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TREATISE ONLINE Number 29

Part N, Revised, Volume 1, Chapter 24:

Extinction in the Marine

Paul G. Harnik and Rowan Lockwood 2011

Lawrence, Kansas, USA ISSN 2153-4012 (online) paleo.ku.edu/treatiseonline

PART N, REVISED, VOLUME 1, CHAPTER 24: IN THE MARINE BIVALVIA Paul G. Harnik and Rowan Lockwood [Department of Geological and Environmental Sciences, , [email protected]; and Department of , The College of William and Mary, [email protected]] Bivalves are diverse and abundant constit- particularly suitable for investigating diver- uents of modern marine , and they sity dynamics over geologic time. We then have a rich record that spans the introduce recently developed analytical (Hallam & Miller, 1988; methods for estimating rates of extinction McRoberts, 2001; Fraiser & Bottjer, and origination from paleontological data 2007). Due to the high quality of their that account for temporal variation in the fossil record, they are well suited for inves- quality of the preserved and sampled fossil tigating patterns of change and record. Applying these methods to data for the processes that generate these patterns. marine bivalves, we present a new analysis Extinction and its influence on patterns of of extinction, origination, and preservation diversification in the Bivalvia have figured rates for bivalve genera over the Phanerozoic prominently in a number of previous studies. and examine the effect of extinction rate on Did bivalves diversify exponentially over subsequent origination rate. We review the geologic time with little long-term influ- growing literature on extinction risk in fossil ence of mass extinction events (Stanley, marine bivalves and summarize the roles of 1975, 1977), or were catastrophic extinction several biological factors that have proven events critical in shaping their dramatic post- important in predicting survivorship over radiation (Gould & Calloway, geologic time. Although recent and historical 1980)? Extinction is an important process extinction in freshwater mussels has been in the evolutionary history of many . well studied (Williams & others, 1993; Selective or chance survivorship can shape Ricciardi & Rasmussen, 1999; Lydeard morphological, ecological, and phylogenetic & others, 2004; Strayer & others, 2004; diversity and disparity (Roy & Foote, 1997; Bogan, 2006), we focus on marine bivalves Jablonski, 2005; Erwin, 2008). Extinction due to the quality of their fossil record over selectivity can also affect the susceptibility long time scales and the general congru- of lineages to later periods of environmental ence between phylogenetic hypotheses and change (Stanley, 1990a; Jackson, 1995; morphologic taxonomies (Jablonski & Jablonski, 2001; Roy, Hunt, & Jablonski, Finarelli, 2009). 2009). In addition, extinction can open up opportunities for diversification through MARINE BIVALVES AS the removal of incumbent taxa (Walker & A MODEL FOR Valentine, 1984; Rosenzweig & McCord, ECOLOGICAL AND 1991; Bambach, Knoll, & Sepkoski, 2002; EVOLUTIONARY ANALYSIS Jablonski, 2008b). Understanding how and why some organisms, including many The marine bivalve fossil record has been bivalves, and not others became extinct in studied intensively by paleontologists and the past may prove useful in predicting the malacologists for centuries. This body of response of modern marine ecosystems to work has produced a detailed picture of the environmental change (Dietl & Flessa, history of the and has advanced our 2009). general understanding of the processes that Here we briefly review those features generate and maintain biodiversity in marine of the bivalve fossil record that make it systems over time. Data for fossil bivalves

