for long Coloniality yes N.A. N.A. term-climate change as a dominant trigger of the Larval strategy no? no no extinction, because extinctions of z-like corals, which Infaunal vs. epifaunal N.A. no no are very sensitive to climate change today, have no Habitat no no no Size of solitary genera no no ? relationship with paleolatitude. Global cooling would Taxonomic structure yes yes? yes predict enhanced extinctions in high latitudes, Number of species in genus no no no whereas global warming should have led to stronger Geographic range yes yes no extinctions in the tropics. Ecological selectivity of the Endemic to America no no yes extinction is compatible with incident light reduction N.A.—not applicable. (dust and sulfate aerosols) caused by a bolide impact, a Scleractinians only (this study). b Raup and Jablonski (1993), Jablonski and Raup (1995), but several other processes could also explain the Lockwood (2003). pattern. 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