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Direct PNAS a is article This interest. of conflict no declare paper. authors the The wrote S.E.P. and S.F., A.Z., and data; analyzed observed drive to inde- unlikely studies, be are should biodiversity and patterns. Other (11), paleogeography fossil (34). assignments of taxonomic in pendent rock incorrect marine concerns as describ- American quality such series North data time of common independent amount bio- alterna- an the and abundance this with ing paleogeography assess the series our To time in compare 29–33). diversity also fossil variation we (15, in temporal hypothesis, rock bias tive to marine fossil sampling related preserved between a of is correlation reflect that could observed data tectonics an and Therefore, that (25–28). diversity conceivable sedimentation is of patterns it regional on trol uhrcnrbtos ..adSEP eindrsac;AZ efre eerh A.Z. research; performed A.Z. research; designed S.E.P. and S.F. contributions: Author owo orsodnesol eadesd mi:[email protected]. Email: addressed. be should correspondence whom To aieaia iest vrteps 4 ilo er is years million supercon- 443 the Pangaea. of past tinent disassembly and the assembly the of over to attributable trajectory diversity the animal of component continen- marine significant test shifting A to hypothesis. of reconstructions this paleogeographic analyses in quantitative configurations tal and global the distributed data that globally in use fossil shifts hypothesized We crust. driven been continental tectonically of has arrangement by influenced It is biology. biodiversity of cen- goal a is tral biodiversity govern that processes the Understanding Significance ee eepiil ettepaetcoi euainhypothesis regulation tectonic plate the test explicitly we Here, lbltcoisi lopeitdt xr rtodrcon- first-order a exert to predicted also is tectonics Global b eateto nertv ilg,Uiest fCalifornia, of University Biology, Integrative of Department paleobiodb.org PNAS | a 0 2017 30, May (24). ) . 201702297 | o.114 vol. www.pnas.org/lookup/suppl/doi:10. ). | o 22 no. | 5653–5658

EARTH, ATMOSPHERIC, ECOLOGY AND PLANETARY SCIENCES Results motions persist, the index will continue to decline as the last rem- Paleogeography. We developed an index to quantitatively nant of the ancient Tethys Sea, the Mediterranean, is eliminated describe the degree of continental fragmentation during the by the collision of Africa with Eurasia. However, opening of the Phanerozoic (Materials and Methods). The index ranges between East African rift may partially offset this decrease. zero and 1, with 1 representing the state in which all continental blocks are completely separated, and zero representing the state Global Biodiversity. Global biodiversity estimates used here in which all continental blocks are contiguous and arranged in a derive from the Paleobiology Database and are broadly compa- way that minimizes their ocean basin-facing perimeter. We cal- rable to other assessments of marine animal richness (12, 14). culate the index only for those continental blocks that are present The specific Phanerozoic trajectory of marine animal genus rich- throughout the entire duration of the Phanerozoic. Together, ness remains contested (11, 13, 15–17, 21), but analyses consis- these persistent continental fragments comprise 88% of total tently reproduce several long-term patterns. A quick rise to a present-day continental area in the EarthByte model (23). volatile Paleozoic plateau (19), a drop during the end-Permian, The continental fragmentation index (Fig. 1) increases at the a prolonged Early Mesozoic recovery, and a subsequent rise (9, start of the Phanerozoic (541 Ma) in response to the breakup and 20–22) that eventually exceeded the Paleozoic