© 2011, The University of Kansas, Paleontological Institute, ISSN (online) 2153-4012 Harnik, Paul G., & Rowan Lockwood. 2011. Part N, Revised, Volume 1, Chapter 24: Extinction in the marine Bivalvia. Treatise Online 29:1–24, 4 fig., 1 table, 1 appendix. 2 Treatise Online, number 29 have been instrumental in informing debates the living communities from which they concerning the roles of biological factors in are derived (Kidwell, 2001, 2002, 2005; extinction risk (e.g., Stanley, 1986a; Raup Lockwood & Chastant, 2006; Valen- & Jablonski, 1993; Jablonski, 2005; Rivad- tine & others, 2006). Notably, instances eneira & Marquet, 2007; Crampton & in which the ecological agreement between others, 2010), the tempo and mode of evolu- and assemblages is poor tend to tionary change (e.g., Kelley, 1983; Geary, be associated with sites affected by recent 1987; Roopnarine, 1995), the processes and pronounced anthropogenic environ- underlying geographic gradients in diversity mental change (e.g., eutrophication and (e.g., Roy & others, 1998; Crame, 2002; benthic trawling), and not postmortem shell Vermeij, 2005; Jablonski, Roy, & Valen- loss (Kidwell, 2007). These taphonomic tine, 2006), and the role of predation in analyses provide a foundation for examining evolutionary trends (e.g., Stanley, 1986b; ecological shifts in the Bivalvia over geologic Kelley, 1989; Dietl & others, 2002). This time as well as the susceptibility of bivalve depth of study is due in part to the high taxa with particular traits to extinction. preservation potential of bivalve shells in ESTIMATING EXTINCTION shallow marine environments. Marine bivalves are not free from the AND ORIGINATION FROM taphonomic insults experienced by other INCOMPLETE DATA marine taxa, but their fossil Accurately estimating extinction and origi- record is relatively complete (Valentine, nation rates is challenging for all taxa, due to 1989; Foote & Raup, 1996; Harper, 1998; incomplete observations. In paleontological Foote & Sepkoski, 1999; Kidwell, 2005; studies, the observed stratigraphic distribu- Valentine & others, 2006), and the tapho- tion of fossil occurrences is affected by pres- nomic biases that affect this record are increas- ervation and sampling, leading to temporal ingly well understood (Cooper & others, offsets between a ’s true time of origi- 2006; Valentine & others, 2006). Taxa that nation and extinction and its observed first have readily soluble shell microstructures, are and last occurrences (Signor & Lipps, 1982; small-bodied or thin-shelled, geographically Marshall, 1990; Meldahl, 1990; Foote, restricted, commensal or parasitic, epifaunal, 2000; Holland & Patzkowsky, 2002; and/or occur in deeper water are less likely to Foote, 2003). Preservation and sampling be preserved and sampled (Cooper & others, are biased by a number of factors, including 2006; Valentine & others, 2006). Yet, the the rarity and body size of taxa, as well as probability of being preserved and sampled the overall quality and quantity (complete- is relatively high for bivalves living at shelf ness) of the fossil record. The complete- to intertidal depths. Approximately 75% of ness of the fossil record varies systematically all living genera and subgenera of shallow through time with tectonic and/or climatic marine bivalves are also known from the fossil factors (e.g., Smith, Gale, & Monks, 2001; record (Valentine & others, 2006). Although Crampton, Foote, Beu, Cooper, & others, postmortem dissolution of primary shell 2006; S. Peters, 2006). It is also affected by aragonite has resulted in considerable loss the abundance of unlithified versus lithified of molluscan skeletal material from the rock sediments (Hendy, 2009; Sessa, Patzkowsky, record (Cherns & Wright, 2000, 2009), this & Bralower, 2009). that occur taphonomic filter does not appear to have during intervals of poor preservation and biased macroevolutionary patterns inferred sampling appear to happen earlier in time from fossil mollusks (Kidwell, 2005). (back-smearing), whereas originations in Recent studies have shown significant those same intervals appear to happen later agreement between ecological metrics calcu- in time (forward-smearing). This problem lated for molluscan death assemblages and is not unique to studies of the fossil record, Extinction in the Marine Bivalvia 3 but rather it presents a general challenge to zoic using a likelihood-based modeling any attempt to estimate extinction (or origi- approach developed by Foote (2003, 2005). nation) from limited observations (Solow, This approach uses numerical optimization 1993, 2005; Rivadeneira, Hunt, & Roy, to identify the time of extinction, 2009). origination, and preservation rates most The degree of discordance between the likely to have generated the observed data timing of true extinction and origination (i.e., the matrices of forward and backward and the observed stratigraphic ranges of taxa survivorship frequencies calculated from depends on temporal variation in preserva- the observed temporal distribution of first tion and sampling. Accounting for such vari- and last occurrences of genera) under a ation can be critical in reconstructing diver- given model of and preservation. sity dynamics over geologic time. Multiple Our analysis of bivalve diversity dynamics approaches have been developed to account differs from previous studies (e.g., Stanley, for variable preservation and sampling 1977; Gould & Calloway, 1980; Krug, in paleontological studies. The choice of Jablonski, & Valentine, 2009), in that method depends on the specific question completeness of the preserved and sampled being addressed and the spatiotemporal fossil record is explicitly taken into account scale of sampling. For example, at local or in estimating evolutionary rates. Our anal- regional scales, datasets may be partitioned ysis also focuses on rates of extinction and to examine only samples collected from origination rather than diversification rate or comparable taphonomic or stratigraphic standing diversity (cf. Stanley, 1977; Gould contexts (e.g., Scarponi & Kowalewski, & Calloway, 1980; Miller & Sepkoski, 2007; N. Heim, 2009). At the global scale, 1988). two general approaches have been taken We use a global compilation of observed to account for temporal variation in the first and last occurrences of marine bivalve completeness of the known fossil record: genera for rate estimation (Sepkoski, 2002). occurrence-based approaches that rely on Data for the 2861 bivalve genera in Sepkos- sub- or replicate-sampling methods, such ki’s Compendium of Fossil Marine as rarefaction (e.g., Alroy & others, 2001; Genera (2002) were compiled primarily from Bush, Markey, & Marshall, 2004; Alroy the first Treatise on Invertebrate & others, 2008) and capture-mark-recapture devoted to the Bivalvia (Cox & others, (e.g., Connolly & Miller, 2002; Liow & 1969; Stenzel, 1971); data for the revised others, 2008), and modeling approaches Treatise are as yet unavailable. The Paleo- that estimate rates of extinction, origination, Database (Alroy & others, 2001, and preservation simultaneously from the 2008)—a global compilation of spatial and observed paleontological data (Foote, 2000, temporal occurrences of fossil taxa through 2003, 2005). Preservation rate in this last the Phanerozoic—is another dataset that approach describes jointly the probability of could have been used to investigate bivalve preservation and sampling over time. evolutionary rates. Although occurrence- based data can also be analyzed in such a MARINE BIVALVE way as to account for variable sampling EXTINCTION AND and preservation (see above), we chose to ORIGINATION DYNAMICS analyze Sepkoski’s Compendium of first THROUGH THE and last occurrences to provide a bench- mark against which analysis of data from PHANEROZOIC the revised Treatise could be compared in Here we estimate extinction, origination, the future. While we are eager to see how and preservation rates simultaneously for results differ following the taxonomic revi- marine bivalve genera through the Phanero- sions anticipated in the revised Treatise, we 4 Treatise Online, number 29 do not expect substantial changes. Studies through the Phanerozoic are presented in conducted at comparably broad spatial, Figure 1. Rates of preservation vary consid- temporal, and taxonomic scales have shown erably over time, spanning nearly the full that taxonomic errors tend to be randomly range from zero to one, with a median rate distributed and overall macroevolutionary equal to 0.33. The median rate of preserva- patterns are surprisingly robust (Adrain & tion for bivalves in this analysis is lower than Westrop, 2000; Ausich & Peters, 2005; previous estimates (Foote & Sepkoski, 1999; Wagner & others, 2007). Valentine & others, 2006). This moderate Rates of extinction, origination, and pres- rate of preservation overall, combined with ervation for marine bivalve genera were esti- its volatility over time, underscores the mated for 71 time intervals that correspond importance of accounting for temporal roughly to geologic stages. Data for some variation in the completeness of the fossil stages were combined to minimize temporal record when estimating extinction and variation in interval duration (median interval origination rates. duration = 6.4 million ; interquartile In general, bivalves exhibit moderate range, 4.4 to 10.2 million years). Extinction rates of extinction and origination through and origination rates were calculated assuming the Phanerozoic (median rate of extinction a pulsed model of taxonomic turnover in = 0.1; median rate of origination = 0.2), which originations cluster at the start of each with rare intervals of elevated rates (Fig. interval and extinctions at the end of each 1). Prominent peaks in extinction occurred interval (Foote, 2003, 2005). Under this during the late , end-, model, extinction rate equals the number of Late , end-, end-, genera last appearing in an interval divided and end-, all times of elevated by the total diversity of the interval; origina- extinction observed at much broader taxo- tion rate equals the number of new genera nomic scales (Foote, 2003). These extinc- in an interval divided by the number present tion peaks have previously been observed at the start of the interval; and preserva- in studies of the fossil record that assume tion rate equals the estimated probability of perfect preservation, but it is noteworthy being preserved and sampled per genus per that they remain after accounting for the interval; see Foote (2003) for additional dramatic temporal variation observed in methodological details. We have also exam- preservation rate. ined extinction and origination rates generated A secular decline in bivalve extinction under an alternative evolutionary model in and origination rates over the Phanerozoic which taxonomic turnover occurs continu- is observed; the Spearman rank corre- ously (Foote, 2003). These models—pulsed lation between extinction and origination versus continuous turnover—both identify rate and time is 0.26 and 0.25, respectively peaks in origination and extinction rates for (p-value < 0.05 for both correlation tests). marine bivalves through the Phanerozoic, but The Phanerozoic-scale decline in rates of the magnitudes and timing of the peaks differ extinction and origination has also been somewhat. We restrict our discussion to the observed at broader taxonomic scales (Raup results of the pulsed turnover model, as this & Sepkoski, 1982; Van Valen, 1984; Foote, model has greater support globally (Foote, 2003) and may result from the loss of extinc- 2005) and regionally (Crampton, Foote, tion-prone lineages over time (Roy, Hunt, Beu, Maxwell, & others, 2006), and also & Jablonski, 2009), although other mecha- has the advantage of incorporating all genera, nisms have also been proposed (Alroy, 2008 including those confined to a single , in and references therein). Differing taxonomic the estimation of all three rates (Foote, 2003). practice could potentially contribute to the Rates of extinction, origination, and pres- observed temporal variation in bivalve rates ervation estimated for marine bivalve genera of taxonomic extinction and origination. Extinction in the Marine Bivalvia 5 0.8 0.8 0.6 0.6 ate ate 0.4 0.4 Extinction r Extinction r 0.2 0.2 0

C O S D C P Tr J K Cz 0 500 400 300 200 100 0 0 5 10 15 20 25 30 35 Geologic time (Ma) Frequency 7.0 7.0 6.0 6.0 ate ate r r 2.0 2.0 igination igination 1.0 O r O r 1.0 0

C O S D C P Tr J K Cz 0 500 400 300 200 100 0 0 10 20 30 40 50 Geologic time (Ma) Frequency 0.9 0.9 ate ate 0.6 0.6 ation r ation r v v 0.3 0.3 Prese r Prese r 0

C O S D C P Tr J K Cz 0 500 400 300 200 100 0 0 2 4 6 8 10 12 14 Geologic time (Ma) Frequency

Fig. 1. Extinction, origination, and preservation rates per interval for marine bivalves through the Phanerozoic. The left panel presents the mean time series of rate estimates derived from 100 bootstrap replicate samples of the observed data. The right panel presents the frequency distribution of rate magnitudes. Extinction and origination rates were estimated assuming a pulsed model of taxonomic turnover. Under this model, extinction rate equals the number of genera last appearing in an interval divided by the total diversity of the interval, origination rate equals the number of new genera in an interval divided by the number present at the start of an interval, and preservation rate is the estimated probability of preservation per genus per interval (new).

However, previous studies conducted at to expect rates of and comparable scales have generally found pseudoorigination to decline toward the taxonomic errors to be randomly distrib- present day. uted (Adrain & Westrop, 2000; Wagner To identify peaks of extinction that stand & others, 2007), and there is little reason out substantially above the baseline rate for 6 Treatise Online, number 29 2 2 0 0 ate residuals ate residuals r −2 −2 −4 −4 igination −6 Extinction r O r −6 C O S D C P Tr J K Cz C O S D C P Tr J K Cz

500 400 300 200 100 0 500 400 300 200 100 0 Geologic time (Ma) Geologic time (Ma)

Fig. 2. Residual variation in extinction and origination rates remaining after removing the secular decline in both rates through the Phanerozoic using linear regression. The so-called Big Five mass extinctions recognized in marine are denoted with large black diamonds (new). the portion of the Phanerozoic in which turnover at global and regional scales, but they occur, the long-term secular trend in such large events are relatively uncommon both evolutionary rates was removed by in the . The end-Cretaceous fitting a linear regression to the natural (K/Pg, formerly K/T) mass extinction and logarithm of evolutionary rate against time recovery is one example of a large-scale event (Alroy, 2008). When the residual varia- in which the biogeographic fabric of diver- tion in extinction rate is considered, the sity loss and rebound has received detailed late Cambrian, end-Permian, end-Triassic, study. At this time, regions differed little and end-Cretaceous are times at which in the severity of extinction experienced by extinction was particularly severe for marine marine bivalves, but markedly in the timing bivalves (Fig. 2). Depending on the cut-off and process of recovery (Raup & Jablonski, value used to define a severe extinction, the 1993; Jablonski, 1998). Examples of more and Plio– biogeographically differentiated intervals of are also noteworthy. This is consistent with extinction and recovery for marine bivalves previous studies that have documented include the Triassic– (e.g., Hallam, substantial molluscan extinction, at least 1981; Aberhan, 2002) and Plio–Pleistocene regionally, during these times (e.g., Raffi, (e.g., Raffi, Stanley, & Marasti, 1985; Stanley, & Marasti, 1985; Stanley, 1986a; Stanley, 1986a; Allmon & others, 1993; Allmon & others, 1993; Prothero, Ivany, Todd & others, 2002), among others. & Nesbitt, 2002; Dockery & Lozouet, 2003; Hansen, Kelley, & Haasl, 2004). DIVERSITY-DEPENDENT Global patterns such as these emerge DYNAMICS IN THE MARINE as the sum of processes of extinction and BIVALVIA THROUGH THE origination operating at finer spatial scales. Processes of extinction and recovery are PHANEROZOIC biogeographically complex. There are Over their history, marine bivalves have currently insufficient data to address spatial experienced periods of elevated extinc- variation in extinction and recovery in any tion and origination, as well as periods detail for many events. Intervals in which of relative evolutionary quiescence (Fig. extinction triggers are both pronounced 1). To what extent have the dynamics of and widespread should result in greater extinction and origination been coupled congruence between patterns of taxonomic over time? Extinction may facilitate origi- Extinction in the Marine Bivalvia 7 nation through the removal of incumbent cally standardized data with data aggregated taxa and opening up of ecospace. Under- from the literature without taxonomic stan- standing whether extinction and origina- dardization have generally found taxonomic tion rates operate in a diversity-dependent errors to be randomly distributed (Adrain & fashion has important implications for our Westrop, 2000; Wagner & others, 2007), understanding of the role of biotic interac- and rate estimates, as a result, to be affected tions in diversification (Sepkoski, 1978; little by the process of taxonomic standard- Miller & Sepkoski, 1988; Kirchner & ization (Wagner & others, 2007; but see Weil, 2000a, 2000b; Erwin, 2001; Lu, Ausich & Peters, 2005). within Yogo, & Marshall, 2006; Alroy, 2008; genera cannot be fully accounted for until Jablonski, 2008a). If diversity-dependent a comprehensive genus-level phylogenetic dynamics have been important for marine framework exists for the Bivalvia. bivalves through the Phanerozoic, then To determine whether evolutionary rates times of limited extinction should have been were diversity-dependent among marine followed by times of limited origination, bivalves through the Phanerozoic, we exam- and times of elevated extinction by times of ined the variation in rates of extinction elevated origination. Whether the response and origination that remains following the of origination to extinction was immediate removal of the long-term secular decline in or lagged by some period of time depends rates noted above. The effect of extinction on the nature of the recovery process. If on origination was evaluated using the slope extinction empties niches, then origination of a linear regression model relating the rate may respond rapidly to new ecological and of extinction in an interval (t) to the rate evolutionary opportunity. However, if niches of origination in the next interval (t + 1). depend in part upon diversity and need to We evaluated the support for an effect of be reconstructed following major perturba- extinction on subsequent origination by tions, then temporal lags between extinction assessing whether the observed regression and origination peaks are to be expected. slope was significantly greater than zero, Pseudoextinction—the apparent evolu- and whether it differed from that expected tionary turnover of taxa resulting from solely by chance via a permutation test. The anagenetic morphological evolution and/or distribution of null values against which the variation in taxonomic practice—could also observed slope was compared was generated contribute to a positive relationship between by randomly shuffling the detrended rates of extinction and subsequent origination, if extinction and origination and calculating rates of pseudoextinction are elevated over the slope of the extinction versus origination some intervals relative to others. If true, this relationship, and repeating this procedure is not necessarily less significant, as pseudo- 10,000 times. extinction presumably reflects some amount A significant positive relationship is of morphological change. Thus, temporal observed, such that periods of elevated variation in rates of pseudoextinction could extinction are followed by elevated origina- offer insight into the evolutionary response tion, and periods of moderate extinction of taxa to changes in the biotic and abiotic are followed by moderate origination (Fig. environment. In practice, pseudoextinction 3; Table 1). The results of the permutation is probably not a major factor governing test also indicate that the observed relation- the variation we observe in rates of extinc- ship between extinction rate and subsequent tion and origination—as well as their asso- origination rate among marine bivalves is ciation—over the Phanerozoic history of significantly greater than expected by chance marine bivalves. Previous studies conducted (Fig. 4). There is no indication of a lag in at comparable spatial, temporal, and taxo- the response of origination to extinction; nomic scales that have compared taxonomi- rather, origination responds immediately in 8 Treatise Online, number 29

Table 1. Effect of extinction on origination for

2 marine bivalve genera through the Phanero- zoic. Effect was measured as the slope of the

0 linear regression of extinction rate in an inter- ate (t+1)

r val (t) on origination rate in the next interval (t + 1). Rates were detrended prior to analysis; −2 details provided in text. A significant positive −4 igination relationship is observed; intervals of elevated