maximum in the dispersal of crustal blocks that previously formed the Proterozoic Late Mesozoic or Early Cenozoic (Fig. 2 A and B) are all consis- supercontinent of Rodinia (35). Peak Paleozoic continental dis- tent features of the paleontological literature. assembly occurs at ∼470 Ma, near the Early/Middle Ordovician There are a number of qualitative similarities between the boundary. A series of alternating plateaus and declines follows trajectory of Phanerozoic biodiversity (Fig. 2) and continental the Late Ordovician peak in continental fragmentation, reflect- fragmentation (Fig. 1), which is consistent with the original for- ing the progressive reaggregation of continental blocks, which mulations of the hypothesis (1). These similarities are quanti- ultimately culminated in the formation of the Phanerozoic super- tatively expressed by a positive rank–order correlation between continent Pangaea. continental fragmentation and global genus richness (range- The most notable feature of the Mesozoic portion of the frag- through ρ = 0.40; in-bin ρ = 0.51). The correlation improves if the mentation index is a steady rise that begins in the mid-Jurassic Cambrian–Ordovician, which witnessed the initial evolution and and culminates in a Phanerozoic maximum during the Late Cre- diversification of marine animal communities, is removed from taceous. This protracted increase in continental fragmentation the analysis (range-through ρ = 0.64; in-bin ρ = 0.57). The impro- reflects the breakup of Pangaea and, notably, the southern conti- vement in correlation is not because the Cambrian–Ordovician nent of Gondwana. Today, Gondwana is represented by the dis- exhibits a pattern contrary to the hypothesis; it is an interval of tinct continents of South America, Africa, Antarctica, and Aus- rising biodiversity coincident with supercontinent breakup (20, tralia, as well as the subcontinent of India (36). The marked 37, 38). However, the Cambrian–Ordovician deviates from the decrease in the continental fragmentation index during the Late overall pattern in that the magnitude of fragmentation is rel- Cenozoic reflects the collision of dispersed Gondwanan conti- atively small (Fig. 1), but biodiversity accumulates at an un- nental blocks with Northern Hemisphere continents. This tran- usually high rate relative to the rest of the Phanerozoic (Fig. 2A). sition is most dramatically represented by the isolation of India To explore the impact that individual time intervals have on during peak continental fragmentation and its mid-Cenozoic col- the overall strength of the correlation between genus richness lision with Asia to form the Himalayas. If present-day plate and continental fragmentation (Fig. 2 C and D), we performed fragmentation index

Cm Ord S Dev Carb Perm Tr Jur K Pg Ng

0.30 0.35500 0.40 0.45 0.50 0.55 400 300 200 100 0 millions of years ago

Fig. 1. An index of continental block fragmentation derived from the EarthByte paleogeographic reconstruction models calculated in million-year incre- ments. An index value of unity indicates no plates are touching; an index value of zero corresponds to the state that would be achieved if all of continental blocks were contiguous and arranged in a single mass with a minimal ratio of perimeter to area. Example EarthByte-derived configurations (23) for the Early/Middle Ordovician (470 ma), the Middle/Late Triassic (237 ma), and Paleocene/Eocene (56 ma) are shown for visual reference. Smaller plates that do not persist throughout the entire Phanerozoic are not used in the index calculations and are grayed out in the inset maps. Carb, Carboniferous; Cm, Cambrian; Dev, Devonian; Jur, Jurassic; K, Cretaceous; Ng, Neogene; Ord, Ordovician; Perm, Permian; Pg, Paleogene; S, Silurian; Tr, Triassic.