O r extinction are followed by intervals of elevated

−6 origination, and intervals of moderate extinc- tion followed by intervals of moderate origina- −6 −4 −2 0 2 tion. This diversity-dependent relationship be- tween extinction and origination is not driven Extinction rate (t) simply by mass extinctions and their associated recovery intervals. Excluding intervals char- Fig. 3. The effect of extinction on origination for ma- rine bivalve genera through the Phanerozoic. Plotted acterized by elevated extinction (e.g., the top are the rates of extinction in each interval (t) against 5% of extinctions, top 10%) does not mark- origination in the next interval (t + 1). Rates were loga- edly weaken the overall relationship (new). rithmically transformed and detrended prior to analysis; details provided in text. The dashed line is a linear Data Effect p-value regression model fitted to the extinction-origination All 0.3 0.02 relationship. A statistically significant positive relation- excluding top 5% 0.25 0.07 ship is observed, indicating that diversity-dependent excluding top 10% 0.22 0.12 processes have operated over the evolutionary history excluding top 20% 0.26 0.09 of the Bivalvia (new). excluding top 30% 0.33 0.04 the next interval, and this effect subsequently acterized by relatively low extinction rates weakens over time. The association between also exhibit diversity dependence (Table 1). extinction rate in an interval (t) and origi- Extinction has been an important evolu- nation rate in the next interval (t + 1) was tionary process throughout the history of approximately double that of extinction rate marine bivalves, varying considerably in and origination rate two intervals later (t + 2) intensity over time, but contributing consis- (i.e., slopes of 0.30 and 0.16 respectively). tently to bivalve diversification, in part, These results are consistent with studies of through its effect on rates of origination. the relationship between extinction and orig- ination for skeletonized marine invertebrates INFLUENCE OF BIOLOGICAL as a whole (Lu, Yogo, & Marshall, 2006; FACTORS ON EXTINCTION Alroy, 2008), and corroborate previous RISK AMONG MARINE work on marine bivalves that documented BIVALVES hyperexponential bursts of diversification following mass extinction events (Miller & Extinction selectivity, or the selective Sepkoski, 1988; Krug, Jablonski, & Valen- removal of taxa that possess particular tine, 2009). It is important to note, however, ecological or evolutionary traits, also plays that the diversity-dependent relationship an important role in shaping macroevo- between rates of extinction and origination lutionary and macroecological patterns for marine bivalves is not limited to mass through time. Extinction selectivity can extinctions and their associated recoveries. contribute to ecosystem reorganization by While removing the most extreme extinc- eliminating dominant taxa and allowing tion events from the analysis somewhat subordinate ones to diversify (Gould & weakens the relationship between extinction Calloway, 1980; Jablonski, 1986, 1989; and subsequent origination, intervals char- Droser, Bottjer, & Sheehan, 1997); it can Extinction in the Marine Bivalvia 9 redirect evolutionary or ecological trends by

eliminating important innovations (Pojeta 1500 & Palmer, 1976; Fürsich & Jablonski, 1984; Jablonski, 1986); and it can limit the