5654 | www.pnas.org/cgi/doi/10.1073/pnas.1702297114 Zaffos et al. Downloaded by guest on September 30, 2021 Downloaded by guest on September 30, 2021 eprldnmc fterltosi.Frt epromda performed we First, relationship. the of dynamics temporal and fragmentation weaken continental strongly richness. not richness between marine does correlation low points However, overall data relatively Pangaea. these the of of supercontinent removal interval the the with long even relation- concurrent a the is record of that drivers they important as most Triassic–Early ship, final the the before are rela- Conversely, stages epochs with 1). last Jurassic (Fig. trend, the Pangaea during overall of 2A) the coalescence (Fig. from richness high most epoch (Early tively Cisuralian the single The departs no 3A). (Fig. Permian) Indeed, correlation effect. observed the quantitative the drives from little Pliocene–Pleistocene be the has may removing analysis that this find (39); we compilations has but data fundamen- so, fossil it is in Pliocene–Pleistocene example, undersampled the tally For bio- that between fragmentation. hypothesized correlation been continental overall con- the and the to without diversity assess epoch recalculated specifically each to correlation of attempts tribution the analysis This and removed data. series successively those time were epochs the which from in analysis jackknife a Ord, Triassic. Neogene; Tr, Silurian; Ng, S, Cm, Cretaceous; Paleogene; Carboniferous; Pg, K, Permian; Carb, Perm, Jurassic; points). Ordovician; Jur, gray Devonian; (light Dev, Ordovician Cambrian; and points) gray s h rgetto ne;teidxvlefrteitra ipitis midpoint interval the for value index the Spearman’s richness shown. index; generic fragmentation in-bin of the Scatterplot vs. (D) duration index. interval fragmentation make intervals. continental to confidence (C introduced 95% balanced. epochs are more custom points identify 1). on circles Bars quo- Black (Fig. (18). shareholder using (SQS) index standardized increments. subsampling richness rum million-year fragmentation genus in in-bin calculated continental Epoch-level, (B) richness the generic Range-through with (A) relationships and 2. Fig. afse al. et Zaffos range-through genus richness in-bin genus richness range-through genus richness at ecnutdbsctm eisaaye oeaiethe examine to analyses series time basic conducted we Last, 0 500 1000 1500 200 300 400 500 600 500 1000 1500 C B A mOdSDvCr emT u K Jur Tr Perm Carb Dev S K Ord Cm Jur Tr Perm Carb Dev S Ord Cm 0 0 0 0 0 0 100 200 300 400 500 .504 .505 0.55 0.50 0.45 0.40 0.35 lbltm eiso kltnzdmrn eu richness genus marine skeletonized of series time Global fragmentation index ctepo frnetruhgnrcrcns s the vs. richness generic range-through of Scatterplot ) ρ ausin values ρ =0.64 C millions ofyearsago and D in-bin genus richness onticueteCmra (dark Cambrian the include not do 0 100 200 300 400 500 600 D .504 .50.50 0.45 0.40 0.35 fragmentation index ρ =0.57 gNg Pg Ng Pg inidxadgnsrcns hntedt r transformed are data the Second, (ρ when differences (1–7). richness first fragmenta- taking years genus continental by and of the index between millions correlation tion of no of tens observe rearrangements we by presage crust should biodiversity continental consis- reason in intuitive direction no changes is the there why expectations; in model correlation general in with case, statisti- continued run tent this of observe In to spurious. only correlations indicator are for we relationships an when common is directions is lag Asymmetry it both because S1). robustness (Fig. stage cal the used at comparable is biodiversity and subsampled level if asymmetry apparent of the also pattern as is lag-times zero same toward The fragmen- steadily the increases. continental declines that lag of correlation such the state reversed, millions then is the tation, of direction precedes lag tens biodiversity the several of If state by 3B). tectonics (Fig. to years time of relative biodiversity the lagged correlation if is even The comparable series remains tem- versa. it a but without vice lag, high poral is and fragmentation and tectonics richness genus to response between relative biological the lagged when is frag- biodiversity supercontinent global of and similarity mentation the assess to cross-correlation hsrsl scnitn ihtehptei htapicplmode principal a that (ρ hypothesis the with index consistent correlated is fragmentation result also This is continental (34) continent the the with on rocks marine of quantity Record. longer Rock expecta- Marine over model changes general only with configuration timescales. consistent continental is as timescales. tions, too, million-year shorter, finding, is on contrast, This richness series In fragmen- continental marine trends. time between and long-term relationship raw tation their observed the no of is in similarity there correlation the by observed driven the that cates in em ema;P,Ploee ,Slra;T,Triassic. Tr, Silurian; S, Paleogene; Ordovi- Pg, Ord, Permian; Neogene; Perm, Ng, Cam- cian; Cretaceous; K, Cm, Jurassic; Spearman’s Jur, Carboniferous; Devonian; series. Carb, Dev, time brian; relationships. the post-Ordovician lagging only when denote rock marine and (C index series. lag- time when richness the global ging and index fragmentation the 2σ between removed represent correlation is lines epoch Dotted the series. geologic time between that the if correlation from richness the marine represents and point index Each fragmentation (A) biodiversity. and ics 3. Fig. spearman's ρ spearman's ρ 0.0 0.2 0.4 0.6 0.0 0.2 0.4 0.6 0.8 1.0 B A tectonics lagsbiology mOdSDvCr emT u K Jur Tr Perm Carb Dev S Ord Cm 0 0 0 0 0 0 100 200 300 400 500 vdnefrln-vru hr-emcvrainbtentecton- between covariation short-term versus long- for Evidence 5 50 0 -50 lag timeinmillionsofyears biology lagstectonics hnigcreainbtentefragmentation the between correlation Changing ) PNAS nidpnettm eisdsrbn the describing series time independent An | millions ofyearsago a 0 2017 30, May 0 = .01, spearman's ρ 0.0 0.2 0.4 0.6 P C tectonics lagsrock 0 = rmtema.(B mean. the from | 5 50 0 -50 .Ti nigindi- finding This .79). o.114 vol. lag timeinmillionsofyears 0 = | rock lagstectonics . i.3C). Fig. 33; o 22 no. Changing ) ρ gNg Pg | values 5655

EARTH, ATMOSPHERIC, ECOLOGY AND PLANETARY SCIENCES of variability in the amount of shallow marine sedimentation in biodiversity is concurrent with peak-Phanerozoic fragmenta- occurring on the continents is attributable to the superconti- tion following the breakup of Pangaea. nent coalescence–breakup cycle (28). However, this result also Evidence that continental coalescence phases cause declines in raises the possibility that there is a rock quantity-related sam- biodiversity is less robust. The latter half of the Paleozoic records pling bias in the fossil data (15, 29–33), and the observed correla- a prolonged period of plate collision to form Pangaea (Fig. 1), tion between continental fragmentation and biodiversity instead but this interval of supercontinent assembly is not matched by a reflects a confounded relationship with marine rock quantity. corresponding decrease in marine richness (Fig. 2). Indeed, it has However, the raw time series describing the quantity of marine been argued that richness was on a slight upward trend during the rock is uncorrelated with global marine richness (ρ = 0.01). Only Late Paleozoic (48). Continental fragmentation has decreased when the marine richness and rock quantity time series are since reaching a peak during the Late Cretaceous. Although bio- detrended by a first-differences transformation does any corre- diversity also appears to be in a declining phase (Fig. 2), it is lation emerge (ρ = 0.22, P  0.01). not clear to what extent this decline reflects an actual downturn Together, these analyses indicate that there are fundamentally in biodiversity (22, 49) versus idiosyncratic paleontological sam- different temporal patterns of correlation among the three time pling of the tropics (39). series of continental fragmentation, marine sedimentation on The suggestion that continental fragmentation and coal- the continents, and global biodiversity. Genus richness remains escence establish different biodiversity regimes has several impli- correlated with continental fragmentation when the biodiver- cations for understanding the history of Phanerozoic marine sity time series is lagged relative to the fragmentation index by richness. First, it implies that the Late Permian–mid-Jurassic bio- tens of millions of years, but there is no comparable lagged diversity minimum was initiated by