potential evolution of clades by reducing 1000 variability (Norris, 1991; Liu & Olsson, 1992). By studying extinction selectivity Frequency over long time scales, one can assess not 500 only which taxa went extinct, but poten- tially how and why they did so. This link between extinction pattern and process can 0 help to bridge the gap between paleontology −0.6 −0.3 0.0 0.3 0.6 and (see papers in Effect of extinction (t) on origination (t+1) Dietl & Flessa, 2009). If we can determine which traits have influenced susceptibility to Fig. 4. The effect of extinction on subsequent origina- tion for marine bivalve genera through the Phanerozoic. extinction during periods of past environ- Effect was measured using the slope of the linear regres- mental change, then we may be better able sion of extinction rate in an interval (t) on origination to predict which organisms are most likely rate in the next interval (t + 1). The gray frequency distribution presents the randomized values, the solid to go extinct or persist in the present day. arrow denotes the observed effect, and the dashed line Extinction selectivity is thought to have indicates the 95th quantile of randomized values. The significantly influenced the evolutionary observed effect is significantly greater than expected by trajectory of marine invertebrates and the chance (new). ecological structure of marine ecosystems through geologic time. Bivalves are no tests) and (114 tests) bivalves, exception; indeed, several classic studies with the Paleozoic receiving considerably of extinction selectivity have focused on less attention (18 tests). The extinctions the long and relatively well-preserved fossil represented in our database range from the record of marine bivalves (e.g., Bretsky, largest and most catastrophic mass extinc- 1973; Kauffman, 1978; Jablonski, 1986; tions (including all of the so-called Big Five), Stanley, 1986a; Jablonski, 2005). This to six smaller, possibly regional-scale, events is in part because bivalves display suffi- (e.g., Eocene/Oligocene and Plio/Pleisto- cient variation among taxa in traits such as cene) and background intervals. Selectivity morphology, feeding mode, life habit, larval has been examined at both the (98 type, geographic range, and stratigraphic tests) and genus (80 tests) levels. Examining range to allow workers to independently the specific traits tested for selectivity, four test the extent to which these traits relate to traits have received the most attention: taxon survivorship. geographic range (27 tests), life habit (28 A review of the literature on extinction tests), body size (21 tests), and feeding mode selectivity in fossil marine bivalves published (15 tests). in 2010 and before (see Appendix, p. 20) If extinction is defined as the point in demonstrates that selectivity among taxa time at which a taxon’s geographic range has been explored with respect to a wide and abundance decrease to zero, taxa variety of traits, such as abundance, feeding with broader geographic ranges should mode, life habit, geographic range, body be less prone to extinction. The primary size, temperature tolerance, species richness, role that geographic distribution plays in and habitat breadth, among many others determining survivorship has long been (33 traits in total). This review includes 170 recognized for both modern and fossil taxa tests of extinction selectivity published in (see references in Gaston, 1994; Rosen- 69 studies. The vast majority of selectivity zweig, 1995; Payne & Finnegan, 2007). studies have focused on (120 An early assessment of global survivorship 10 Treatise Online, number 29 across four mass extinctions—the end- & Aberhan, 2007; for exceptions, see Rode Ordovician, Late Devonian, end-Permian, & Lieberman, 2004, for the Late Devonian, and end-Triassic events—concluded that and Stilwell, 2003, for the K/Pg). Late geographically widespread bivalve genera extinctions, intermediate in scale were more likely to survive, at least in the between the Big Five events and background initial stages of an , before extinction, differ in the effects of geographic drastic deterioration of the physical envi- range on selectivity among regions. ronment (Bretsky, 1973). The event that Narrowly distributed species were more has been most thoroughly examined for likely to become extinct in western South geographic range selectivity is the K/Pg mass America (Rivadeneira & Marquet, 2007) extinction, in conjunction with the interval and tropical America (Roopnarine, 1997), of background extinction leading up to it. but not in western , where K/Pg survivorship patterns in bivalves and there is no evidence of selective extinction gastropods, along the United States Gulf (Stanley, 1986b). These complex patterns and Atlantic Coastal Plain and globally, highlight the importance of assessing selec- suggest that the only trait reliably associ- tivity across a range of extinctions that differ ated with genus survivorship is geographic in magnitude, as well as across a range of range, whereas several traits are associated taxonomic levels. It is possible that thresh- with genus- and species-level survivorship olds exist, such that geographic range no during the preceding background interval longer ensures the survival of a species, when (Jablonski, 1986, 1987, 1989; Raup & the scale of environmental perturbation Jablonski, 1993; Jablonski & Raup, 1995; and extinction exceed a critical magnitude. Jablonski, 2005; Jablonski & Hunt, 2006). If true, this has enormous implications for Data for Southern Hemisphere bivalves assessment of extinction risk and develop- across the K/Pg, also compiled at the genus ment of effective management strategies in level, support this general pattern (Stilwell, modern ecosystems. 2003). In contrast, during the end-Triassic Bivalve life habit, specifically the sites the extinction, European bivalve species with occupy relative to the sediment- broader distributions appear no more likely water interface, is another ecological trait to have survived this event than narrowly that is thought to affect survivorship. distributed taxa (McRoberts & Newton, Which life habit favors survival probably 1995; McRoberts, Newton, & Alla- depends on the mechanism of extinction. sinaz, 1995). When the spatial scale of For example, epifaunal taxa are thought to analysis is expanded to global coverage, the be more vulnerable to predation pressure same nonsignificant species-level pattern is (Stanley, 1977, 1982, 1986b; Vermeij, observed for the end-Triassic (Kiessling & 1987), which could lead to decreased popu- Aberhan, 2007). lation size and an increase in extinction risk. Taxonomic level and the intensity In contrast, an epifaunal life habit may be of extinction complicate the pattern of advantageous in escaping sudden changes in geographic range selectivity. The data bottom water chemistry and/or oxygenation. compiled here strongly suggest that wide- Of the studies compiled here, 14 suggested spread bivalve species were significantly more that epifaunal taxa were more likely to likely to survive background (Jablonski, become extinct than infaunal, 5 suggested 1986, 1987; Jablonski & Hunt, 2006; the opposite, and 8 found no evidence of Kiessling & Aberhan, 2007; Crampton selectivity either way. When these results are & others, 2010; Harnik, 2011), but not parsed according to the size of the extinction mass extinction events (Jablonski, 1986; event, an interesting pattern emerges. The Hansen & others, 1993; McRoberts & majority of background intervals studied Newton, 1995; McRoberts, Newton, & (8 out of 10) suggest that infaunal bivalves Allasinaz, 1995; Jablonski, 2005; Kiessling were less likely to go extinct, while mass Extinction in the Marine Bivalvia 11 extinctions yield contradictory results (4 increased body size is often associated negative, 5 positive, 6 nonsignificant). When with increased fecundity, broader environ- the mass extinction events are broken down mental tolerance, and wider geographic by specific event, the results remain mixed. range (Stanley, 1986b; McKinney, 1990; For example, while bivalve genera glob- Rosenzweig, 1995; Hildrew, Raffaelli, & ally did not exhibit differential survival Edmonds-Brown, 2007), which suggests with respect to life habit across the end- that larger taxa should have increased rates Permian mass extinction (Jablonski, 2005), of survivorship. Among marine bivalves, regional patterns from China suggested however, large body size is not generally asso- greater losses of epifaunal than infaunal ciated with either extinction risk or survivor- bivalve genera (Knoll & others, 2007). ship. Body size and extinction are positively Perhaps these differences reflect the extent linked in only 7 and negatively linked in only to which different geographic regions were 2 (out of a total of 21) studies. Four of the 7 affected by environmental deterioration. In studies that documented selective extinction another example, preferential extinction of of large taxa focused on regional patterns infaunal bivalve species was documented during intervals characterized by background across the K/Pg boundary in New Jersey rates of extinction; these include the Jurassic and the Delmarva Peninsula of the United (Hallam, 1975), (Anderson & States (Gallagher, 1991), but subsequent Roopnarine, 2003, for the Western Atlantic work found the opposite pattern for bivalve and , but not the Eastern Pacific), species in Denmark (Heinberg, 1999) and and Pleistocene (Stanley, 1986a, 1990b). the Southern Hemisphere (Stilwell, 2003). Interestingly, not a single one of the 10 Although global analyses of selectivity studies that considered size across a mass can be very useful in seeking possible causes extinction event found a strong, conclu- of extinction, they can obscure regional sive link between body size and extinction, patterns that may be less predictable and yet although the only 2 events investigated likely to provide more information about thus far are the end-Triassic (McRoberts & the interacting effects of biotic and abiotic Newton, 1995; McRoberts, Newton, & factors on survivorship. Spatial variation in Allasinaz, 1995; McRoberts, Krystyn, & environmental change, coupled with spatial Shea, 2008) and K/Pg (Hansen & others, heterogeneity in the distributions of taxa and 1987; Raup & Jablonski, 1993; Jablonski associated biological traits, effectively ensure & Raup, 1995; McClure & Bohonak, that patterns of selectivity will vary region- 1995; Jablonski, 1996a; Lockwood, 2005; ally (see Fritz, Bininda-Emonds, & Purvis, Aberhan & others, 2007) events. Three 2009, for an example of geographic variation of these 10 studies documented a decrease in extinction risk among extant ). in bivalve size across a mass extinction Spatial variation may provide useful informa- boundary (Norian–Rhaetian: McRoberts, tion about gradients of environmental change Krystyn, & Shea, 2008; K/Pg: Hansen & and the existence of environmental thresholds others, 1987; Aberhan & others, 2007), affecting taxon survivorship. Despite the but it is unclear in each case whether these clear importance of regional-scale studies in patterns were driven by extinction selectivity, modern conservation biology, paleontological within lineage size change, or size-biased examples are few and far between. origination. Although large body size is widely thought One of the few instances in which a to increase extinction risk in vertebrates, connection between large body size and the link between size and extinction risk in survivorship has been documented convinc- marine invertebrates is considerably more ingly focused on scallops across the Plio– ambiguous (Hallam, 1975; Stanley, 1986b; Pleistocene extinction in California (Smith Budd & Johnson, 1991; Jablonski, 1996b; & Roy, 2006). This positive relationship was Smith & Roy, 2006). Among invertebrates, not apparent until phylogenetic relationships 12 Treatise Online, number 29 were considered. This emphasizes an under- & Thayer, 1991; Raup & Jablonski, 1993; appreciated problem that may affect many Jablonski & Raup, 1995; Jablonski, 1996a; selectivity studies. Patterns of selectivity can Stilwell, 2003; Aberhan & others, 2007). sometimes be masked or artificially exagger- There is preliminary evidence to suggest ated when phylogenetic relationships are not that the strength of the selectivity may taken into account (Purvis, 2008). Taxa may have increased with distance away from the share a particular trait and similar pattern United States Gulf Coastal Plain, which of survivorship because they are related raises the question as to whether proximity to each other and not necessarily because to the killing agent, in this case the K/Pg the trait under consideration, by itself, bolide impact that occurred in the Yucatan, confers survivorship. A recent analysis (Roy, has an effect on selectivity patterns. Studies Hunt, & Jablonski, 2009) of Jurassic to limited to eastern Texas (Hansen & others, Recent bivalves demonstrated conclusively 1987; Hansen, Farrell, & Upshaw, 1993; that phylogenetic clustering of extinction Hansen & others, 1993, for exception see occurs. Phylogenetic relationships do not Sheehan & Hansen, 1986) or the United always affect patterns of selectivity, however; States Gulf and Atlantic Coastal Plain for example, patterns of selectivity among (McClure & Bohonak, 1995), have yielded Cenozoic mollusks from New Zealand did either weak or no evidence for selectivity. not change appreciably after accounting On the other hand, regional studies in the for phylogeny (Foote & others, 2008; Southern Hemisphere (e.g., Stilwell, 2003) Crampton & others, 2010). The fact that and Argentina (Aberhan & others, 2007) taxa in some clades are significantly more have suggested that proximity matters, as extinction-prone than others does strongly they have shown strong evidence for selec- suggest that future paleontological studies tivity. Although Jablonski’s work (Raup & of selectivity should explicitly account for Jablonski, 1993; Jablonski & Raup, 1995; phylogenetic effects. Jablonski, 1996a) clearly supports a global In general, deposit feeding is thought pattern of selective extinction of suspension- to represent a more generalized dietary feeding bivalves at the K/Pg, he has argued mode than suspension feeding and could that this was driven by taxonomic factors, promote survivorship, especially across rather than selectivity according to feeding events that involve a collapse in primary mode. He and his colleagues pointed to productivity. Levinton’s (1974) observation anomalously low rates of extinction in the that genera of deposit-feeding bivalves were two bivalve orders Nuculoida and Lucinoida geologically longer-lived than suspension- and argued that other attributes of these two feeders, has inspired an ongoing debate clades helped to promote their survivorship. over whether bivalves with different feeding Sheehan and Hansen (1986), Hansen and modes experience different extinction trajec- others (1987, 1993), and Hansen, Farrell, tories. Building on this work, a qualitative and Upshaw (1993) emphasized the shift examination of background extinction in from molluscan communities dominated by Cretaceous bivalve species documented suspension-feeders to those dominated by particularly slow rates of evolution and long deposit-feeders across the K/Pg boundary, a stratigraphic durations in deposit-feeders pattern that could have been caused by selec- relative to suspension-feeders (Kauffman, tive extinction against suspension-feeders, 1978). The majority of studies that have preferential recovery of deposit-feeders, explicitly tested for extinction selectivity or some combination of the two. Explicit according to feeding mode have focused on evaluation of this possibility, in addition to the K/Pg event, with mixed results. Seven detailed tracking of feeding mode across the out of 11 studies have reported selective recovery interval, is still lacking. Although extinction of suspension-feeding genera early studies heralded the usefulness of (e.g., Sheehan & Hansen, 1986; Rhodes patterns of extinction selectivity based on Extinction in the Marine Bivalvia 13 feeding habits in differentiating among was geographic range. Such multivariate possible extinction mechanisms, this poten- approaches are crucial to studies of selec- tial has seldom been realized (but see Knoll tivity, offering considerable insight into the & others, 1996, 2007, for exceptions). As direct and indirect effects of extinction on our understanding of changes in primary the evolution of correlated traits. A clear productivity associated with mass extinc- understanding as to how traits interact, tions deepens, aided by geochemical proxies, influencing extinction risk across a range of it should be possible to further refine and past events, is needed, if bivalve workers are test hypotheses bearing on the relationship to make such patterns relevant to managers between feeding mode and extinction risk predicting biotic response to current extinc- across an array of marine environments. tion pressures. Most of the studies outlined above focus on the selectivity of single traits and do not CONCLUSIONS consider the potential interactions among One of the major insights of paleontology multiple traits. We have every reason to is the importance of extinction in shaping believe, based on ecological studies of extant the diversity of life through time. The bivalves and many other clades, that several effects of extinction on diversity dynamics of these traits, for example, body size and have been intensively studied in the marine geographic range (Jablonski & Roy, 2003; Bivalvia because of their relatively complete Crampton & others, 2010; Harnik, 2011), fossil record, the considerable biological are linked to one another. This raises the variation that exists among taxa, and their question—to what extent do these interac- diversity and abundance in shallow marine tions influence patterns of selectivity? A environments today and in the past. In this handful of recent studies have tackled this contribution, we provide new estimates of question for marine bivalves (Jablonski & global extinction and origination rates for Hunt, 2006; Rivadeneira & Marquet, marine bivalve genera through the Phanero- 2007; Jablonski, 2008a; Crampton & zoic that explicitly account for temporal others, 2010; Harnik, 2011). Almost all variation in preservation. These analyses, of them have found that geographic range using data compiled primarily from the first played a more important role in survivorship Treatise on Invertebrate Paleontology (Part than any other ecological trait. For example, N: Cox & others, 1969; Stenzel, 1971), in a genus-level analysis of selectivity across underscore the important contributions of the K/Pg mass extinction, Jablonski (2008a) the Treatise to our understanding of bivalve independently tested the effects of body macroevolutionary history. While rates of size, geographic range, and species richness, extinction and origination are moderate and found that the last two traits were both for marine bivalves overall, times of severe statistically significantly correlated with extinction and times of general quiescence survivorship. However, once the covariation are observed through the Phanerozoic. Inter- among these three traits was controlled for, vals of elevated global extinction for marine geographic range yielded the only significant bivalves correspond to intervals of elevated evidence for selectivity. In what is perhaps extinction for marine invertebrates more the most extensive multivariate selectivity broadly, and bivalves exhibit secular declines study to date, Crampton and others (2010) in rates of extinction and origination over assessed the relative importance of several the Phanerozoic that are also observed at traits, including geographic range, body size, much broader taxonomic scales. Throughout feeding mode, life habit, and larval type, in their history, marine bivalves exhibited promoting survivorship among Cenozoic coupled dynamics of extinction and origi- bivalve species from New Zealand. 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Fisheries 18(9):6–22. 20 Treatise Online, number 29 Notes nuculoids and lucinids. May or may not be due to extinction selectivity. nuculoids and lucinids. May pectinoids, not significant in carditoids and glossoids. and glossoids. Size decrease may or not be due to extinction selectivity. decrease Size Geo range more important than abundance in determining extinction risk. range more Geo extinction rates. and abundant genera exhibit elevated Rare excluding boundary, lifestyles across of individuals with active Decline may or not be due to extinction selectivity. decrease Size or may not be extinction selectivity. May to smaller bivalves. Shift in negative in veneroids, and duration positive size between Correlation arcticoids, include veneroids, Large taxa radiate after extinction. Groups arcticoids, include veneroids, taxa radiate after extinction. Groups Small significant only when adjusted for phylogenetic bias. Pattern only. bivalves Siphonate only. bivalves Siphonate feeders only. Suspension taxa radiate during recovery. burrowing Deeper Location

Austria US Gulf CP US Gulf Global Global US CP US CP Global Argentina Global Tibet US CP Global Global Argentina Carib/W Atl E Pacific NZ W Euro Texas E CP US Gulf Global Global US CP N Am, Euro N Am, Euro US CP Euro Euro Global California SE US W N Am, Japan W N Am, Japan N Amer N Am, Euro

Age Nor/Rhaet Eo T/J Tri-Jur K/Pg K/Pg post Pz K/Pg K/Pg T/J K/Pg K/Pg K/Pg K/Pg Mio Mio Cz Jur K/Pg Eo K/Pg K/Pg Late Cret E/O K/Pg K/Pg T/J T/J K/Pg Plio/Pleist Plio/Pleist Pleist-Rec Pleist-Rec Cret K/Pg

Taxon level sp sp sp sp gen gen gen sp gen sp gen gen gen sp gen gen sp gen, sp sp sp gen gen sp gen gen gen sp sp gen gen sp sp sp sp gen

and should not be considered exhaustive (new). exhaustive and should not be considered Taxon 3 superfm biv biv biv biv biv moll dom biv biv dom biv gast biv, biv moll dom corbulids corbulids biv ammon biv, biv 3 superfm biv biv gast biv, 3 groups 3 groups biv biv biv Monotis biv scallops biv biv biv biv 3 groups NS NS Neg NS NS Nonlinear Pos? Pos Pos NS NS NS Pos? Pos NS NS Pos Pos? Varies NS NS NS NS NS NS NS NS Pos? NS Neg Neg Pos Pos Neg NS Pattern Reference 2011 Harnik, 2007 Kiessling & Aberhan, 2007 Kiessling & Aberhan, Lockwood, 2003 & Bohonak, 1995 McClure 2009 & Harnik, Simpson & others, 2007 Aberhan 1991 Thayer, Rhodes & & others, 2008 Hautmann 1995 & Raup, Jablonski 1996a Jablonski, 1993 Raup & Jablonski, & others, 2007 Aberhan 2003 Anderson & Roopnarine, 2003 Anderson & Roopnarine, & others, 2010 Crampton 1975 Hallam, & others, 1987 Hansen 2011 Harnik, 1995 & Raup, Jablonski 1996a Jablonski, 1996b Jablonski, Lockwood, 2005 Lockwood, 2005 & Bohonak, 1995 McClure 1995 & Newton, McRoberts & Allasinaz, 1995 Newton, McRoberts, Krystyn, 2008 & Shea, McRoberts, 1993 Raup & Jablonski, 2006 & Roy, Smith 1986a Stanley, 1986a Stanley, 1990b Stanley, Kauffman, 1978 Lockwood, 2004 Database of extinction selectivity studies in fossil marine bivalves; please note that the compilation of past work summarized here is ongoing ongoing is here summarized work past of compilation the that note please bivalves; marine fossil in studies selectivity extinction of Database . endix

pp A Trait Abundance Abundance Abundance Abundance Abundance Abundance locom Active locom Active Aragonite range Bathy range Bathy range Bathy Body size Body size Body size Body size Body size Body size Body size Body size Body size Body size Body size Body size Body size Body size Body size Body size Body size Body size Body size Body size Body size depth Bur depth Bur Extinction in the Marine Bivalvia 21 but not taxa inhabiting open shelf. change. Both infaunal and epifaunal selectivity or environmental boundary. across suspension-feeders decrease extinction selectivity. may or not be due to extinction selectivity. boundary, water epifaunal taxa. to shallow and detritus feeders (Nuculoida). Veneracea) Pteriacea, (Pectinacea, may be artifactual. Increased survival for taxa inhabiting middle-outer platform environments, Increased in infaunal suspension feeders may be due to extinction Long-term increase may or not be due to in abundance of infaunal bivalves Increase higher latitudes. In in infaunal species across increase habitat comparison shows Within . Excluding rudists. Excluding evolutionary rates among infaunal and deep water epifaunal, relative Slower suspension-feeders Comparison of mortality selective rates between infaunal taxa indicates that selectivity preservation of early Jurassic Poor

US CP W N Am, Japan Andean basin Andean basin Global SE US N Atl N Amer Andes NW Euro, Andean basin Global Global Global NZ CP US Atl Tibet Denmark Global Global Global Global Global W Interior N Amer China York New Global NW Euro US CP Euro Global Euro Global W S Amer (Continued).