the end-Permian mass extinc- relationship between marine rock and continental fragmenta- tion (50), and the subsequent recovery was prolonged because tion (Fig. 3 B and C). Furthermore, marine richness and North the world was in a coalescence regime. Without the end-Permian American sedimentation are correlated when they are detrended mass extinction, the Paleozoic biodiversity plateau (51) would by taking first differences, but there is no correlation between likely have persisted until the start of the next fragmentation continental fragmentation and biodiversity when the data are regime. Second, if coalescence does serve to stabilize biodiver- detrended. The opposing nature of these results is not consis- sity, then the increase in genus richness observed during the tent with a confounded three-way relationship in which rock Cenozoic will likely slow or cease sometime in the relatively near record-controlled sampling biases have driven a spurious corre- future (22, 49). A new interval of plate aggregation has been lation between marine richness and continental fragmentation. underway since the Late Cretaceous; this also suggests that any There is also no evidence to support the hypothesis that the long- downturn in present-day taxonomic richness (52) may be fol- term trajectory of Phanerozoic marine richness is an artifact of lowed by a protracted recovery phase that is more analogous to increasing rock quantities (11, 12, 14, 29, 30). Results instead the Triassic than it is to the Paleogene, at least from the per- suggest that only a small percentage of variance in biodiversity spective of long-term plate tectonic boundary conditions. Finally, is explained by rock quantity, and only over short timescales (40, these results help to bridge long-debated, contrasting models for 41). Longer-term trends in biodiversity are instead well predicted Phanerozoic fossil diversity, namely diversity-dependent logistic by the supercontinent coalescence–breakup cycle. growth (12, 14, 51) versus nearly unbridled exponential increase (1, 21, 53, 54). Under our hybrid model, whether biodiversity is Discussion in an asymptotic or growth phase depends on the current paleo- geographic continental fragmentation–coalescence regime. The history of marine biodiversity broadly mirrors patterns of The identification of paleogeographically mediated regimes supercontinent coalescence–breakup. This similarity is most dra- that control biodiversity does not preclude most alternative matically expressed by a pronounced Late Mesozoic rise in both hypotheses about long-term macroevolutionary mechanisms. For time series. This observation naturally raises a question (1–7): example, our results are consistent with models that empha- What aspects or correlates of continental crustal fragmentation size the importance of mass extinctions and recovery intervals are ultimately responsible for driving the trajectory of Phanero- in governing the history of life (55). Indeed, we attribute the zoic marine richness? Late Permian–Middle Jurassic biodiversity low to a coupling of The creation and destruction of intercontinental barriers to the end-Permian mass extinction and a continental coalescence biological dispersal was originally envisioned as a primary causal regime. This view is also compatible with many hypotheses that connection between tectonically driven shifts in paleogeogra- are primarily biotic in nature. Changing productivity patterns (8), phy and global biodiversity (1–7). Under this model, continen- coevolutionary dynamics (9), latitudinal biodiversity gradients tal fragmentation results in the formation of geographic barri- (48), incumbency and invasibility dynamics (56), and morpho- ers that separate formerly well-mixed faunas. Vicariance then logical/ecological character displacement (57) may all be facil- leads to evolution of new lineages and contributes to an over- itated or triggered by changing geographic connectivity. Even all increase in global taxonomic richness. However, eustatic sea biotic hypotheses that do not have an obvious connection to the level (3, 42–45) and ocean–atmospheric circulation (e.g., carbon geographic arrangement of continental crust, such as the emer- cycling, global climate) (3, 38, 46, 47) can also be directly or indi- gence of sexual characteristics (22), are, at worst, independent rectly affected by shifts in the geographic arrangement of conti- of the tectonic dynamics presented here and should not be con- nental crust. Because each of these tectonically linked processes sidered mutually exclusive. Nevertheless, our results do provide has been invoked separately as a driver of global biodiversity, it compelling evidence that the shifting distribution and connectiv- is difficult to identify a single causal mechanism. ity of continental landmass, either as the primary forcer or as a The correlations documented here are driven primarily by complement to other macroevolutionary processes, has been a broad temporal alignment of Phanerozoic highs and lows in fundamental driver of long-term global Phanerozoic biodiversity biodiversity and continental fragmentation. Similarities are par- patterns. ticularly well expressed during intervals of increasing conti- nental fragmentation and increasing biodiversity. Continental Materials and Methods fragmentation-forced increases in biodiversity have been posited Paleogeographic Reconstruction. Paleocontinent geometries and posi- in one form or another for several different times in Earth history tions were obtained from a curated version of an EarthByte (earthbyte.org) (20, 37, 38). The Cambrian–Ordovician is coincident with the tectonic plate reconstruction model (23). Modifications involved the correc- final fragmentation of Rodinia, and the Late Mesozoic increase tion of basic geometry errors (e.g., crossing plate polygon vertices) and the

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Global (1972) EM Moores JW, diver- Valentine marine Phanerozoic of 6. model provincial A an (1978) endemism, D Peart and TC, Foin diversity JW, Valentine marine shallow 5. and tetonics Plate (1971) JW Valentine 4. speciation con- in of climate and eustasy rate-control movements, plate of the importance Relative (1981) for A Hallam theory 3. nonequilibrium A (1982) J sea Cracraft and diversity 2. faunal of regulation Plate-tectonic (1970) EM Moores JW, Valentine 1. eew s h ubro nqegnr,ie,gnsrcns,a our as richness, genus i.e., genera, unique of number the use we Here h aegorpi eosrcin o ahtm nevlwr summa- were interval time each for reconstructions paleogeographic The aiediversification. marine 26:53–73. biotas. grazers. and ecosystem. marine the in ductivity and Colloquium, Biology Annual Papers Thirty-seventh Selected the of Proceedings Environment: ing 167–184. sity. model. actualistic Critique Mesozoic. A Early raphy: the since changes biogeographical major patterns. trolling macroevolutionary of origin the 365. and extinction and model. A level: Paleobiology brates. itn h el ftpooi,lttdnl n niomna biases. environmental and latitudinal, taphonomic, 625–642. of veils the Lifting studies. palaeobiodiversity for important are Paleobiology Paleobiology Science Paleobiology 35:51–62. 321:97–100. Nature Oeo tt nvPes ovli,OR). Corvallis, Press, Univ State (Oregon Clmi nvPes e ok,p 303–330. pp York), New Press, Univ (Columbia ytBiol Syst 4:55–66. 35:612–630. rcNt cdSiUSA Sci Acad Natl Proc 228:657–659. osldt dwlae aur 6 07 comes 2017) 16, January (downloaded data Fossil itrclBoegah,PaeTcois n h Chang- the and Tectonics, Plate Biogeography, Historical 3:245–258. 20:253–264. Paleobiology paleobiodb.org elScSe Publ Spec Soc Geol 98:6261–6266. 19:372–397. a n “min and ma” aasrie(24). service data ) 358:1–7. iaineBiogeog- Vicariance ytBiol Syst a fields, ma” Paleobiology Geol J Geol J 31:348– 112: 80: 3 mt B coa J(07 h hp ftePaeoocmrn palaeodiversity marine Phanerozoic the of shape The (2007) AJ McGowan reinterpretation. AB, A for Smith phanerozoic: Implications the 33. in record: Biodiversity (2001) fossil M Foote the SE, of Peters heterogeneity 32. Large–scale (2001) AB Smith interpretation. 31. An phanerozoic: the in diversity Species (1976) phanerozoic. DM sedimenta- the Raup American during diversity North Taxonomic 30. in (1972) rhythm DM Raup year million 29. 56 A (2011) SE the Peters for SR, trigger Meyers a as Unconformity’ 28. ‘Great the of Formation (2012) RR Gaines America. SE, North Peters of Macrostratigraphy 27. (2006) America. SE North Peters of interior 26. cratonic the in Sequences (1963) L Sloss programming application 25. database paleobiology The (2016) M McClennen SE, Peters 24. of dynamics M biodiversity S, shifting Zahirovic the N, and Wright Sex 23. (2016) RK Bambach G, Hunt AM, Bush marine Mesozoic- of 22. diversification and Mesozoic–Cenozoic Sustained Ordovician (2015) RK the Bambach AM, and Bush Palaeobiogeography 21. (2002) fossil J marine Phanerozoic Crame the A, of Owen description analytic 20. factor A (1981) Jr JJ, Sepkoski orig- of 19. estimation unbiased and richness taxonomic of sampling Fair (2010) The J curves: Alroy diversity 18. from bias Removing (2004) CR Marshall MJ, Markey AM, Bush 17. onainGatER1502(oSEP) hsi aebooyDatabase Paleobiology thank is Science This also National S.E.P.). for by We (to supported Gurnis 282. edi- 1150082 primarily Publication services. the EAR M. was and Grant work referees, rotation and anonymous This Foundation two comments. GPlates Turner Scalfani, for J. M. tor initial Marshall, M. and establishing Christie, M. infrastructure, in data help Macrostrat to ACKNOWLEDGMENTS. (10 sequences stratigraphic fourth-order packages” or 10 “sediment to the third- to unconformity-bound of comparable 4,585 portion are into that American grouped North units, marine are the stratigraphic 17,996 which to include data analysis the restricted, record Thus there- database. rock We in America. our global North limit for are only fore data complete some is Although coverage 62). Macrostrat’s scope, 61, pub- (26, from derived columns units stratigraphic geologic lished of attributes temporal and spatial, mental, Rock. 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(3, shifts this long-term level in be of biases sea cannot sampling changing (48), potential continents as as of only such distribution viewed tectonics, latitudinal completely changing to of the a linked or nature are 42–45) with the that governing correlations record factors fossil systematic other the analysis yield Moreover, our series. to time in as independent errors so other combine or should sampling, variable geographically practice, diversity. defini- single fossil a Phanerozoic environmental not of is and history there Consequently, biological biodiversity tive 40). real (21, distort estimates remove standardiza- the that from even size signals idiosyncrasies possibly sample or other some (17), that introduce estimates noted can capacity been methods some has in tion it in-bin, 18) However, on 14, 2B). rely (12, (Fig. instead estimates and biodiversity 2A), standardized (Fig. sampling estimates richness range-through use Accuracy. Analytic eetees hr r e roiraost eiv httaxonomic that believe to reasons priori a few are there Nevertheless, uv:Hwmc a epeitdfo h eietr okrcr fWestern of record rock sedimentary the from predicted be Europe? can much How curve: biology studies. biodiversity phanerozoic 2:289–297. Phanerozoic. the during tion explosion. Cambrian 74:93–114. interface. tectonics. plate with data paleobiol- ogy and paleogeographic open-access Integrating reconstructions: geographic time. deep in animals marine record. fossil the from signal consistent A metazoa: radiations. biotic Cenozoic record. Papers Society rates. extinction and ination sampling-standardization. on 30:666–686. biodiversity organized spatially of effects 7 ydrto) l aaaeacsil i h arsrtdt service. data Macrostrat the via accessible are data All duration). -y Paleobiology 27:583–601. Palaeontology Paleobiology PlotlSc ehsa D,Vl1,p 55–80. pp 16, Vol MD), Bethesda, Soc, (Paleontol h arsrtdtbs eod h ihlgc environ- lithologic, the records database Macrostrat The Nature 7(1):36–53. otrcn idvriyaaye yial onot do typically analyses biodiversity recent Most 42:1–7. 50:765–774. PNAS etakJ zpesifrciia contributions critical for Czaplewski J. thank We ile ,StnM(03 oad omnt-rvnpaleo- community-driven Towards (2013) M Seton R, Aijller ˜ elScSe Publ Spec Soc Geol 484:363–366. uniaieMtosi aebooy Paleontological Paleobiology, in Methods Quantitative at lntSiLett Sci Planet Earth rcNt cdSiUSA Sci Acad Natl Proc Biogeosciences | hlsTasRScLno e B Ser London Soc R Trans Philos a 0 2017 30, May 10:1529–1541. 194:1–11. 303:174–180. | Geology 113:14073–14078. Geol J o.114 vol. 114:391–412. 43:979–982. Science | 356:351–367. elScA Bull Am Soc Geol o 22 no. 177:1065–1071. Paleobiology Paleobiology | Paleo- 5657 5 -

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