K/Pg Pleist-Rec Pliens-Toar Pliens-Toar T/J Plio/Pleist Plio F/F early Jur Pliens-Toar Perm O/S Cret Cz K/Pg T/J K/Pg K/Pg K/Pg K/Pg P/T T/J-K/Pg Cret Cret P/T Rec Phan T/J K/Pg T/J T/J T/J K/Pg Late Ng endix pp A

gen sp sp sp gen sp sp sp sp sp gen gen gen, sp sp sp sp sp gen gen gen gen gen sp sp gen sp gen gen gen sp gen sp gen sp biv biv biv biv biv biv moll brach biv, biv biv gast biv, biv biv biv moll biv dom biv biv biv biv biv biv biv biv biv 6 species 4 groups biv biv biv biv biv biv biv NS Neg Pos Pos Neg Pos Pos notes See Neg Pos? Pos? Pos Pos NS Neg Neg Pos? NS NS NS NS Pos Pos Pos Pos Pos Pos - NS Neg Pos Neg NS Pos McClure & Bohonak, 1995 McClure 1990b Stanley, & Fürsich, 1997 Aberhan & Fürsich, 2000 Aberhan 1981 Hallam, 1986a Stanley, 1986 Vermeij, & Lieberman, 2004 Rode 2003 & Baumiller, Aberhan & Fürsich, 1997 Aberhan 2007 Clapham & Bottjer, Cope, 2002 2002 Crame, & others, 2010 Crampton 1991 Gallagher, & others, 2008 Hautmann 1999 Heinberg, 1995 & Raup, Jablonski 1996a Jablonski, 2005 Jablonski, 2005 Jablonski, 2005 Jablonski, Kauffman, 1977 Kauffman, 1978 Knoll & others, 2007 Levinton, 1973 Levinton, 1974 2008 Twitchett, & Mander & Bohonak, 1995 McClure 1995 & Newton, McRoberts 2001 McRoberts, & Allasinaz, 1995 Newton, McRoberts, 1993 Raup & Jablonski, 2007 & Marquet, Rivadeneira Bur depth Bur depth Bur Endemicity Endemicity Endemicity Endemicity Endemicity Environment Epifaunal Epifaunal Epifaunal Epifaunal Epifaunal Epifaunal Epifaunal Epifaunal Epifaunal Epifaunal Epifaunal Epifaunal Epifaunal Epifaunal Epifaunal Epifaunal Epifaunal Epifaunal Epifaunal Epifaunal Epifaunal Epifaunal Epifaunal Epifaunal Epifaunal Epifaunal 22 Treatise Online, number 29 , may or not be due to extinction selectivity. 2 S and CO 2 Notes margins) associated with higher extinction risk in bivalves. H Used morphology as proxy for escalation. morphology as proxy Used for escalation. morphology as proxy Used for escalation. morphology as proxy Used for escalation. morphology as proxy Used for escalation, only one trait (crenulate morphology as proxy Used risk. extinction predicting in mode larval than important more is range Geo extinction resistant. species more containing widespread Genera risk. extinction determining in abundance than important more range Geo severely. of endemism affected more with higher degrees Regions tolerant of probably genera flourish after P/T, 4 cosmopolitan bivalve Location

SE US S hemis US CP US CP US CP US CP Mississippi Global Global Global Global NZ Global Texas E CP US Gulf US CP Global US CP US CP US CP US CP N Am US CP US CP Global Global Euro Euro Global W S Amer N Amer Amer trop W N Amer S hemis Global China China N Amer (Continued).

Age

Plio/Pleist K/Pg E/O K/Pg Mio Plio/Pleist K/Pg F/F O/S P/T T/J Cz T/J K/Pg Eo Cret-Cz K/Pg K/Pg K/Pg Late Cret Cret K/Pg K/Pg Late Cret T/J Tri-Jur T/J T/J K/Pg Late Ng F/F Late Ng Late Ng K/Pg P/T P/T P/T F/F endix pp A Taxon level

sp gen, sp sp sp sp sp gen, sp gen gen gen gen sp gen sp sp sp gen gen sp w/in gen sp w/in gen sp gen sp w/in gen sp w/in gen sp sp sp sp gen sp sp sp sp gen, sp gen gen gen sp Taxon biv moll gast biv, gast biv, gast biv, gast biv, gast biv, biv biv biv biv biv biv moll 3 superfm moll biv gast biv, gast biv, gast biv, gast biv, gast biv, gast biv, gast biv, biv biv biv biv biv biv brach biv, chionines biv moll biv biv biv brach biv, Pattern NS Pos NS NS NS Pos NS Neg Neg Neg Neg Neg Neg NS Neg Neg Neg NS NS Neg Neg Neg NS Neg NS Neg NS NS Neg Neg Neg Neg NS Neg Pos? Pos Pos Neg Reference Stanley, 1986a Stanley, 2003 Stilwell, & others, 1999 Hansen & others, 1999 Hansen & others, 1999 Hansen & others, 1999 Hansen 2005 & Kelley, Reinhold 1973 Bretsky, 1973 Bretsky, 1973 Bretsky, 1973 Bretsky, & others, 2010 Crampton 1997 Wignall, & Hallam & others, 1993 Hansen 2011 Harnik, 2006 & Hunt, Jablonski 1995 & Raup, Jablonski 1986 Jablonski, 1986 Jablonski, 1986 Jablonski, 1987 Jablonski, 1989 Jablonski, 2005 Jablonski, 2005 Jablonski, 2007 Kiessling & Aberhan, 2007 Kiessling & Aberhan, 1995 & Newton, McRoberts & Allasinaz, 1995 Newton, McRoberts, 1993 Raup & Jablonski, 2007 & Marquet, Rivadeneira & Lieberman, 2004 Rode 1997 Roopnarine, 1986b Stanley, 2003 Stilwell, Bottjer & others, 2008 Knoll & others, 1996 Knoll & others, 1996 & Lieberman, 2004 Rode

Trait Epifaunal Epifaunal Escalation Escalation Escalation Escalation Escalation range Geo range Geo range Geo range Geo range Geo range Geo range Geo range Geo range Geo range Geo range Geo range Geo range Geo range Geo range Geo range Geo range Geo range Geo range Geo range Geo range Geo range Geo range Geo range Geo range Geo range Geo range Geo Hypercapnia Hypercapnia Hypercapnia Invaders Extinction in the Marine Bivalvia 23 can be as strong as geographic range in specific superfamilies.can be as strong planktonic larva not significant. substantial period of time. in determining risk. in determining risk. and detritus feeders (Nuculoida). Veneracea) Pteriacea, (Pectinacea, Epifaunal taxa only. Epifaunal and body size, Epifaunal, suspension-feeder, range significant. Geographic but overall weak Body size weak. Abundance range strong. Geographic important than larval range more Geographic type. and species richness nonsignificant. range significant. Body size Geographic significant. geographic range, body size Epifaunal, been in decline for a already those that have risk are at greatest Species important range is more be example of trait hitch-hiking. Geographic May important range is more be example of trait hitch-hiking. Geographic May within phylogenetic framework. Examined for stenotopy. lithofacies tolerance as proxy Used suspension-feeders Comparison of mortality selective rates between

Global China N Amer Florida NZ CP US Gulf US CP Global W S Amer NZ Global Global NZ CP US Atl US CP US CP W N Amer US CP Global US CP Euro US CP US CP N Am, Euro US CP US CP US CP California N Atl Global US CP SE US W Atl E Pacific Texas E W Interior N Amer Global (Continued).

Cret P/T Cret Late Ng Cz Eo Cret-Cz K/Pg Late Ng Cz K/Pg Jur-Rec Cz K/Pg Cret-Cz Cret Late Ng K/Pg K/Pg K/Pg K/Pg K/Pg Late Cret K/Pg K/Pg Late Cret K/Pg Plio/Pleist Plio T/J K/Pg Plio/Pleist Late Ng Mio K/Pg Cret Cret Phan endix pp A

gen, sp gen sp sp sp sp sp gen sp sp gen fam, gen sp sp sp sp sp gen gen gen gen sp sp gen gen gen gen gen sp gen gen sp sp gen sp sp sp gen biv biv biv veneroids biv 3 superfm moll biv biv moll rudists biv biv moll moll gast biv, biv gast biv, biv biv biv gast biv, gast biv, gast biv, gast biv, gast biv, biv scallops moll biv biv biv biv corbulids moll biv biv 4 groups

Pos Neg NS Neg notes See notes See notes See notes See notes See Neg Pos Pos NS Pos NS NS NS NS Pos NS NS NS Neg NS NS Neg NS Neg NS Pos NS Pos Pos NS NS Pos Mixed Pos Crame, 2002 Crame, Knoll & others, 2007 Kauffman, 1978 2011 Lockwood, & Hunt, Kolbe, & others, 2010 Crampton 2011 Harnik, 2006 & Hunt, Jablonski 2008a Jablonski, 2007 & Marquet, Rivadeneira & others, 2007 Foote 2008a Jablonski, 2009 & Jablonski, Hunt, Roy, & others, 2010 Crampton 1991 Gallagher, 2006 & Hunt, Jablonski 1987 Jablonski, 1986b Stanley, 1986 & Jablonski, Valentine 2008a Jablonski, & Bohonak, 1995 McClure 1986 Hoffman, 1986 Jablonski, 1986 Jablonski, 1989 Jablonski, 2005 Jablonski, 2005 Jablonski, & Bohonak, 1995 McClure 2006 & Roy, Smith 1986 Vermeij, 1981 Hallam, & Bohonak, 1995 McClure 1986a Stanley, 1990a Stanley, 2003 Anderson & Roopnarine, & others, 1993 Hansen Kauffman, 1977 Kauffman, 1978 Levinton, 1974

Latitude rate Meta com Morph var Morph Multivariate Multivariate Multivariate Multivariate Multivariate Occupancy Pachyodont Phylogenetic larvaPlank larvaPlank larvaPlank larvaPlank larvaPlank larvaPlank Schizodont thick Shell richness Sp richness Sp richness Sp richness Sp richness Sp richness Sp richness Sp richness Sp richness Sp Stenohaline Stenotherm Stenotherm Stenotherm Stenotopic Stenotopic Stenotopic Stenotopic Stenotopic 24 Treatise Online, number 29 Notes selectivity. and detritus feeders (Nuculoida). Veneracea) Pteriacea, (Pectinacea, Selectivity varies according to site. according varies Selectivity or may not be extinction selectivity. susp to dep feeders. May from Shift or may not be extinction susp to dep feeders at one site. May from Shift susp to dep feeders. Both go extinct at boundary. Long-term shift from and Lucinoida. extinction in Nuculoida due to low Selectivity pattern. Argues that this is a taxonomic suspension-feeders Comparison of mortality selective rates between and Lucinoida. extinction in Nuculoida due to low Pattern taxa only. Epifaunal

Location

Tethys W S hemis Argentina NZ Texas E Texas E Texas E Tibet Global Global N Am Global US CP Global Global Texas E S hemis W S Amer SE US US CP Global York New US CP Global California

(Continued).

Age Perm K/Pg K/Pg Cz K/Pg K/Pg K/Pg T/J K/Pg K/Pg Cret Phan K/Pg K/Pg K/Pg K/Pg K/Pg Late Ng Plio/Pleist K/Pg K/Pg Rec K/Pg K/Pg Pleist endix pp A

Taxon level gen gen, sp sp sp sp sp sp sp gen gen sp gen gen gen gen sp gen, sp sp sp gen gen sp gen gen sp

Taxon brach biv, moll moll dom biv biv moll moll biv dom biv biv biv 4 groups biv biv biv moll moll biv biv biv gast biv, 6 species biv biv moll Varies Neg Pos NS Pos? Pos? NS Pos Pos Pos Pos Pos NS Pos Pos Pos Pos NS Pos NS NS Pos NS NS Pos Pattern

Reference 2009 Posenato, 2003 Stilwell, & others, 2007 Aberhan & others, 2010 Crampton & others, 1987 Hansen 1993 & Upshaw, Farrell, Hansen, & others, 1993 Hansen & others, 2008 Hautmann 1995 & Raup, Jablonski 1996a Jablonski, Kauffman, 1978 Levinton, 1974 & Bohonak, 1995 McClure 1993 Raup & Jablonski, 1991 Thayer, Rhodes & 1986 & Hansen, Sheehan 2003 Stilwell, 2007 & Marquet, Rivadeneira 1986a Stanley, 1995 & Raup, Jablonski 1996a Jablonski, Levinton, 1973 & Bohonak, 1995 McClure 1993 Raup & Jablonski, 1993 & Jablonski, Valentine larva Plank approaches; = multivariate Multivariate = morphological variation; var com = morphological complexity; Morph rate = metabolic rate; Morph Meta = sensitivity to hypercapnia; Hypercapnia = taxonomic structure Tax feeding = suspension feeding; range = stratigraphic range; Susp Strat = Stenothermal; richness = species richness; Stenotherm = planktonic larva;thick shell thickness; Sp Shell structure. likely to go extinct); ? = unclear whether this pattern is due extinction selectivity. more these traits are Rhaet = = Recent; Rec Pz = Paleozoic; = Pliocene; Plio = Pliensbachian; Pliens = Pleistocene; Pleist = Phanerozoic; extinction; Phan P/T = end-Permian = Permian; Perm O/S = end-Ordovician; Norian; Triassic. = Tri Toarcian; = Toar extinction; T/J = end-Triassic Rhaetian; Trait Stenotopic range Strat feeding Susp feeding Susp feeding Susp feeding Susp feeding Susp feeding Susp feeding Susp feeding Susp feeding Susp feeding Susp feeding Susp feeding Susp feeding Susp feeding Susp feeding Susp structure Tax Tropical depth Water depth Water depth Water depth Water depth Water depth Water Key: range = geographic range; depth; Escalation = escalated traits; Geo depth = burrowing range = bathymetric range; Bur locomotion; Aragonite = aragonitic shell composition; Bathy locom = active Active Traits: of selectivity (i.e., these taxa or with higher levels Pos = positive less likely to go extinct); NS = not significant; of these traits are selectivity (i.e., these taxa or with higher levels = negative Neg Pattern: moll = mollusks; superfm brach = brachiopods; dom dominated; gast gastropods; = superfamilies. ammon = ammonites; biv bivalve; Taxon: fam = families; gen genera; sp species. level: Taxon = Nor Ng = Neogene; = Miocene; extinction; Mio K/Pg = end-Cretaceous = Jurassic; extinction; Jur extinction; F/F = late Devonian E/O = end-Eocene = Eocene; Eo = Cenozoic; Cz = Cretaceous; Age: Cret West. W = = tropical; trop S = South; Zealand; NZ = New N = North; hemis = hemisphere; = Europe; E = East; Euro Carib = Caribbean; CP Coastal Plain; = Atlantic; Location: Am = America; Atl dep = deposit; susp suspension. Notes: