Marine

In this paper a methodology is presented for measuring diversity based on rarefaction of actual samples. By the use of this technique, a within-habitat analysis was made of the bivalve and polychaete components of soft-bottom marine faunas which differed in latitude, depth, temperature, and salinity. The resulting diversity values were highly correlated with the physical stability and past history of these environments. A stability-time hypothesis was invoked to fit these findings, and, with this hypothesis, predictions were made about the diversities present in certain other environments as yet unstudied. The two types of diversity, based on numerical percentage composition and on number of species, were compared and shown to be poorly correlated with each other. Our data indicated that species number is the more valid diversity measurement. The rarefaction methodology was compared with a number of diversity indexes using identical data. Many of these indexes were markedly influenced by sample size. Good agreement was found between the rarefaction methodology and the Shannon-Wiener information function.

@article{doi101086282541,
    author = "Sanders, Howard L.",
    title = "Marine Benthic Diversity: A Comparative Study",
    year = "1968",
    journal = "The American Naturalist",
    abstract = "In this paper a methodology is presented for measuring diversity based on rarefaction of actual samples. By the use of this technique, a within-habitat analysis was made of the bivalve and polychaete components of soft-bottom marine faunas which differed in latitude, depth, temperature, and salinity. The resulting diversity values were highly correlated with the physical stability and past history of these environments. A stability-time hypothesis was invoked to fit these findings, and, with this hypothesis, predictions were made about the diversities present in certain other environments as yet unstudied. The two types of diversity, based on numerical percentage composition and on number of species, were compared and shown to be poorly correlated with each other. Our data indicated that species number is the more valid diversity measurement. The rarefaction methodology was compared with a number of diversity indexes using identical data. Many of these indexes were markedly influenced by sample size. Good agreement was found between the rarefaction methodology and the Shannon-Wiener information function.",
    url = "https://doi.org/10.1086/282541",
    doi = "10.1086/282541",
    openalex = "W2067564219",
    references = "connell1961effects, doi101086282114, doi101086282398, doi101111j1469185x1965tb00815x, doi101111j155856461960tb03057x, doi101126science1473655250, doi1023071950746, doi1023072628, doi104319lo1960520138"
}

@article{doi101126science17740541065,
    author = "Raup, David M.",
    title = "Taxonomic Diversity during the Phanerozoic",
    year = "1972",
    journal = "Science",
    url = "https://doi.org/10.1126/science.177.4054.1065",
    doi = "10.1126/science.177.4054.1065",
    openalex = "W1968883957",
    references = "doi101038185749a0, doi10113000167606194960561dombig20co2, doi10113000167606196879273crosda20co2, doi101130spe89p93, doi1023071931976, doi105281zenodo16226412, openalexw2352235109, openalexw2596278645, openalexw2603991543, openalexw2614067154"
}

On the basis of about 70,000 species citations in the Zoological Record, it is estimated that about 190,000 fossil invertebrate species were described and named through 1970. The true figure may be higher because of incompleteness of the Zoological Record or lower because the estimate does not account for synonymy. About 70% of the species were described from USSR, Europe, and North America. About 42% are Paleozoic, 28% Mesozoic, and 30% Cenozoic. In the Cambrian part of the sample, 75% of the species are trilobites. In the Mesozoic and Cenozoic, about 70% are either molluscs or protozoans. When the data are normalized for absolute time, diversity (species per million years) shows a Paleozoic high in the Devonian which is approximately four-tenths of the Cenozoic level.

@article{doi101017s0094837300004917,
    author = "Raup, David M.",
    title = "Species diversity in the Phanerozoic: a tabulation",
    year = "1976",
    journal = "Paleobiology",
    abstract = "On the basis of about 70,000 species citations in the Zoological Record, it is estimated that about 190,000 fossil invertebrate species were described and named through 1970. The true figure may be higher because of incompleteness of the Zoological Record or lower because the estimate does not account for synonymy. About 70\% of the species were described from USSR, Europe, and North America. About 42\% are Paleozoic, 28\% Mesozoic, and 30\% Cenozoic. In the Cambrian part of the sample, 75\% of the species are trilobites. In the Mesozoic and Cenozoic, about 70\% are either molluscs or protozoans. When the data are normalized for absolute time, diversity (species per million years) shows a Paleozoic high in the Devonian which is approximately four-tenths of the Cenozoic level.",
    url = "https://doi.org/10.1017/s0094837300004917",
    doi = "10.1017/s0094837300004917",
    openalex = "W2467392096",
    references = "doi101016s0016787876800077, doi101017s0094837300002633, doi101017s0094837300004929, doi101126science17740541065, doi1023072412725, doi1023072412728, openalexw1522518756, openalexw1523079602, openalexw2596278645, openalexw645218623"
}

Species diversity among fossil invertebrates of the Phanerozoic is highly correlated with volume and area of sedimentary rocks. The correlations are statistically significant at the 1% level. The relationship holds even in regions (such as Canada) where the area and volume of rock do not increase through time. These results are interpreted as indicating that the apparent number of species is strongly dependent on sampling and that many of the changes in diversity seen in the Phanerozoic are artifactual. Consequently, there is no compelling evidence for a general increase in the number of invertebrate species from Paleozoic to Recent. This conclusion applies primarily to marine organisms. Diversity may have been in dynamic equilibrium throughout much of this time. A few intervals of the Phanerozoic have consistently fewer invertebrate species than would be predicted from the amount of sedimentary rock available for study. The Silurian, Permian, and Cretaceous stand out in this regard. This may result either from lower than normal diversity during these periods or from an unusual abundance of unfossiliferous rocks (evaporites, red beds, etc.).

@article{doi101017s0094837300004929,
    author = "Raup, David M.",
    title = "Species diversity in the Phanerozoic: an interpretation",
    year = "1976",
    journal = "Paleobiology",
    abstract = "Species diversity among fossil invertebrates of the Phanerozoic is highly correlated with volume and area of sedimentary rocks. The correlations are statistically significant at the 1\% level. The relationship holds even in regions (such as Canada) where the area and volume of rock do not increase through time. These results are interpreted as indicating that the apparent number of species is strongly dependent on sampling and that many of the changes in diversity seen in the Phanerozoic are artifactual. Consequently, there is no compelling evidence for a general increase in the number of invertebrate species from Paleozoic to Recent. This conclusion applies primarily to marine organisms. Diversity may have been in dynamic equilibrium throughout much of this time. A few intervals of the Phanerozoic have consistently fewer invertebrate species than would be predicted from the amount of sedimentary rock available for study. The Silurian, Permian, and Cretaceous stand out in this regard. This may result either from lower than normal diversity during these periods or from an unusual abundance of unfossiliferous rocks (evaporites, red beds, etc.).",
    url = "https://doi.org/10.1017/s0094837300004929",
    doi = "10.1017/s0094837300004929",
    openalex = "W1506867616",
    references = "doi1010160037073876900282, doi101016b0080437516071036, doi101017s0094837300002633, doi101017s0094837300004917, doi101086282541, doi101126science17740541065, doi1015159781400881376, doi1023072412725, doi1023072412728, openalexw1523079602"
}

Raup's (1976a) data on Phanerozoic species numbers are examined for species-area relationships, using published estimates of areas of continental seas. By means of multiple regression, species numbers are regressed on both estimated areas of seas and amounts of available rock for sampling, as measured by outcrop area and rock volume. Although the sampling effects apparently have the strongest influence on fossil species diversity, areas of seas substantially increase the total correlation, suggesting that Phanerozoic species numbers were in equilibrium with habitat area. This is further supported by the fact that estimated parameters in the regressions are fairly consistent with established island biogeographic theory. Much of the remaining residual variation can be explained by periods of disequilibrium.

@article{doi101017s0094837300004930,
    author = "Sepkoski, J. John",
    title = "Species diversity in the Phanerozoic: species-area effects",
    year = "1976",
    journal = "Paleobiology",
    abstract = "Raup's (1976a) data on Phanerozoic species numbers are examined for species-area relationships, using published estimates of areas of continental seas. By means of multiple regression, species numbers are regressed on both estimated areas of seas and amounts of available rock for sampling, as measured by outcrop area and rock volume. Although the sampling effects apparently have the strongest influence on fossil species diversity, areas of seas substantially increase the total correlation, suggesting that Phanerozoic species numbers were in equilibrium with habitat area. This is further supported by the fact that estimated parameters in the regressions are fairly consistent with established island biogeographic theory. Much of the remaining residual variation can be explained by periods of disequilibrium.",
    url = "https://doi.org/10.1017/s0094837300004930",
    doi = "10.1017/s0094837300004930",
    openalex = "W2488451723",
    references = "doi1010160012825273900226, doi101017s0094837300004929, doi101086282738, doi101146annureves05110174001113, doi1015159781400881376, doi1023071269470, doi1023071931976, openalexw3035987306, openalexw623436458"
}

The distribution of numbers of species and the median number of species from 386 selected fossil communities are tabulated for high stress, variable nearshore, and open marine environments during the Lower, Middle, and Upper Paleozoic, the Mesozoic and the Cenozoic. The number of species always increases from high stress to variable nearshore to open marine environments. Within-habitat variation in number of species is small for long intervals of the Phanerozoic. The median number of species in communities from high stress environments remains fixed at about 8 from the Cambrian to the Pleistocene. In open marine environments, the median is near 30 for the Middle and Upper Paleozoic and almost the same for the Mesozoic. Increases of 50% in median number of species between the Lower and Middle Paleozoic and 2 times between the Mesozoic and Cenozoic occur in open marine environments with parallel, but less pronounced, increases in variable nearshore environments. Conditions controlling overall within-habitat species richness changed at those times. These changes do not correlate directly with evolution of new major taxa, change in physical conditions, predation, space availability or oxygen supply. They may be related to changes in resource availability influenced by factors such as the developing terrestrial flora, to lag-time inherent in the evolutionary process of diversification, or to as yet undetermined factors. Although provinciality determines total species richness for the biosphere, the within-habitat data suggest that the number of marine invertebrate species in the world has increased since the Middle Paleozoic, contrary to Raup's (1976b) contention, but possibly only by about 4 times, not the order of magnitude or more suggested by Valentine (1970).

@article{doi101017s0094837300005236,
    author = "Bambach, Richard K.",
    title = "Species richness in marine benthic habitats through the Phanerozoic",
    year = "1977",
    journal = "Paleobiology",
    abstract = "The distribution of numbers of species and the median number of species from 386 selected fossil communities are tabulated for high stress, variable nearshore, and open marine environments during the Lower, Middle, and Upper Paleozoic, the Mesozoic and the Cenozoic. The number of species always increases from high stress to variable nearshore to open marine environments. Within-habitat variation in number of species is small for long intervals of the Phanerozoic. The median number of species in communities from high stress environments remains fixed at about 8 from the Cambrian to the Pleistocene. In open marine environments, the median is near 30 for the Middle and Upper Paleozoic and almost the same for the Mesozoic. Increases of 50\% in median number of species between the Lower and Middle Paleozoic and 2 times between the Mesozoic and Cenozoic occur in open marine environments with parallel, but less pronounced, increases in variable nearshore environments. Conditions controlling overall within-habitat species richness changed at those times. These changes do not correlate directly with evolution of new major taxa, change in physical conditions, predation, space availability or oxygen supply. They may be related to changes in resource availability influenced by factors such as the developing terrestrial flora, to lag-time inherent in the evolutionary process of diversification, or to as yet undetermined factors. Although provinciality determines total species richness for the biosphere, the within-habitat data suggest that the number of marine invertebrate species in the world has increased since the Middle Paleozoic, contrary to Raup's (1976b) contention, but possibly only by about 4 times, not the order of magnitude or more suggested by Valentine (1970).",
    url = "https://doi.org/10.1017/s0094837300005236",
    doi = "10.1017/s0094837300005236",
    openalex = "W2148395366",
    references = "dayton1971competition, doi1010079783642659232, doi1010160031018268900473, doi101017s0094837300004930, doi101086282070, doi101086282400, doi101086282541, doi101086627723, doi101111j155856461963tb03295x, doi101130001676061968791315tailif20co2, doi101146annureves05110174001441, doi1015159781400881376, doi1023071948498, doi1023072258550, doi1023072485224, doi103133pp323, doi105281zenodo16238847, openalexw574363047"
}

@article{hallam1977secular,
    author = "Hallam, A.",
    title = "Secular changes in marine inundation of USSR and North America through the Phanerozoic",
    year = "1977",
    journal = "Nature",
    url = "https://doi.org/10.1038/269769a0",
    doi = "10.1038/269769a0",
    number = "5631",
    pages = "769-772",
    volume = "269"
}

@misc{hallam1977secular1,
    author = "Hallam, A",
    title = "Secular changes in marine inundation of USSR and North America during the Phanerozoic",
    year = "1977",
    howpublished = "Nature, v. 269, p. 769-772",
    note = "talkorigins\_source = {true}; raw\_reference = {Hallam, A., 1977, Secular changes in marine inundation of USSR and North America during the Phanerozoic: Nature, v. 269, p. 769-772.}"
}

A simple equilibrial model for the growth and maintenance of Phanerozoic global marine taxonomic diversity can be constructed from considerations of the behavior of origination and extinction rates with respect to diversity. An initial postulate that total rate of diversification is proportional to number of taxa extant leads to an exponential model for early phases of diversification. This model appears to describe adequately the “explosive” diversification of known metazoan orders across the Precambrian-Cambrian Boundary, suggesting that no special event, other than the initial appearance of Metazoa, is necessary to explain this phenomenon. As numbers of taxa increase, the rate of diversification should become “diversity dependent.” Ecological factors should cause the per taxon rate of origination to decline and the per taxon rate of extinction to increase. If these relationships are modeled as simple linear functions, a logistic description of the behavior of taxonomic diversity through time results. This model appears remarkably consistent with the known pattern of Phanerozoic marine ordinal diversity as a whole. Analysis of observed rates of ordinal origination also indicates these are to a large extent diversity dependent; however, diversity dependence is not immediately evident in rates of ordinal extinction. Possible explanations for this pattern are derived from considerations of the size of higher taxa and from simulations of their diversification. These suggest that both the standing diversity and the pattern of origination of orders may adequately reflect the behavior of species diversity through time; however, correspondence between rates of ordinal and species extinction may deteriorate with progressive loss of information resulting from incomplete sampling of the fossil record.

@article{doi101017s0094837300005972,
    author = "Sepkoski, J. John",
    title = "A kinetic model of Phanerozoic taxonomic diversity I. Analysis of marine orders",
    year = "1978",
    journal = "Paleobiology",
    abstract = "A simple equilibrial model for the growth and maintenance of Phanerozoic global marine taxonomic diversity can be constructed from considerations of the behavior of origination and extinction rates with respect to diversity. An initial postulate that total rate of diversification is proportional to number of taxa extant leads to an exponential model for early phases of diversification. This model appears to describe adequately the “explosive” diversification of known metazoan orders across the Precambrian-Cambrian Boundary, suggesting that no special event, other than the initial appearance of Metazoa, is necessary to explain this phenomenon. As numbers of taxa increase, the rate of diversification should become “diversity dependent.” Ecological factors should cause the per taxon rate of origination to decline and the per taxon rate of extinction to increase. If these relationships are modeled as simple linear functions, a logistic description of the behavior of taxonomic diversity through time results. This model appears remarkably consistent with the known pattern of Phanerozoic marine ordinal diversity as a whole. Analysis of observed rates of ordinal origination also indicates these are to a large extent diversity dependent; however, diversity dependence is not immediately evident in rates of ordinal extinction. Possible explanations for this pattern are derived from considerations of the size of higher taxa and from simulations of their diversification. These suggest that both the standing diversity and the pattern of origination of orders may adequately reflect the behavior of species diversity through time; however, correspondence between rates of ordinal and species extinction may deteriorate with progressive loss of information resulting from incomplete sampling of the fossil record.",
    url = "https://doi.org/10.1017/s0094837300005972",
    doi = "10.1017/s0094837300005972",
    openalex = "W2336833489",
    references = "bengtson1976the, doi1010160012825266900407, doi1010160012825272900724, doi101017s0094837300004917, doi101017s0094837300004929, doi101017s0094837300004930, doi101017s009483730000508x, doi101017s0094837300005224, doi101017s0094837300005236, doi101017s0094837300005649, doi101086282505, doi101086282762, doi101086627723, doi101093sysbio233305, doi101111j150239311969tb01258x, doi101111j150239311971tb01864x, doi101126science16238591265, doi101130gsab49195, doi101144gsjgs13130289, doi1015159780691206912, doi1015159781400881376, doi1023071218194, doi1023071483846, doi1023072412538, doi1023072412725, doi1023072412728, doi102475ajs2728752, doi102475ajs2748833, doi104095100784, doi104159harvard9780674865327, doi105281zenodo16238847, doi105962bhltitle66379, openalexw2145250129, openalexw2506868775, openalexw2601410785, openalexw3126336940, openalexw3135630760"
}

Much new empirical evidence on the levels of Phanerozoic paleoprovinciality and of species diversity within paleocommunities now permits a reevaluation of marine diversity patterns. Data on paleoprovincial patterns are assembled from the literature and evaluated by means of a stochastic computer simulation model. The simulation is based on the statistics of modern patterns of diversity and endemism extrapolated conservatively to the paleoprovincial patterns and on estimates of species duration from the fossil record. The species diversities associated with the paleoprovincial patterns are then corrected for temporal changes in species packing in communities as determined by Bambach (1977) from studies of paleocommunities. The model thus has an empirical basis throughout. Furthermore it is free of biases that can arise due to the differential preservation of taxa in space and time. The Paleozoic and Mesozoic were characterized by low provinciality and low average species diversity, on the order of 38,000 to 40,000 species, although there were significant fluctuations in standing diversities. In the Cenozoic, provinciality rose markedly, primarily through the appearance of latitudinal provincial chains, and average species diversity rose to about 240,000. Today it stands over 350,000; this is an order of magnitude greater than the Paleozoic average.

@article{valentine1978a,
    author = "Valentine, James W. and Foin, Theodore C. and Peart, David",
    title = "A provincial model of Phanerozoic marine diversity",
    year = "1978",
    journal = "Paleobiology",
    abstract = "Much new empirical evidence on the levels of Phanerozoic paleoprovinciality and of species diversity within paleocommunities now permits a reevaluation of marine diversity patterns. Data on paleoprovincial patterns are assembled from the literature and evaluated by means of a stochastic computer simulation model. The simulation is based on the statistics of modern patterns of diversity and endemism extrapolated conservatively to the paleoprovincial patterns and on estimates of species duration from the fossil record. The species diversities associated with the paleoprovincial patterns are then corrected for temporal changes in species packing in communities as determined by Bambach (1977) from studies of paleocommunities. The model thus has an empirical basis throughout. Furthermore it is free of biases that can arise due to the differential preservation of taxa in space and time. The Paleozoic and Mesozoic were characterized by low provinciality and low average species diversity, on the order of 38,000 to 40,000 species, although there were significant fluctuations in standing diversities. In the Cenozoic, provinciality rose markedly, primarily through the appearance of latitudinal provincial chains, and average species diversity rose to about 240,000. Today it stands over 350,000; this is an order of magnitude greater than the Paleozoic average.",
    url = "https://doi.org/10.1017/s0094837300005686",
    doi = "10.1017/s0094837300005686",
    number = "1",
    pages = "55-66",
    volume = "4"
}

The kinetic model of taxonomic diversity predicts that the long-term diversification of taxa within any large and essentially closed ecological system should approximate a logistic process controlled by changes in origination and extinction rates with changing numbers of taxa. This model is tested with a new compilation of numbers of metazoan families known from Paleozoic stages (including stage-level subdivisions of the Cambrian). These data indicate the occurrence of two intervals of logistic diversification within the Paleozoic. The first interval, spanning the Vendian and Cambrian, includes an approximately exponential increase in families across the Precambrian-Cambrian Boundary and a “pseudo-equilibrium” through the Middle and Late Cambrian, caused by diversity-dependent decrease in origination rate and increase in extinction rate. The second interval begins with a rapid re-diversification in the Ordovician, which leads to a tripling of familial diversity during a span of 50 Myr; by the end of the Ordovician diversity attains a new dynamic equilibrium that is maintained, except for several extinction events, for nearly 200 Myr until near the end of the Paleozoic. A “two-phase” kinetic model is constructed to describe this heterogeneous pattern of early Phanerozoic diversification. The model adequately describes the “multiple equilibria,” the asymmetrical history of the “Cambrian fauna,” the extremely slow initial diversification of the later “Paleozoic fauna,” and the combined patterns of origination and extinction in both faunas. It is suggested that this entire pattern of diversification reflects the early success of ecologically generalized taxa and their later replacement by more specialized taxa.

@article{doi101017s0094837300006539,
    author = "Sepkoski, J. John",
    title = "A kinetic model of Phanerozoic taxonomic diversity II. Early Phanerozoic families and multiple equilibria",
    year = "1979",
    journal = "Paleobiology",
    abstract = "The kinetic model of taxonomic diversity predicts that the long-term diversification of taxa within any large and essentially closed ecological system should approximate a logistic process controlled by changes in origination and extinction rates with changing numbers of taxa. This model is tested with a new compilation of numbers of metazoan families known from Paleozoic stages (including stage-level subdivisions of the Cambrian). These data indicate the occurrence of two intervals of logistic diversification within the Paleozoic. The first interval, spanning the Vendian and Cambrian, includes an approximately exponential increase in families across the Precambrian-Cambrian Boundary and a “pseudo-equilibrium” through the Middle and Late Cambrian, caused by diversity-dependent decrease in origination rate and increase in extinction rate. The second interval begins with a rapid re-diversification in the Ordovician, which leads to a tripling of familial diversity during a span of 50 Myr; by the end of the Ordovician diversity attains a new dynamic equilibrium that is maintained, except for several extinction events, for nearly 200 Myr until near the end of the Paleozoic. A “two-phase” kinetic model is constructed to describe this heterogeneous pattern of early Phanerozoic diversification. The model adequately describes the “multiple equilibria,” the asymmetrical history of the “Cambrian fauna,” the extremely slow initial diversification of the later “Paleozoic fauna,” and the combined patterns of origination and extinction in both faunas. It is suggested that this entire pattern of diversification reflects the early success of ecologically generalized taxa and their later replacement by more specialized taxa.",
    url = "https://doi.org/10.1017/s0094837300006539",
    doi = "10.1017/s0094837300006539",
    openalex = "W2504649407",
    references = "bretsky1968evolution, doi1010160012825272900724, doi101017s0094837300004917, doi101038260204c0, doi101086627723, doi101093aesa383396, doi101111j155856461963tb03295x, doi101144gsjgs13130289, doi101306m12367, doi1015159781400881376, doi10182618200093301197301, doi1023071483846, doi1023072405671, doi1023072412728, doi102475ajs2748833, doi104095100784, doi105281zenodo16238847, doi105962bhltitle4489, doi105962bhltitle66379, doi107312simp93764, openalexw2145250129"
}

A strong correlation exists between the outcrop area of nonmarine rocks deposited during a given geologic period and the observed vascular plant diversity for the same period; however, diversity residuals characteristic of certain periods may have underlying biological causes. Within-flora diversity changes through time indicate that stepwise increases in community species packing have accompanied major tracheophyte evolutionary innovations. Total and within-flora data suggest that the track of North American land-plant diversity has been similar in nature, but not in timing, to that inferred for marine invertebrates.

@article{knoll1979phanerozoic,
    author = "Knoll, Andrew H. and Niklas, Karl J. and Tiffney, Bruce H.",
    title = "Phanerozoic Land-Plant Diversity in North America",
    year = "1979",
    journal = "Science",
    abstract = "A strong correlation exists between the outcrop area of nonmarine rocks deposited during a given geologic period and the observed vascular plant diversity for the same period; however, diversity residuals characteristic of certain periods may have underlying biological causes. Within-flora diversity changes through time indicate that stepwise increases in community species packing have accompanied major tracheophyte evolutionary innovations. Total and within-flora data suggest that the track of North American land-plant diversity has been similar in nature, but not in timing, to that inferred for marine invertebrates.",
    url = "https://doi.org/10.1126/science.206.4425.1400",
    doi = "10.1126/science.206.4425.1400",
    number = "4425",
    pages = "1400-1402",
    volume = "206"
}

Data on numbers of marine families within 91 metazoan classes known from the Phanerozoic fossil record are analyzed. The distribution of the 2800 fossil families among the classes is very uneven, with most belonging to a small minority of classes. Similarly, the stratigraphic distribution of the classes is very uneven, with most first appearing early in the Paleozoic and with many of the smaller classes becoming extinct before the end of that era. However, despite this unevenness, a Q -mode factor analysis indicates that the structure of these data is rather simple. Only three factors are needed to account for more than 90% of the data. These factors are interpreted as reflecting the three great “evolutionary faunas” of the Phanerozoic marine record: a trilobite-dominated Cambrian fauna, a brachiopod-dominated later Paleozoic fauna, and a mollusc-dominated Mesozoic-Cenozoic, or “modern,” fauna. Lesser factors relate to slow taxonomic turnover within the major faunas through time and to unique aspects of particular taxa and times. Each of the three major faunas seems to have its own characteristic diversity so that its expansion or contraction appears as being intimately associated with a particular phase in the history of total marine diversity. The Cambrian fauna expands rapidly during the Early Cambrian radiations and maintains dominance during the Middle to Late Cambrian “equilibrium.” The Paleozoic fauna then ascends to dominance during the Ordovician radiations, which increase diversity dramatically; this new fauna then maintains dominance throughout the long interval of apparent equilibrium that lasts until the end of the Paleozoic Era. The modern fauna, which slowly increases in importance during the Paleozoic Era, quickly rises to dominance with the Late Permian extinctions and maintains that status during the general rise in diversity to the apparent maximum in the Neogene. The increase in diversity associated with the expansion of each new fauna appears to coincide with an approximately exponential decline of the previously dominant fauna, suggesting possible displacement of each evolutionary fauna by its successor.

@article{doi101017s0094837300003778,
    author = "Sepkoski, J. John",
    title = "A factor analytic description of the Phanerozoic marine fossil record",
    year = "1981",
    journal = "Paleobiology",
    abstract = "Data on numbers of marine families within 91 metazoan classes known from the Phanerozoic fossil record are analyzed. The distribution of the 2800 fossil families among the classes is very uneven, with most belonging to a small minority of classes. Similarly, the stratigraphic distribution of the classes is very uneven, with most first appearing early in the Paleozoic and with many of the smaller classes becoming extinct before the end of that era. However, despite this unevenness, a Q -mode factor analysis indicates that the structure of these data is rather simple. Only three factors are needed to account for more than 90\% of the data. These factors are interpreted as reflecting the three great “evolutionary faunas” of the Phanerozoic marine record: a trilobite-dominated Cambrian fauna, a brachiopod-dominated later Paleozoic fauna, and a mollusc-dominated Mesozoic-Cenozoic, or “modern,” fauna. Lesser factors relate to slow taxonomic turnover within the major faunas through time and to unique aspects of particular taxa and times. Each of the three major faunas seems to have its own characteristic diversity so that its expansion or contraction appears as being intimately associated with a particular phase in the history of total marine diversity. The Cambrian fauna expands rapidly during the Early Cambrian radiations and maintains dominance during the Middle to Late Cambrian “equilibrium.” The Paleozoic fauna then ascends to dominance during the Ordovician radiations, which increase diversity dramatically; this new fauna then maintains dominance throughout the long interval of apparent equilibrium that lasts until the end of the Paleozoic Era. The modern fauna, which slowly increases in importance during the Paleozoic Era, quickly rises to dominance with the Late Permian extinctions and maintains that status during the general rise in diversity to the apparent maximum in the Neogene. The increase in diversity associated with the expansion of each new fauna appears to coincide with an approximately exponential decline of the previously dominant fauna, suggesting possible displacement of each evolutionary fauna by its successor.",
    url = "https://doi.org/10.1017/s0094837300003778",
    doi = "10.1017/s0094837300003778",
    openalex = "W2505144080",
    references = "doi10100797814613088367, doi1010160012825272900724, doi101017s0094837300004917, doi101017s009483730000508x, doi101017s0094837300005236, doi101017s0094837300005352, doi101017s0094837300005649, doi101017s0094837300005972, doi101017s0094837300012549, doi101126science17740541065, doi101126science2064415217, doi101130spe89p63, doi1023071483846, doi1023071796560, doi1023072405671, doi1023072412725, doi1023072412728, doi1023072806339, doi107312simp93764, openalexw1504049102, openalexw645218623"
}

@article{doi101038293435a0,
    author = "Sepkoski, J. John and Bambach, Richard K. and Raup, David M. and Valentine, James W.",
    title = "Phanerozoic marine diversity and the fossil record",
    year = "1981",
    journal = "Nature",
    url = "https://doi.org/10.1038/293435a0",
    doi = "10.1038/293435a0",
    openalex = "W2002352667",
    references = "doi101017s0094837300003778, doi101017s0094837300004917, doi101017s0094837300004929, doi101017s0094837300004930, doi101017s0094837300005236, doi101017s0094837300005972, doi101017s0094837300006539, doi101111j150239311980tb00632x, doi101126science17740541065, doi1023071441916, doi1023072341482, openalexw645218623"
}

@article{hallam1981diversity,
    author = "Hallam, A.",
    title = "Diversity changes of marine organisms through the Phanerozoic",
    year = "1981",
    journal = "Nature",
    url = "https://doi.org/10.1038/293428a0",
    doi = "10.1038/293428a0",
    number = "5832",
    pages = "428-428",
    volume = "293"
}

@misc{sepkoski1981phanerozoic3,
    author = "Sepkoski, J. J. and Bambach, R. K. and Raup, D. M. and Valentine, J. W",
    title = "Phanerozoic marine diversity and the fossil record",
    year = "1981",
    howpublished = "Nature, v. 293, p. 435",
    note = "talkorigins\_source = {true}; raw\_reference = {Sepkoski, J. J., Bambach, R. K., Raup, D. M., and Valentine, J. W., 1981, Phanerozoic marine diversity and the fossil record: Nature, v. 293, p. 435.}"
}

Summary 1. Modes of larval development play important roles in the ecology, biogeography, and evolution of marine benthic organisms. Studies of the larval ecology of fossil organisms can contribute greatly to our understanding of such roles by allowing us to race effects on evolutionary time scales. 2. Modes of development can be inferred for well preserved molluscan fossils because the size of the initial larval shell (Protoconch I in gastropods, Prodissoconch I in bivalves) reflects egg size. Other morphological criteria are also available, and a comparative approach based on related taxa with known development may be the most reliable method. By combining larval and adult traits, it is possible to recognize modes of larval development in at least some fossil bryozoans, brachiopods, and echinoderms as well. (a) Planktotrophic larvae arise from small eggs, are released in enormous numbers with little parental investment per offspring, and suffer tremendous mortality during and shortly after a planktic existence. These larvae feed on the plankton during development, and are commonly capable of a prolonged free‐swimming existence, and thus wide dispersal. (b) Nonplanktotrophic larvae (which include both planktic lecithotrophic forms and ‘direct developers’) generally arise from large eggs, with relatively few young produced per parent. Relative to planktotrophic larvae, nonplanktotrophic larvae generally receive greater parental investment per larva, and larval mortality is generally lower. These larvae rely on yolk for nutrition during development, and planktic durations are generally much briefer than for species with planktotrophic larvae, so that dispersal capability is considerably less. Energetic investment per egg is generally higher than in planktotrophs, but as there are lower fecundities as well it is difficult to generalize about the total energetic cost of one mode of reproduction against the other. 3. Owing to the high dispersal capability of planktotrophic larvae, it has been suggested that species with such larvae will be geographically widespread, geologically long‐ranging, and exhibit low speciation and extinction rates. Species with nonplanktotrophic larvae will tend to be geographically more restricted, geologically short‐ranging, and exhibit high speciation and extinction rates (again, as a consequence of their characteristically low larval dispersal capabilities). 4. Recognition of differential dispersal capabilities can play a role in paleobiogeo‐graphic analyses. Concurrent study of the distribution of groups with contrasting modes of development will permit testing of hypotheses concerning timing, magnitudes and frequencies of migration and vicariance events. 5. Larval types are not randomly distributed in the oceans, but relationships with other aspects of the organisms' biology and habitats are very complex. Mode of development varies with: (a) Ecology. A simple r–––K model of adaptive strategies is clearly insufficient to explain the observed relationships: while many ‘equilibrium’ species have nonplanktotrophic larvae, and organisms living in less prdictable environments often have planktotrophic larvae, some of the most opportunistic marine species have nonplanktotrophic larvae. Nonetheless, planktotrophic development seems most suited for exploitation of patchy but widespread habitats. (b) Latitude. At shelf depths, planktotrophy is predominant in the tropics, and decreases sharply at high latitudes. (c) Depth. Incidence of planktotrophy decreases with depth across the continental shelf, at least in some taxa. Beyond the shelf, many deep‐sea organisms are nonplanktotrophic (e.g. most bivalves, peracarid crustaceans), but planktotrophic development appears to be present in other groups (prosobranch gastropods, ophiuroids, and bivalves inhabiting transient habitats such as sunken wood and hydrothermal vents). These trends in developmental types will be accompanied by trends in evolutionary rates and patterns as outlined above. The study of larval ecology by paleobiologists will yield insights into the processes that gave rise to ancient evolutionary and biogeographic patterns, and will permit the development and testing of hypotheses on the origins of the patterns observed in modern seas.

@article{jablonski1983larval,
    author = "JABLONSKI, DAVID and LUTZ, RICHARD A.",
    title = "LARVAL ECOLOGY OF MARINE BENTHIC INVERTEBRATES: PALEOBIOLOGICAL IMPLICATIONS",
    year = "1983",
    journal = "Biological Reviews",
    abstract = "Summary 1. Modes of larval development play important roles in the ecology, biogeography, and evolution of marine benthic organisms. Studies of the larval ecology of fossil organisms can contribute greatly to our understanding of such roles by allowing us to race effects on evolutionary time scales. 2. Modes of development can be inferred for well preserved molluscan fossils because the size of the initial larval shell (Protoconch I in gastropods, Prodissoconch I in bivalves) reflects egg size. Other morphological criteria are also available, and a comparative approach based on related taxa with known development may be the most reliable method. By combining larval and adult traits, it is possible to recognize modes of larval development in at least some fossil bryozoans, brachiopods, and echinoderms as well. (a) Planktotrophic larvae arise from small eggs, are released in enormous numbers with little parental investment per offspring, and suffer tremendous mortality during and shortly after a planktic existence. These larvae feed on the plankton during development, and are commonly capable of a prolonged free‐swimming existence, and thus wide dispersal. (b) Nonplanktotrophic larvae (which include both planktic lecithotrophic forms and ‘direct developers’) generally arise from large eggs, with relatively few young produced per parent. Relative to planktotrophic larvae, nonplanktotrophic larvae generally receive greater parental investment per larva, and larval mortality is generally lower. These larvae rely on yolk for nutrition during development, and planktic durations are generally much briefer than for species with planktotrophic larvae, so that dispersal capability is considerably less. Energetic investment per egg is generally higher than in planktotrophs, but as there are lower fecundities as well it is difficult to generalize about the total energetic cost of one mode of reproduction against the other. 3. Owing to the high dispersal capability of planktotrophic larvae, it has been suggested that species with such larvae will be geographically widespread, geologically long‐ranging, and exhibit low speciation and extinction rates. Species with nonplanktotrophic larvae will tend to be geographically more restricted, geologically short‐ranging, and exhibit high speciation and extinction rates (again, as a consequence of their characteristically low larval dispersal capabilities). 4. Recognition of differential dispersal capabilities can play a role in paleobiogeo‐graphic analyses. Concurrent study of the distribution of groups with contrasting modes of development will permit testing of hypotheses concerning timing, magnitudes and frequencies of migration and vicariance events. 5. Larval types are not randomly distributed in the oceans, but relationships with other aspects of the organisms' biology and habitats are very complex. Mode of development varies with: (a) Ecology. A simple r–––K model of adaptive strategies is clearly insufficient to explain the observed relationships: while many ‘equilibrium’ species have nonplanktotrophic larvae, and organisms living in less prdictable environments often have planktotrophic larvae, some of the most opportunistic marine species have nonplanktotrophic larvae. Nonetheless, planktotrophic development seems most suited for exploitation of patchy but widespread habitats. (b) Latitude. At shelf depths, planktotrophy is predominant in the tropics, and decreases sharply at high latitudes. (c) Depth. Incidence of planktotrophy decreases with depth across the continental shelf, at least in some taxa. Beyond the shelf, many deep‐sea organisms are nonplanktotrophic (e.g. most bivalves, peracarid crustaceans), but planktotrophic development appears to be present in other groups (prosobranch gastropods, ophiuroids, and bivalves inhabiting transient habitats such as sunken wood and hydrothermal vents). These trends in developmental types will be accompanied by trends in evolutionary rates and patterns as outlined above. The study of larval ecology by paleobiologists will yield insights into the processes that gave rise to ancient evolutionary and biogeographic patterns, and will permit the development and testing of hypotheses on the origins of the patterns observed in modern seas.",
    url = "https://doi.org/10.1111/j.1469-185x.1983.tb00380.x",
    doi = "10.1111/j.1469-185x.1983.tb00380.x",
    number = "1",
    openalex = "W2060347297",
    pages = "21-89",
    volume = "58",
    references = "doi101016b9780122825057x50015, doi101017s0094837300005224, doi101017s0094837300005236, doi101086282697, doi101086409052, doi101126science150369228, doi1015159781400881376, doi1023071483846, doi104159harvard9780674865327, openalexw2506868775, openalexw3135630760"
}

A three-phase kinetic model with time-specific perturbations is used to describe large-scale patterns in the diversification of Phanerozoic marine families. The basic model assumes that the Cambrian, Paleozoic, and Modern evolutionary faunas each diversified logistically as a consequence of early exponential growth and of later slowing of growth as the ecosystems became filled; it also assumes interaction among the evolutionary faunas such that expansion of the combined diversities of all three faunas above any single fauna's equilibrium caused that fauna's diversity to begin to decline. This basic model adequately describes the diversification of the evolutionary faunas through the Paleozoic Era as well as the asymmetrical rise and fall of background extinction rates through the entire Phanerozoic. Declines in diversity and changes in faunal dominance associated with mass extinctions can be accommodated in the model with short-term accelerations in extinction rates or declines in equilibria. Such accelerations, or perturbations, cause diversity to decline exponentially and then to rebound sigmoidally following release. The amount of decline is dependent on the magnitude and duration of the perturbation, the timing of the perturbation with respect to the diversification of the system, and the system's initial per-taxon rates of diversification and turnover. When applied to the three-phase model, such perturbations describe the changes in diversity and faunal dominance during and after major mass extinctions, the long-term rise in total diversity following the Late Permian and Norian mass extinctions, and the peculiar diversification and then decline of the remnants of the Paleozoic fauna during the Mesozoic and Cenozoic Eras. The good fit of this model to data on Phanerozoic familial diversity suggests that many of the large-scale patterns of diversification seen in the marine fossil record of animal families are simple consequences of nonlinear interrelationships among a small number of parameters that are intrinsic to the evolutionary faunas and are largely (but not completely) invariant through time.

@article{doi101017s0094837300008186,
    author = "Sepkoski, J. John",
    title = "A kinetic model of Phanerozoic taxonomic diversity. III. Post-Paleozoic families and mass extinctions",
    year = "1984",
    journal = "Paleobiology",
    abstract = "A three-phase kinetic model with time-specific perturbations is used to describe large-scale patterns in the diversification of Phanerozoic marine families. The basic model assumes that the Cambrian, Paleozoic, and Modern evolutionary faunas each diversified logistically as a consequence of early exponential growth and of later slowing of growth as the ecosystems became filled; it also assumes interaction among the evolutionary faunas such that expansion of the combined diversities of all three faunas above any single fauna's equilibrium caused that fauna's diversity to begin to decline. This basic model adequately describes the diversification of the evolutionary faunas through the Paleozoic Era as well as the asymmetrical rise and fall of background extinction rates through the entire Phanerozoic. Declines in diversity and changes in faunal dominance associated with mass extinctions can be accommodated in the model with short-term accelerations in extinction rates or declines in equilibria. Such accelerations, or perturbations, cause diversity to decline exponentially and then to rebound sigmoidally following release. The amount of decline is dependent on the magnitude and duration of the perturbation, the timing of the perturbation with respect to the diversification of the system, and the system's initial per-taxon rates of diversification and turnover. When applied to the three-phase model, such perturbations describe the changes in diversity and faunal dominance during and after major mass extinctions, the long-term rise in total diversity following the Late Permian and Norian mass extinctions, and the peculiar diversification and then decline of the remnants of the Paleozoic fauna during the Mesozoic and Cenozoic Eras. The good fit of this model to data on Phanerozoic familial diversity suggests that many of the large-scale patterns of diversification seen in the marine fossil record of animal families are simple consequences of nonlinear interrelationships among a small number of parameters that are intrinsic to the evolutionary faunas and are largely (but not completely) invariant through time.",
    url = "https://doi.org/10.1017/s0094837300008186",
    doi = "10.1017/s0094837300008186",
    openalex = "W2221600847",
    references = "doi1010079781475707403, doi1010160012825272900724, doi1010160031018281900924, doi101017s0094837300003778, doi101017s0094837300004917, doi101017s0094837300004929, doi101017s009483730000508x, doi101017s0094837300005236, doi101017s0094837300005352, doi101017s0094837300005649, doi101017s0094837300005972, doi101017s0094837300006539, doi101017s0094837300008174, doi101038260204c0, doi101038293435a0, doi101038303614a0, doi101073pnas722646, doi101073pnas813801, doi101086627905, doi101111j1469185x1983tb00380x, doi101111j150239311977tb00628x, doi101126science2064415217, doi101126science21545391501, doi101126science2164542173, doi101126science22246281123, doi101130spe89p63, doi1015159780691206912, doi102110pec77250019, doi1023071441916, doi1023072412725, jablonski1983larval, openalexw2145250129, openalexw2989049194"
}

@techreport{jablonski1986larval2,
    author = "Jablonski, D",
    title = "Larval ecology and macroevolution in marine invertebrates",
    year = "1986",
    howpublished = "Bulletin of Marine Science, v. 39, p. 565-587",
    note = "talkorigins\_source = {true}; raw\_reference = {Jablonski, D., 1986, Larval ecology and macroevolution in marine invertebrates: Bulletin of Marine Science, v. 39, p. 565-587.}"
}

A kill curve for Phanerozoic species is developed from an analysis of the stratigraphic ranges of 17,621 genera, as compiled by Sepkoski. The kill curve shows that a typical species' risk of extinction varies greatly, with most time intervals being characterized by very low risk. The mean extinction rate of 0.25/m.y. is thus a mixture of long periods of negligible extinction and occasional pulses of much higher rate. Because the kill curve is merely a description of the fossil record, it does not speak directly to the causes of extinction. The kill curve may be useful, however, to li¿mit choices of extinction mechanisms.

@article{doi101017s0094837300010332,
    author = "Raup, David M.",
    title = "A kill curve for Phanerozoic marine species",
    year = "1991",
    journal = "Paleobiology",
    abstract = "A kill curve for Phanerozoic species is developed from an analysis of the stratigraphic ranges of 17,621 genera, as compiled by Sepkoski. The kill curve shows that a typical species' risk of extinction varies greatly, with most time intervals being characterized by very low risk. The mean extinction rate of 0.25/m.y. is thus a mixture of long periods of negligible extinction and occasional pulses of much higher rate. Because the kill curve is merely a description of the fossil record, it does not speak directly to the causes of extinction. The kill curve may be useful, however, to li¿mit choices of extinction mechanisms.",
    url = "https://doi.org/10.1017/s0094837300010332",
    doi = "10.1017/s0094837300010332",
    openalex = "W1891084190",
    references = "valentine1978a"
}

Perturbations in ecosystems consist of a sequence of 2 events: the disturbance, marked by the application of the disturbing forces, and the response shown by the biota to the damage inflicted by the disturbance. The disturbance must be effectively characterized, without confounding it with the response, for progress to be made in the study of the disturbance ecology of streams. A disturbance may take the form of a pulse, a press, or a ramp, and the consequent trajectory of the response may be a pulse, a press, or a ramp.Floods and droughts are the major forms of natural disturbance in flowing waters and, although the effects of floods have been relatively well studied, those of droughts have been largely neglected. Floods accentuate downstream and lateral transport links, often with damaging consequences, whereas droughts fragment the continuity of streams. Both floods and droughts destroy and generate habitat patchiness and patchiness of the biota. During recovery, there are changes in the biotic composition and spatial configuration in patches. Resistance and resilience of the biota to disturbance may be facilitated by the use of refugia. The characterization of flood refugia is much more advanced than that of drought refugia.Recovery from floods is marked by the rapid attainment of relatively constant levels of diversity at the local scale of individual patches. At the regional scale of streams and their catchments, several studies have reported negative correlations between diversity and levels of flood disturbance, whereas other studies have reported unimodal diversity–disturbance curves consistent with patterns expected of the intermediate disturbance hypothesis. Such a unimodal relationship may be generated in several ways that await testing. In flowing waters at the regional scale, disturbance may play a central role in regulating species diversity. A predicted increase in the severity and frequency of disturbances with global climate change requires a comprehensive understanding of the disturbance ecology of running waters.

@article{doi1023071468118,
    author = "Lake, P. S.",
    title = "Disturbance, patchiness, and diversity in streams",
    year = "2000",
    journal = "Journal of the North American Benthological Society",
    abstract = "Perturbations in ecosystems consist of a sequence of 2 events: the disturbance, marked by the application of the disturbing forces, and the response shown by the biota to the damage inflicted by the disturbance. The disturbance must be effectively characterized, without confounding it with the response, for progress to be made in the study of the disturbance ecology of streams. A disturbance may take the form of a pulse, a press, or a ramp, and the consequent trajectory of the response may be a pulse, a press, or a ramp.Floods and droughts are the major forms of natural disturbance in flowing waters and, although the effects of floods have been relatively well studied, those of droughts have been largely neglected. Floods accentuate downstream and lateral transport links, often with damaging consequences, whereas droughts fragment the continuity of streams. Both floods and droughts destroy and generate habitat patchiness and patchiness of the biota. During recovery, there are changes in the biotic composition and spatial configuration in patches. Resistance and resilience of the biota to disturbance may be facilitated by the use of refugia. The characterization of flood refugia is much more advanced than that of drought refugia.Recovery from floods is marked by the rapid attainment of relatively constant levels of diversity at the local scale of individual patches. At the regional scale of streams and their catchments, several studies have reported negative correlations between diversity and levels of flood disturbance, whereas other studies have reported unimodal diversity–disturbance curves consistent with patterns expected of the intermediate disturbance hypothesis. Such a unimodal relationship may be generated in several ways that await testing. In flowing waters at the regional scale, disturbance may play a central role in regulating species diversity. A predicted increase in the severity and frequency of disturbances with global climate change requires a comprehensive understanding of the disturbance ecology of running waters.",
    url = "https://doi.org/10.2307/1468118",
    doi = "10.2307/1468118",
    openalex = "W2075197853",
    references = "doi101126science972526482b, doi1023075503, mcauliffe1984competition"
}

Global diversity curves reflect more than just the number of taxa that have existed through time: they also mirror variation in the nature of the fossil record and the way the record is reported. These sampling effects are best quantified by assembling and analyzing large numbers of locality-specific biotic inventories. Here, we introduce a new database of this kind for the Phanerozoic fossil record of marine invertebrates. We apply four substantially distinct analytical methods that estimate taxonomic diversity by quantifying and correcting for variation through time in the number and nature of inventories. Variation introduced by the use of two dramatically different counting protocols also is explored. We present sampling-standardized diversity estimates for two long intervals that sum to 300 Myr (Middle Ordovician-Carboniferous; Late Jurassic-Paleogene). Our new curves differ considerably from traditional, synoptic curves. For example, some of them imply unexpectedly low late Cretaceous and early Tertiary diversity levels. However, such factors as the current emphasis in the database on North America and Europe still obscure our view of the global history of marine biodiversity. These limitations will be addressed as the database and methods are refined.

@article{doi101073pnas111144698,
    author = "Alroy, John and Marshall, Charles R. and Bambach, Richard K. and Bezusko, Karen M. and Foote, Michael and Fürsich, Franz T. and Hansen, Thor A. and Holland, Steven M. and Ivany, Linda C. and Jablonski, David and Jacobs, David K. and Jones, Donna C. and Kosnik, Matthew A. and Lidgard, Scott and Low, Shook Ling and Miller, Arnold I. and Novack‐Gottshall, Philip M. and Olszewski, Thomas D. and Patzkowsky, Mark E. and Raup, David M. and Roy, Kaustuv and Sepkoski, J. John and Sommers, M. G. and Wagner, Peter J. and Webber, Andrew J.",
    title = "Effects of sampling standardization on estimates of Phanerozoic marine diversification",
    year = "2001",
    journal = "Proceedings of the National Academy of Sciences",
    abstract = "Global diversity curves reflect more than just the number of taxa that have existed through time: they also mirror variation in the nature of the fossil record and the way the record is reported. These sampling effects are best quantified by assembling and analyzing large numbers of locality-specific biotic inventories. Here, we introduce a new database of this kind for the Phanerozoic fossil record of marine invertebrates. We apply four substantially distinct analytical methods that estimate taxonomic diversity by quantifying and correcting for variation through time in the number and nature of inventories. Variation introduced by the use of two dramatically different counting protocols also is explored. We present sampling-standardized diversity estimates for two long intervals that sum to 300 Myr (Middle Ordovician-Carboniferous; Late Jurassic-Paleogene). Our new curves differ considerably from traditional, synoptic curves. For example, some of them imply unexpectedly low late Cretaceous and early Tertiary diversity levels. However, such factors as the current emphasis in the database on North America and Europe still obscure our view of the global history of marine biodiversity. These limitations will be addressed as the database and methods are refined.",
    url = "https://doi.org/10.1073/pnas.111144698",
    doi = "10.1073/pnas.111144698",
    openalex = "W2160525739",
    references = "doi101017s0022336000040026, doi101017s0094837300003778, doi101017s0094837300004929, doi101017s0094837300005236, doi101017s0094837300005972, doi101017s0094837300006539, doi101017s0094837300008186, doi10102994jb01889, doi101038293435a0, doi101086282541, doi101098rstb19940091, doi101126science11536722, doi101126science11537491, doi101126science21545391501, doi1015159780691224244, doi102110pec9504, doi1023072412725, doi1023073515180, openalexw1528487914, openalexw2989049194, valentine1978a"
}

Many features of global diversity compilations have proven robust to continued sampling and taxonomic revision. Inherent biases in the stratigraphic record may nevertheless substantially affect estimates of global taxonomic diversity. Here we focus on short-term (epoch-level) changes in apparent diversity. We use a simple estimate of the amount of marine sedimentary rock available for sampling: the number of formations in the stratigraphic Lexicon of the United States Geological Survey. We find this to be positively correlated with two independent estimates of rock availability: global outcrop area derived from the Paleogeographic Atlas Project (University of Chicago) database, and percent continental flooding. Epoch-to-epoch changes in the number of formations are positively correlated with changes in sampled Phanerozoic marine diversity at the genus level. We agree with previous workers in finding evidence of a diversity-area effect that is substantially weaker than the effect of the amount of preserved sedimentary rock. Once the mutual correlation among change in formation numbers, in diversity, and in area flooded is taken into consideration, there is relatively little residual correlation between change in diversity and in the extent of continental flooding. These results suggest that much of the observed short-term variation in marine diversity may be an artifact of variation in the amount of rock available for study. Preliminary results suggest the same possibility for terrestrial data. Like the comparison between change in number of formations and change in sampled diversity, which addresses short-term variation in apparent diversity, the comparison between absolute values of these quantities, which relates to longer-term patterns, also shows a positive correlation. Moreover, there is no clear temporal trend in the residuals of the regression of sampled diversity on number of formations. This raises the possibility that taxonomic diversity may not have increased substantially since the early Paleozoic. Because of limitations in our data, however, this question must remain open.

@article{doi1016660094837320010270583bitpar20co2,
    author = "Peters, Shanan E. and Foote, Michael",
    title = "Biodiversity in the Phanerozoic: a reinterpretation",
    year = "2001",
    journal = "Paleobiology",
    abstract = "Many features of global diversity compilations have proven robust to continued sampling and taxonomic revision. Inherent biases in the stratigraphic record may nevertheless substantially affect estimates of global taxonomic diversity. Here we focus on short-term (epoch-level) changes in apparent diversity. We use a simple estimate of the amount of marine sedimentary rock available for sampling: the number of formations in the stratigraphic Lexicon of the United States Geological Survey. We find this to be positively correlated with two independent estimates of rock availability: global outcrop area derived from the Paleogeographic Atlas Project (University of Chicago) database, and percent continental flooding. Epoch-to-epoch changes in the number of formations are positively correlated with changes in sampled Phanerozoic marine diversity at the genus level. We agree with previous workers in finding evidence of a diversity-area effect that is substantially weaker than the effect of the amount of preserved sedimentary rock. Once the mutual correlation among change in formation numbers, in diversity, and in area flooded is taken into consideration, there is relatively little residual correlation between change in diversity and in the extent of continental flooding. These results suggest that much of the observed short-term variation in marine diversity may be an artifact of variation in the amount of rock available for study. Preliminary results suggest the same possibility for terrestrial data. Like the comparison between change in number of formations and change in sampled diversity, which addresses short-term variation in apparent diversity, the comparison between absolute values of these quantities, which relates to longer-term patterns, also shows a positive correlation. Moreover, there is no clear temporal trend in the residuals of the regression of sampled diversity on number of formations. This raises the possibility that taxonomic diversity may not have increased substantially since the early Paleozoic. Because of limitations in our data, however, this question must remain open.",
    url = "https://doi.org/10.1666/0094-8373(2001)027<0583:bitpar>2.0.co;2",
    doi = "10.1666/0094-8373(2001)027<0583:bitpar>2.0.co;2",
    openalex = "W2174571031",
    references = "doi1010160377839895000100, doi101017s0022336000040026, doi101017s0094837300004929, doi101017s0094837300004930, doi10103818872, doi101073pnas111144698, doi101146annurevea12050184001225, hallam1977secular"
}

Up to 50% of the increase in marine animal biodiversity through the Cenozoic at the genus level has been attributed to a sampling bias termed "the Pull of the Recent," the extension of stratigraphic ranges of fossil taxa by the relatively complete sampling of the Recent biota. However, 906 of 958 living genera and subgenera of bivalve mollusks having a fossil record occur in the Pliocene or Pleistocene. The Pull of the Recent thus accounts for only 5% of the Cenozoic increase in bivalve diversity, a major component of the marine record, suggesting that the diversity increase is likely to be a genuine biological pattern.

@article{doi101126science1083246,
    author = "Jablonski, David and Roy, Kaustuv and Valentine, James W. and Price, Rebecca and Anderson, Philip S. L.",
    title = "The Impact of the Pull of the Recent on the History of Marine Diversity",
    year = "2003",
    journal = "Science",
    abstract = {Up to 50\% of the increase in marine animal biodiversity through the Cenozoic at the genus level has been attributed to a sampling bias termed "the Pull of the Recent," the extension of stratigraphic ranges of fossil taxa by the relatively complete sampling of the Recent biota. However, 906 of 958 living genera and subgenera of bivalve mollusks having a fossil record occur in the Pliocene or Pleistocene. The Pull of the Recent thus accounts for only 5\% of the Cenozoic increase in bivalve diversity, a major component of the marine record, suggesting that the diversity increase is likely to be a genuine biological pattern.},
    url = "https://doi.org/10.1126/science.1083246",
    doi = "10.1126/science.1083246",
    openalex = "W2115328825",
    references = "doi101017s0094837300026907, valentine1978a"
}

A central goal of evolutionary biology is the development of a general predictive theory that incorporates the various forces structuring biological diversity. In this article, we argue that a focus on allometry and metabolic scaling theory provides the basis to begin to mechanistically link and understand patterns observed at ecological and evolutionary timescales. To make our case, we draw on and review several recently published papers and present new analyses. We argue that together, this synthesized work supports the notion that selection has maximized the scaling of resource exchange surfaces (e.g., photosynthetic surface areas) yet simultaneously minimizes the scaling of transport times and resistances. As a result, the scaling of plant metabolism, in turn, has profoundly influenced the evolution and ecology of plant form, function, and diversity, probably since the inception of the chlorophytes. In particular, we discuss preliminary support for the notion that stabilizing selection for 3/4‐power scaling of metabolism, in addition to competition for similar limiting resources, has led to the emergence of regular ecological patterns that have likely been prevalent throughout plant evolution.

@article{doi101086513479,
    author = "Enquist, Brian J. and Tiffney, Bruce H. and Niklas, Karl J.",
    title = "Metabolic Scaling and the Evolutionary Dynamics of Plant Size, Form, and Diversity: Toward a Synthesis of Ecology, Evolution, and Paleontology",
    year = "2007",
    journal = "International Journal of Plant Sciences",
    abstract = "A central goal of evolutionary biology is the development of a general predictive theory that incorporates the various forces structuring biological diversity. In this article, we argue that a focus on allometry and metabolic scaling theory provides the basis to begin to mechanistically link and understand patterns observed at ecological and evolutionary timescales. To make our case, we draw on and review several recently published papers and present new analyses. We argue that together, this synthesized work supports the notion that selection has maximized the scaling of resource exchange surfaces (e.g., photosynthetic surface areas) yet simultaneously minimizes the scaling of transport times and resistances. As a result, the scaling of plant metabolism, in turn, has profoundly influenced the evolution and ecology of plant form, function, and diversity, probably since the inception of the chlorophytes. In particular, we discuss preliminary support for the notion that stabilizing selection for 3/4‐power scaling of metabolism, in addition to competition for similar limiting resources, has led to the emergence of regular ecological patterns that have likely been prevalent throughout plant evolution.",
    url = "https://doi.org/10.1086/513479",
    doi = "10.1086/513479",
    openalex = "W2008786418",
    references = "doi101017cbo9780511608551, doi101086282505, doi101111j155856461983tb00236x, doi101126science1064088, doi101126science2765309122, doi101890039000, doi1023073071998, doi1023074549, doi105860choice332720, knoll1979phanerozoic, openalexw2077454220"
}

Abstract: Palaeodiversity curves are constructed from counts of fossils collected at outcrop and thus potentially biased by variation in the rock record, specifically by the amount of sedimentary rock representative of different time intervals that has been preserved at outcrop. To investigate how much of a problem this poses we have compiled a high‐resolution record of marine rock outcrop area in Western Europe for the Phanerozoic and use this to generate a model that predicts the sampled diversity curve. We find that we can predict with high accuracy the variance of the marine genus diversity curve (itself dominated by European taxa) from rock outcrop data and a three‐step model of diversity that tracks supercontinent fragmentation, coalescence and fragmentation. The size and position of two of the five major mass extinction spikes are largely predicted by rock outcrop data. We conclude that the long‐term trends in taxonomic diversity and the end‐Cretaceous extinction are not the result of rock area bias, but cannot rule out that rock outcrop area bias explains many of the short‐term rises and falls in sampled diversity that palaeontologists have previously sought to explain biologically.

@article{doi101111j14754983200700693x,
    author = "Smith, Andrew B. and McGowan, Alistair J.",
    title = "THE SHAPE OF THE PHANEROZOIC MARINE PALAEODIVERSITY CURVE: HOW MUCH CAN BE PREDICTED FROM THE SEDIMENTARY ROCK RECORD OF WESTERN EUROPE?",
    year = "2007",
    journal = "Palaeontology",
    abstract = "Abstract: Palaeodiversity curves are constructed from counts of fossils collected at outcrop and thus potentially biased by variation in the rock record, specifically by the amount of sedimentary rock representative of different time intervals that has been preserved at outcrop. To investigate how much of a problem this poses we have compiled a high‐resolution record of marine rock outcrop area in Western Europe for the Phanerozoic and use this to generate a model that predicts the sampled diversity curve. We find that we can predict with high accuracy the variance of the marine genus diversity curve (itself dominated by European taxa) from rock outcrop data and a three‐step model of diversity that tracks supercontinent fragmentation, coalescence and fragmentation. The size and position of two of the five major mass extinction spikes are largely predicted by rock outcrop data. We conclude that the long‐term trends in taxonomic diversity and the end‐Cretaceous extinction are not the result of rock area bias, but cannot rule out that rock outcrop area bias explains many of the short‐term rises and falls in sampled diversity that palaeontologists have previously sought to explain biologically.",
    url = "https://doi.org/10.1111/j.1475-4983.2007.00693.x",
    doi = "10.1111/j.1475-4983.2007.00693.x",
    openalex = "W2126310917",
    references = "doi101146annurevecolsys33030602152151, doi1016660094837320050310006poaeit20co2, smith2007marine"
}

Abstract According to when they attained high diversity, major taxa of marine animals have been clustered into three groups, the Cambrian, Paleozoic, and Modern Faunas. Because the Cambrian Fauna was a relatively minor component of the total fauna after mid-Ordovician time, the Phanerozoic history of marine animal diversity is largely a matter of the fates of the Paleozoic and Modern Faunas. The fact that most late Cenozoic genera belong to taxa that have been radiating for tens of millions of years indicates that the post-Paleozoic increase in diversity indicated by fossil data is real, rather than an artifact of improvement of the fossil record toward the present. Assuming that ecological crowding produced the so-called Paleozoic plateau for family diversity, various workers have used the logistic equation of ecology to model marine animal diversification as damped exponential increase. Several lines of evidence indicate that this procedure is inappropriate. A plot of the diversity of marine animal gene...

@article{doi101666060201,
    author = "Stanley, Steven M.",
    title = "An Analysis of the History of Marine Animal Diversity",
    year = "2007",
    journal = "Paleobiology",
    abstract = "Abstract According to when they attained high diversity, major taxa of marine animals have been clustered into three groups, the Cambrian, Paleozoic, and Modern Faunas. Because the Cambrian Fauna was a relatively minor component of the total fauna after mid-Ordovician time, the Phanerozoic history of marine animal diversity is largely a matter of the fates of the Paleozoic and Modern Faunas. The fact that most late Cenozoic genera belong to taxa that have been radiating for tens of millions of years indicates that the post-Paleozoic increase in diversity indicated by fossil data is real, rather than an artifact of improvement of the fossil record toward the present. Assuming that ecological crowding produced the so-called Paleozoic plateau for family diversity, various workers have used the logistic equation of ecology to model marine animal diversification as damped exponential increase. Several lines of evidence indicate that this procedure is inappropriate. A plot of the diversity of marine animal gene...",
    url = "https://doi.org/10.1666/06020.1",
    doi = "10.1666/06020.1",
    openalex = "W2153512785",
    references = "doi101017s0094837300005248, doi101111j1469185x1973tb00979x, doi101130g211551, doi101144pygs5211, valentine1978a"
}

The fossil record provides direct evidence of how diversity has changed over time, but cannot be taken at face value. Diversity curves constructed from counting taxa in the rock record are seriously biased by unevenness of geographical and stratigraphical sampling effort, inequality in the rock record available for sampling, and inconsistent taxonomic practice. Sample standardization removes some bias, but does not overcome more general incompleteness problems. Modelling that accounts for potential biases is a newer approach but needs accurate estimates of rock record and consistent taxonomic data. Uncertainty remains over whether the steep rise in diversity over the last 100 Ma is real or reflects sampling bias. The repeated rise and fall of marine diversity over time correlates closely with the areal extent of sedimentary deposits and independent estimates of the quality of the fossil record, implying a common driving factor, namely tectonically mediated sea-level change. However, whether changes in diversity are primarily biological in origin, or reflect sampling artefact, remains contentious. There is a distinct possibility that many of the apparent rises and falls in diversity over the Phanerozoic, including most of the ‘mass extinctions’, arise either partially or entirely from rock record bias.

@article{smith2007marine,
    author = "SMITH, ANDREW B.",
    title = "Marine diversity through the Phanerozoic: problems and prospects",
    year = "2007",
    journal = "Journal of the Geological Society",
    abstract = "The fossil record provides direct evidence of how diversity has changed over time, but cannot be taken at face value. Diversity curves constructed from counting taxa in the rock record are seriously biased by unevenness of geographical and stratigraphical sampling effort, inequality in the rock record available for sampling, and inconsistent taxonomic practice. Sample standardization removes some bias, but does not overcome more general incompleteness problems. Modelling that accounts for potential biases is a newer approach but needs accurate estimates of rock record and consistent taxonomic data. Uncertainty remains over whether the steep rise in diversity over the last 100 Ma is real or reflects sampling bias. The repeated rise and fall of marine diversity over time correlates closely with the areal extent of sedimentary deposits and independent estimates of the quality of the fossil record, implying a common driving factor, namely tectonically mediated sea-level change. However, whether changes in diversity are primarily biological in origin, or reflect sampling artefact, remains contentious. There is a distinct possibility that many of the apparent rises and falls in diversity over the Phanerozoic, including most of the ‘mass extinctions’, arise either partially or entirely from rock record bias.",
    url = "https://doi.org/10.1144/0016/76492006-184",
    doi = "10.1144/0016/76492006-184",
    number = "4",
    pages = "731-745",
    volume = "164"
}

It has previously been thought that there was a steep Cretaceous and Cenozoic radiation of marine invertebrates. This pattern can be replicated with a new data set of fossil occurrences representing 3.5 million specimens, but only when older analytical protocols are used. Moreover, analyses that employ sampling standardization and more robust counting methods show a modest rise in diversity with no clear trend after the mid-Cretaceous. Globally, locally, and at both high and low latitudes, diversity was less than twice as high in the Neogene as in the mid-Paleozoic. The ratio of global to local richness has changed little, and a latitudinal diversity gradient was present in the early Paleozoic.

@article{doi101126science1156963,
    author = "Alroy, John and Aberhan, Martin and Bottjer, David J. and Foote, Michael and Fürsich, Franz T. and Harries, Peter J. and Hendy, Austin and Holland, Steven M. and Ivany, Linda C. and Kiessling, Wolfgang and Kosnik, Matthew A. and Marshall, Charles R. and McGowan, Alistair J. and Miller, Arnold I. and Olszewski, Thomas D. and Patzkowsky, Mark E. and Peters, Shanan E. and Villier, Loïc and Wagner, Peter J. and Bonuso, Nicole and Borkow, Philip S. and Brenneis, Benjamin and Clapham, Matthew E. and Fall, Leigh M. and Ferguson, Chad Allen and Hanson, Victoria L. and Krug, Andrew Z. and Layou, Karen M. and Leckey, Erin and Nürnberg, Sabine and Powers, Catherine M. and Sessa, Jocelyn A. and Simpson, Carl and Tomášových, Adam and Visaggi, Christy C.",
    title = "Phanerozoic Trends in the Global Diversity of Marine Invertebrates",
    year = "2008",
    journal = "Science",
    abstract = "It has previously been thought that there was a steep Cretaceous and Cenozoic radiation of marine invertebrates. This pattern can be replicated with a new data set of fossil occurrences representing 3.5 million specimens, but only when older analytical protocols are used. Moreover, analyses that employ sampling standardization and more robust counting methods show a modest rise in diversity with no clear trend after the mid-Cretaceous. Globally, locally, and at both high and low latitudes, diversity was less than twice as high in the Neogene as in the mid-Paleozoic. The ratio of global to local richness has changed little, and a latitudinal diversity gradient was present in the early Paleozoic.",
    url = "https://doi.org/10.1126/science.1156963",
    doi = "10.1126/science.1156963",
    openalex = "W2019736558",
    references = "doi1010079781475707403, doi101016s001282520000026x, doi101017s0094837300004929, doi101017s0094837300008186, doi101038293435a0, doi101073pnas092150999, doi101073pnas111144698, doi101126science17740541065, doi101126science21545391501, doi101126science2164542173, doi101126science7701342, doi101130g211551, doi1016660094837320002674oaecot20co2, doi1016660094837320010270583bitpar20co2"
}

The fossil record is our only direct means for evaluating shifts in biodiversity through Earth's history. However, analyses of fossil marine invertebrates have demonstrated that geological megabiases profoundly influence fossil preservation and discovery, obscuring true diversity signals. Comparable studies of vertebrate palaeodiversity patterns remain in their infancy. A new species-level dataset of Mesozoic marine tetrapod occurrences was compared with a proxy for temporal variation in the volume and facies diversity of fossiliferous rock (number of marine fossiliferous formations: FMF). A strong correlation between taxic diversity and FMF is present during the Cretaceous. Weak or no correlation of Jurassic data suggests a qualitatively different sampling regime resulting from five apparent peaks in Triassic-Jurassic diversity. These correspond to a small number of European formations that have been the subject of intensive collecting, and represent 'Lagerstätten effects'. Consideration of sampling biases allows re-evaluation of proposed mass extinction events. Marine tetrapod diversity declined during the Carnian or Norian. However, the proposed end-Triassic extinction event cannot be recognized with confidence. Some evidence supports an extinction event near the Jurassic/Cretaceous boundary, but the proposed end-Cenomanian extinction is probably an artefact of poor sampling. Marine tetrapod diversity underwent a long-term decline prior to the Cretaceous-Palaeogene extinction.

@article{doi101098rspb20091845,
    author = "Benson, Roger and Butler, Richard J. and Lindgren, Johan and Smith, Adam S.",
    title = "Mesozoic marine tetrapod diversity: mass extinctions and temporal heterogeneity in geological megabiases affecting vertebrates",
    year = "2009",
    journal = "Proceedings of the Royal Society B Biological Sciences",
    abstract = "The fossil record is our only direct means for evaluating shifts in biodiversity through Earth's history. However, analyses of fossil marine invertebrates have demonstrated that geological megabiases profoundly influence fossil preservation and discovery, obscuring true diversity signals. Comparable studies of vertebrate palaeodiversity patterns remain in their infancy. A new species-level dataset of Mesozoic marine tetrapod occurrences was compared with a proxy for temporal variation in the volume and facies diversity of fossiliferous rock (number of marine fossiliferous formations: FMF). A strong correlation between taxic diversity and FMF is present during the Cretaceous. Weak or no correlation of Jurassic data suggests a qualitatively different sampling regime resulting from five apparent peaks in Triassic-Jurassic diversity. These correspond to a small number of European formations that have been the subject of intensive collecting, and represent 'Lagerstätten effects'. Consideration of sampling biases allows re-evaluation of proposed mass extinction events. Marine tetrapod diversity declined during the Carnian or Norian. However, the proposed end-Triassic extinction event cannot be recognized with confidence. Some evidence supports an extinction event near the Jurassic/Cretaceous boundary, but the proposed end-Cenomanian extinction is probably an artefact of poor sampling. Marine tetrapod diversity underwent a long-term decline prior to the Cretaceous-Palaeogene extinction.",
    url = "https://doi.org/10.1098/rspb.2009.1845",
    doi = "10.1098/rspb.2009.1845",
    openalex = "W2110009186",
    references = "doi101017s0094837300004929, doi101038274661a0, doi101038293435a0, doi10108002724634198710011647, doi10108010292389409380462, doi101098rspb20080715, doi101111j10963642200900571x, doi101111j251761611995tb02031x, doi101126science1156963, doi101126science17740541065, doi101130spe190p291, doi101525california97805202462320010001, doi1016660094837320010270583bitpar20co2, openalexw586972754, openalexw592572837, smith2007marine"
}

Abstract: The Paleobiology Database now includes enough data on fossil collections to produce useful time series of geographical and environmental variables in addition to a robust global Phanerozoic marine diversity curve. The curve is produced by a new ‘shareholder quorum’ method of sampling standardization that removes biases but avoids overcompensating for them by imposing entirely uniform data quotas. It involves drawing fossil collections until the taxa that have been sampled at least once (the ‘shareholders’) have a summed total of frequencies (i.e. coverage) that meets a target (the ‘quorum’). Coverage of each interval’s entire data set is estimated prior to subsampling using a variant of a standard index, Good’s u. This variant employs counts of occurrences of taxa described in only one publication instead of taxa found in only one collection. Each taxon’s frequency within an interval is multiplied by the interval’s index value, which limits the maximum possible sampling level and thereby creates the need for subsampling. Analyses focus on a global diversity curve and curves for northern, southern and ‘tropical’ (30°N to 30°S) palaeolatitudinal belts. Tropical genus richness is remarkably static, so most large shifts in the curve reflect trends at higher latitudes. Changes in diversity are analysed as a function of standing diversity; the number, spacing and palaeolatitudinal position of sampled geographical cells; the mean onshore–offshore position of cells; and proportions of cells from carbonate, onshore and reefal environments. Redundancy among the variables is eliminated by performing a principal components analysis of each data set and using the axis scores in multiple regressions. The key factors are standing diversity and the dominance of onshore environments such as reefs. These factors combine to produce logistic growth patterns with slowly changing equilibrium values. There is no evidence of unregulated exponential growth across any long stretch of the Phanerozoic, and in particular there was no large Cenozoic radiation beyond the Eocene. The end-Ordovician, Permo–Triassic and Cretaceous–Palaeogene mass extinctions had relatively short-term albeit severe effects. However, reef collapse was involved in these events and also may have caused large, longer term global diversity decreases in the mid-Devonian and across the Triassic/Jurassic boundary. Conversely, the expansion of reef ecosystems may explain newly recognized major radiations in the mid-Permian and mid-Jurassic. Reef ecosystems are particularly vulnerable to current environmental disturbances such as ocean acidification, and their decimation might prolong the recovery from today’s mass extinction by millions or even tens of millions of years.

@article{doi101111j14754983201001011x,
    author = "Alroy, John",
    title = "Geographical, environmental and intrinsic biotic controls on Phanerozoic marine diversification",
    year = "2010",
    journal = "Palaeontology",
    abstract = "Abstract: The Paleobiology Database now includes enough data on fossil collections to produce useful time series of geographical and environmental variables in addition to a robust global Phanerozoic marine diversity curve. The curve is produced by a new ‘shareholder quorum’ method of sampling standardization that removes biases but avoids overcompensating for them by imposing entirely uniform data quotas. It involves drawing fossil collections until the taxa that have been sampled at least once (the ‘shareholders’) have a summed total of frequencies (i.e. coverage) that meets a target (the ‘quorum’). Coverage of each interval’s entire data set is estimated prior to subsampling using a variant of a standard index, Good’s u. This variant employs counts of occurrences of taxa described in only one publication instead of taxa found in only one collection. Each taxon’s frequency within an interval is multiplied by the interval’s index value, which limits the maximum possible sampling level and thereby creates the need for subsampling. Analyses focus on a global diversity curve and curves for northern, southern and ‘tropical’ (30°N to 30°S) palaeolatitudinal belts. Tropical genus richness is remarkably static, so most large shifts in the curve reflect trends at higher latitudes. Changes in diversity are analysed as a function of standing diversity; the number, spacing and palaeolatitudinal position of sampled geographical cells; the mean onshore–offshore position of cells; and proportions of cells from carbonate, onshore and reefal environments. Redundancy among the variables is eliminated by performing a principal components analysis of each data set and using the axis scores in multiple regressions. The key factors are standing diversity and the dominance of onshore environments such as reefs. These factors combine to produce logistic growth patterns with slowly changing equilibrium values. There is no evidence of unregulated exponential growth across any long stretch of the Phanerozoic, and in particular there was no large Cenozoic radiation beyond the Eocene. The end-Ordovician, Permo–Triassic and Cretaceous–Palaeogene mass extinctions had relatively short-term albeit severe effects. However, reef collapse was involved in these events and also may have caused large, longer term global diversity decreases in the mid-Devonian and across the Triassic/Jurassic boundary. Conversely, the expansion of reef ecosystems may explain newly recognized major radiations in the mid-Permian and mid-Jurassic. Reef ecosystems are particularly vulnerable to current environmental disturbances such as ocean acidification, and their decimation might prolong the recovery from today’s mass extinction by millions or even tens of millions of years.",
    url = "https://doi.org/10.1111/j.1475-4983.2010.01011.x",
    doi = "10.1111/j.1475-4983.2010.01011.x",
    openalex = "W2122289512",
    references = "doi101017s0094837300004930, doi101046j14610248200100230x, doi101093biomet4034237, doi101098rstb19940091, doi101111j14754983200600612x, doi101126science1152509, doi101126science1177265, doi101126science21545391501, doi1015159781400881376, doi1023071934145, knoll1979phanerozoic, openalexw2145250129"
}

The stratigraphic record from both deep-sea and shallow-water depositional environments indicates that during late Aptian through Cenomanian time (1) global climates were considerably warmer than at present; (2) latitudinal gradients of atmospheric and oceanic temperatures were considerably less than at present; (3) rates of accumulation of organic matter of both marine and terrestrial origin were as high as or higher than during any other interval in the Mesozoic or Cenozoic; (4) the rate and volume of accumulation of CaCO3 in the deep sea were reduced in response to a marked shoaling of the carbonate compensation depth; (5) seafloor spreading rates were somewhat more rapid than at any other time in the Cretaceous or Cenozoic; (6) off-ridge volcanism was intense and widespread, particularly in the ancestral Pacific Ocean basin; and (7) sea level was relatively high, forming widespread areas of shallow shelf seas. A marked increase in the rate of CO2 outgassing due to volcanic activity between about 110 and 70 m.y. ago may have resulted in a buildup of atmospheric CO2. A significant fraction of this atmospheric CO2 may have been reduced by an increase in the production and burial of terrestrial organic carbon. Some excess CO2 may have been consumed by marine algal photosynthesis, but marine productivity apparently was low during the Aptian-Albian relative to terrestrial productivity. Terrestrial productivity also may have been stimulated by increased rainfall that resulted from a warm global climate and increased marine transgression as well as by the higher CO2.

@incollection{doi101029gm032p0504,
    author = "Arthur, M. A. and Dean, Walter E. and Schlanger, S. O.",
    title = "Variations in the Global Carbon Cycle During the Cretaceous Related to Climate, Volcanism, and Changes in Atmospheric CO 2",
    year = "2011",
    booktitle = "Geophysical monograph",
    abstract = "The stratigraphic record from both deep-sea and shallow-water depositional environments indicates that during late Aptian through Cenomanian time (1) global climates were considerably warmer than at present; (2) latitudinal gradients of atmospheric and oceanic temperatures were considerably less than at present; (3) rates of accumulation of organic matter of both marine and terrestrial origin were as high as or higher than during any other interval in the Mesozoic or Cenozoic; (4) the rate and volume of accumulation of CaCO3 in the deep sea were reduced in response to a marked shoaling of the carbonate compensation depth; (5) seafloor spreading rates were somewhat more rapid than at any other time in the Cretaceous or Cenozoic; (6) off-ridge volcanism was intense and widespread, particularly in the ancestral Pacific Ocean basin; and (7) sea level was relatively high, forming widespread areas of shallow shelf seas. A marked increase in the rate of CO2 outgassing due to volcanic activity between about 110 and 70 m.y. ago may have resulted in a buildup of atmospheric CO2. A significant fraction of this atmospheric CO2 may have been reduced by an increase in the production and burial of terrestrial organic carbon. Some excess CO2 may have been consumed by marine algal photosynthesis, but marine productivity apparently was low during the Aptian-Albian relative to terrestrial productivity. Terrestrial productivity also may have been stimulated by increased rainfall that resulted from a warm global climate and increased marine transgression as well as by the higher CO2.",
    url = "https://doi.org/10.1029/gm032p0504",
    doi = "10.1029/gm032p0504",
    openalex = "W1518673644",
    references = "hallam1977secular"
}

@article{doi101007s1091401292221,
    author = "Woodburne, Michael O. and Goin, Francisco J. and Bond, Mariano and Carlini, Alfredo A. and Gelfo, Javier N. and López, Guillermo M. and Iglesias, Ari and Zimicz, Natalia",
    title = "Paleogene Land Mammal Faunas of South America; a Response to Global Climatic Changes and Indigenous Floral Diversity",
    year = "2013",
    journal = "Journal of Mammalian Evolution",
    url = "https://doi.org/10.1007/s10914-012-9222-1",
    doi = "10.1007/s10914-012-9222-1",
    openalex = "W2112038109",
    references = "doi101016s003101829800056x, doi101086430055, doi101111j10958312201101657x, romero1976a"
}

Latitudinal diversity gradients are underlain by complex combinations of origination, extinction, and shifts in geographic distribution and therefore are best analyzed by integrating paleontological and neontological data. The fossil record of marine bivalves shows, in three successive late Cenozoic time slices, that most clades (operationally here, genera) tend to originate in the tropics and then expand out of the tropics (OTT) to higher latitudes while retaining their tropical presence. This OTT pattern is robust both to assumptions on the preservation potential of taxa and to taxonomic revisions of extant and fossil species. Range expansion of clades may occur via "bridge species," which violate climate-niche conservatism to bridge the tropical-temperate boundary in most OTT genera. Substantial time lags (∼5 Myr) between the origins of tropical clades and their entry into the temperate zone suggest that OTT events are rare on a per-clade basis. Clades with higher diversification rates within the tropics are the most likely to expand OTT and the most likely to produce multiple bridge species, suggesting that high speciation rates promote the OTT dynamic. Although expansion of thermal tolerances is key to the OTT dynamic, most latitudinally widespread species instead achieve their broad ranges by tracking widespread, spatially-uniform temperatures within the tropics (yielding, via the nonlinear relation between temperature and latitude, a pattern opposite to Rapoport's rule). This decoupling of range size and temperature tolerance may also explain the differing roles of species and clade ranges in buffering species from background and mass extinctions.

@article{doi101073pnas1308997110,
    author = "Jablonski, David and Belanger, Christina L. and Berke, Sarah K. and Huang, Shan and Krug, Andrew Z. and Roy, Kaustuv and Tomášových, Adam and Valentine, James W.",
    title = "Out of the tropics, but how? Fossils, bridge species, and thermal ranges in the dynamics of the marine latitudinal diversity gradient",
    year = "2013",
    journal = "Proceedings of the National Academy of Sciences",
    abstract = {Latitudinal diversity gradients are underlain by complex combinations of origination, extinction, and shifts in geographic distribution and therefore are best analyzed by integrating paleontological and neontological data. The fossil record of marine bivalves shows, in three successive late Cenozoic time slices, that most clades (operationally here, genera) tend to originate in the tropics and then expand out of the tropics (OTT) to higher latitudes while retaining their tropical presence. This OTT pattern is robust both to assumptions on the preservation potential of taxa and to taxonomic revisions of extant and fossil species. Range expansion of clades may occur via "bridge species," which violate climate-niche conservatism to bridge the tropical-temperate boundary in most OTT genera. Substantial time lags (∼5 Myr) between the origins of tropical clades and their entry into the temperate zone suggest that OTT events are rare on a per-clade basis. Clades with higher diversification rates within the tropics are the most likely to expand OTT and the most likely to produce multiple bridge species, suggesting that high speciation rates promote the OTT dynamic. Although expansion of thermal tolerances is key to the OTT dynamic, most latitudinally widespread species instead achieve their broad ranges by tracking widespread, spatially-uniform temperatures within the tropics (yielding, via the nonlinear relation between temperature and latitude, a pattern opposite to Rapoport's rule). This decoupling of range size and temperature tolerance may also explain the differing roles of species and clade ranges in buffering species from background and mass extinctions.},
    url = "https://doi.org/10.1073/pnas.1308997110",
    doi = "10.1073/pnas.1308997110",
    openalex = "W2150741383",
    references = "doi101093icbicq078"
}

A number of methods have been developed to infer differential rates of species diversification through time and among clades using time-calibrated phylogenetic trees. However, we lack a general framework that can delineate and quantify heterogeneous mixtures of dynamic processes within single phylogenies. I developed a method that can identify arbitrary numbers of time-varying diversification processes on phylogenies without specifying their locations in advance. The method uses reversible-jump Markov Chain Monte Carlo to move between model subspaces that vary in the number of distinct diversification regimes. The model assumes that changes in evolutionary regimes occur across the branches of phylogenetic trees under a compound Poisson process and explicitly accounts for rate variation through time and among lineages. Using simulated datasets, I demonstrate that the method can be used to quantify complex mixtures of time-dependent, diversity-dependent, and constant-rate diversification processes. I compared the performance of the method to the MEDUSA model of rate variation among lineages. As an empirical example, I analyzed the history of speciation and extinction during the radiation of modern whales. The method described here will greatly facilitate the exploration of macroevolutionary dynamics across large phylogenetic trees, which may have been shaped by heterogeneous mixtures of distinct evolutionary processes.

@article{doi101371journalpone0089543,
    author = "Rabosky, Daniel L.",
    title = "Automatic Detection of Key Innovations, Rate Shifts, and Diversity-Dependence on Phylogenetic Trees",
    year = "2014",
    journal = "PLoS ONE",
    abstract = "A number of methods have been developed to infer differential rates of species diversification through time and among clades using time-calibrated phylogenetic trees. However, we lack a general framework that can delineate and quantify heterogeneous mixtures of dynamic processes within single phylogenies. I developed a method that can identify arbitrary numbers of time-varying diversification processes on phylogenies without specifying their locations in advance. The method uses reversible-jump Markov Chain Monte Carlo to move between model subspaces that vary in the number of distinct diversification regimes. The model assumes that changes in evolutionary regimes occur across the branches of phylogenetic trees under a compound Poisson process and explicitly accounts for rate variation through time and among lineages. Using simulated datasets, I demonstrate that the method can be used to quantify complex mixtures of time-dependent, diversity-dependent, and constant-rate diversification processes. I compared the performance of the method to the MEDUSA model of rate variation among lineages. As an empirical example, I analyzed the history of speciation and extinction during the radiation of modern whales. The method described here will greatly facilitate the exploration of macroevolutionary dynamics across large phylogenetic trees, which may have been shaped by heterogeneous mixtures of distinct evolutionary processes.",
    url = "https://doi.org/10.1371/journal.pone.0089543",
    doi = "10.1371/journal.pone.0089543",
    openalex = "W2042449834",
    references = "doi10100797814612066756, doi101017s0094837300005972, doi101038nature05634, doi101038nature11631, doi10106311699114, doi101093bioinformaticsbtm538, doi101093biomet57197, doi101093biomet824711, doi101111j14610248200701020x, doi101126science2354785167, doi101186147121481393"
}

228:657-659] hypothesized that plate tectonics regulates global biodiversity by changing the geographic arrangement of continental crust, but the data required to fully test the hypothesis were not available. Here, we use a global database of marine animal fossil occurrences and a paleogeographic reconstruction model to test the hypothesis that temporal patterns of continental fragmentation have impacted global Phanerozoic biodiversity. We find a positive correlation between global marine invertebrate genus richness and an independently derived quantitative index describing the fragmentation of continental crust during supercontinental coalescence-breakup cycles. The observed positive correlation between global biodiversity and continental fragmentation is not readily attributable to commonly cited vagaries of the fossil record, including changing quantities of marine rock or time-variable sampling effort. Because many different environmental and biotic factors may covary with changes in the geographic arrangement of continental crust, it is difficult to identify a specific causal mechanism. However, cross-correlation indicates that the state of continental fragmentation at a given time is positively correlated with the state of global biodiversity for tens of millions of years afterward. There is also evidence to suggest that continental fragmentation promotes increasing marine richness, but that coalescence alone has only a small negative or stabilizing effect. Together, these results suggest that continental fragmentation, particularly during the Mesozoic breakup of the supercontinent Pangaea, has exerted a first-order control on the long-term trajectory of Phanerozoic marine animal diversity.

@article{doi101073pnas1702297114,
    author = "Zaffos, Andrew and Finnegan, Seth and Peters, Shanan E.",
    title = "Plate tectonic regulation of global marine animal diversity",
    year = "2017",
    journal = "Proceedings of the National Academy of Sciences",
    abstract = "228:657-659] hypothesized that plate tectonics regulates global biodiversity by changing the geographic arrangement of continental crust, but the data required to fully test the hypothesis were not available. Here, we use a global database of marine animal fossil occurrences and a paleogeographic reconstruction model to test the hypothesis that temporal patterns of continental fragmentation have impacted global Phanerozoic biodiversity. We find a positive correlation between global marine invertebrate genus richness and an independently derived quantitative index describing the fragmentation of continental crust during supercontinental coalescence-breakup cycles. The observed positive correlation between global biodiversity and continental fragmentation is not readily attributable to commonly cited vagaries of the fossil record, including changing quantities of marine rock or time-variable sampling effort. Because many different environmental and biotic factors may covary with changes in the geographic arrangement of continental crust, it is difficult to identify a specific causal mechanism. However, cross-correlation indicates that the state of continental fragmentation at a given time is positively correlated with the state of global biodiversity for tens of millions of years afterward. There is also evidence to suggest that continental fragmentation promotes increasing marine richness, but that coalescence alone has only a small negative or stabilizing effect. Together, these results suggest that continental fragmentation, particularly during the Mesozoic breakup of the supercontinent Pangaea, has exerted a first-order control on the long-term trajectory of Phanerozoic marine animal diversity.",
    url = "https://doi.org/10.1073/pnas.1702297114",
    doi = "10.1073/pnas.1702297114",
    openalex = "W2615085513",
    references = "doi101038nature10969, valentine1978a"
}

Biotic interactions such as competition, predation, and niche construction are fundamental drivers of biodiversity at the local scale, yet their long-term effect during earth history remains controversial. To test their role and explore potential limits to biodiversity, we determine within-habitat (alpha), between-habitat (beta), and overall (gamma) diversity of benthic marine invertebrates for Phanerozoic geological formations. We show that an increase in gamma diversity is consistently generated by an increase in alpha diversity throughout the Phanerozoic. Beta diversity drives gamma diversity only at early stages of diversification but remains stationary once a certain gamma level is reached. This mode is prevalent during early- to mid-Paleozoic periods, whereas coupling of beta and gamma diversity becomes increasingly weak toward the recent. Generally, increases in overall biodiversity were accomplished by adding more species to local habitats, and apparently this process never reached saturation during the Phanerozoic. Our results provide general support for an ecological model in which diversification occurs in successive phases of progressing levels of biotic interactions.

@article{hofmann2019diversity,
    author = "Hofmann, Richard and Tietje, Melanie and Aberhan, Martin",
    title = "Diversity partitioning in Phanerozoic benthic marine communities",
    year = "2019",
    journal = "Proceedings of the National Academy of Sciences",
    abstract = "Biotic interactions such as competition, predation, and niche construction are fundamental drivers of biodiversity at the local scale, yet their long-term effect during earth history remains controversial. To test their role and explore potential limits to biodiversity, we determine within-habitat (alpha), between-habitat (beta), and overall (gamma) diversity of benthic marine invertebrates for Phanerozoic geological formations. We show that an increase in gamma diversity is consistently generated by an increase in alpha diversity throughout the Phanerozoic. Beta diversity drives gamma diversity only at early stages of diversification but remains stationary once a certain gamma level is reached. This mode is prevalent during early- to mid-Paleozoic periods, whereas coupling of beta and gamma diversity becomes increasingly weak toward the recent. Generally, increases in overall biodiversity were accomplished by adding more species to local habitats, and apparently this process never reached saturation during the Phanerozoic. Our results provide general support for an ecological model in which diversification occurs in successive phases of progressing levels of biotic interactions.",
    url = "https://doi.org/10.1073/pnas.1814487116",
    doi = "10.1073/pnas.1814487116",
    number = "1",
    pages = "79-83",
    volume = "116"
}

Across time, but also across space Fossils, especially those from marine systems, have long been used to estimate changes in patterns of diversity over time. However, fossils are patchy in their occurrence, so such temporal estimates generally have not included variations due to space. Such a singular examination has the potential to simplify, or even misrepresent, patterns. Close et al. used a spatially explicit approach to measure diversity changes in marine fossils across time and space. They found that, like modern systems, diversity varies considerably across space, with reefs increasing diversity levels. Accounting for this spatial-environmental variation will shed new light on the study of diversity over time. Science, this issue p. 420

@article{close2020the,
    author = "Close, R. A. and Benson, R. B. J. and Saupe, E. E. and Clapham, M. E. and Butler, R. J.",
    title = "The spatial structure of Phanerozoic marine animal diversity",
    year = "2020",
    journal = "Science",
    abstract = "Across time, but also across space Fossils, especially those from marine systems, have long been used to estimate changes in patterns of diversity over time. However, fossils are patchy in their occurrence, so such temporal estimates generally have not included variations due to space. Such a singular examination has the potential to simplify, or even misrepresent, patterns. Close et al. used a spatially explicit approach to measure diversity changes in marine fossils across time and space. They found that, like modern systems, diversity varies considerably across space, with reefs increasing diversity levels. Accounting for this spatial-environmental variation will shed new light on the study of diversity over time. Science, this issue p. 420",
    url = "https://doi.org/10.1126/science.aay8309",
    doi = "10.1126/science.aay8309",
    number = "6489",
    openalex = "W3018232557",
    pages = "420-424",
    volume = "368",
    references = "doi101002sim3107, doi101017s0094837300008186, doi101093biomet4034237, doi1011112041210x12613, doi101111j155856461960tb03057x, doi101126science1156963, doi101126science21545391501, doi101126science7701342, doi1023073802723, doi1023075503"
}

Conodonts, one of the longest-lived early groups of vertebrates, have a very complete fossil record ranging from the late Cambrian to the end of the Triassic and persisted through many global climatic and biotic events. In this paper, we analyse a large dataset harvested from the Paleobiology Database to compute global diversity curves at the generic level and explore patterns of conodont paleogeographic distribution. Our results partly confirm the most prominent findings of earlier studies including the occurrence of an Ordovician acme, a Permian nadir and a short-lived Triassic recovery.

@article{doi101016jgloplacha2020103325,
    author = "Ginot, Samuel and Goudemand, Nicolas",
    title = "Global climate changes account for the main trends of conodont diversity but not for their final demise",
    year = "2020",
    journal = "Global and Planetary Change",
    abstract = "Conodonts, one of the longest-lived early groups of vertebrates, have a very complete fossil record ranging from the late Cambrian to the end of the Triassic and persisted through many global climatic and biotic events. In this paper, we analyse a large dataset harvested from the Paleobiology Database to compute global diversity curves at the generic level and explore patterns of conodont paleogeographic distribution. Our results partly confirm the most prominent findings of earlier studies including the occurrence of an Ordovician acme, a Permian nadir and a short-lived Triassic recovery.",
    url = "https://doi.org/10.1016/j.gloplacha.2020.103325",
    doi = "10.1016/j.gloplacha.2020.103325",
    openalex = "W3088285233",
    references = "hofmann2019diversity"
}

@article{doi101016jearscirev2021103503,
    author = "Scotese, Christopher R. and Song, Haijun and Mills, Benjamin and van der Meer, Douwe G.",
    title = "Phanerozoic paleotemperatures: The earth’s changing climate during the last 540 million years",
    year = "2021",
    journal = "Earth-Science Reviews",
    url = "https://doi.org/10.1016/j.earscirev.2021.103503",
    doi = "10.1016/j.earscirev.2021.103503",
    openalex = "W3120450552",
    references = "doi10100797894017960024, doi1010160016703789901506, doi101016003101828790040x, doi1010160031018292901825, doi101016jcretres200805025, doi101016jearscirev201305014, doi101016jepsl200905028, doi101016jgr201212026, doi101016jmargeo200502007, doi101016jpalaeo200606026, doi101016jpalaeo200911006, doi101016jpalaeo201005036, doi101016jpalaeo201409013, doi101016jpalaeo201611005, doi101016jpalaeo201703014, doi101016jpalwor200610016, doi101016s0009254199000819, doi101016s0009254199000844, doi101016s0012825200000374, doi101016s0012825202001046, doi101016s0012825299000483, doi101016s1631071303000063, doi101017cbo9780511628948, doi101017s0016756818000110, doi1010292009gc002788, doi101038333547a0, doi101038ncomms14845, doi101038s41467018039961, doi101038s4156101700036, doi101046j13653121200200408x, doi101073pnas1319253111, doi101086648217, doi101111j14754983201201165x, doi101126sciadvaaz1346, doi101126science1155814, doi101126science1161648, doi101126science1177265, doi101126science2064415217, doi101126science21545381351, doi101126science2845414616, doi1011300016760619637493sitcio20co2, doi10113000167606198596567defie20co2, doi1011300016760619991110960cisona23co2, doi1011300091761319880160022lctvan23co2, doi1011300091761319910190867ccapct23co2, doi1011302019254214, doi101130g315791, doi101130gsab471177, doi101146annurevea05050177001535, doi101146annurevearth040610133431, doi1015159781400862924, doi1016660094837320040300522oeamdo20co2, doi105194cp1714832021, doi105194cp76032011, doi105860choice435903, openalexw2106559152, openalexw2139291338, openalexw2989964553"
}

The decline in background extinction rates of marine animals through geologic time is an established but unexplained feature of the Phanerozoic fossil record. There is also growing consensus that the ocean and atmosphere did not become oxygenated to near-modern levels until the mid-Paleozoic, coinciding with the onset of generally lower extinction rates. Physiological theory provides us with a possible causal link between these two observations-predicting that the synergistic impacts of oxygen and temperature on aerobic respiration would have made marine animals more vulnerable to ocean warming events during periods of limited surface oxygenation. Here, we evaluate the hypothesis that changes in surface oxygenation exerted a first-order control on extinction rates through the Phanerozoic using a combined Earth system and ecophysiological modeling approach. We find that although continental configuration, the efficiency of the biological carbon pump in the ocean, and initial climate state all impact the magnitude of modeled biodiversity loss across simulated warming events, atmospheric oxygen is the dominant predictor of extinction vulnerability, with metabolic habitat viability and global ecophysiotype extinction exhibiting inflection points around 40% of present atmospheric oxygen. Given this is the broad upper limit for estimates of early Paleozoic oxygen levels, our results are consistent with the relative frequency of high-magnitude extinction events (particularly those not included in the canonical big five mass extinctions) early in the Phanerozoic being a direct consequence of limited early Paleozoic oxygenation and temperature-dependent hypoxia responses.

@article{doi101073pnas2101900118,
    author = "Stockey, Richard and Pohl, Alexandre and Ridgwell, Andy and Finnegan, Seth and Sperling, Erik A.",
    title = "Decreasing Phanerozoic extinction intensity as a consequence of Earth surface oxygenation and metazoan ecophysiology",
    year = "2021",
    journal = "Proceedings of the National Academy of Sciences",
    abstract = "The decline in background extinction rates of marine animals through geologic time is an established but unexplained feature of the Phanerozoic fossil record. There is also growing consensus that the ocean and atmosphere did not become oxygenated to near-modern levels until the mid-Paleozoic, coinciding with the onset of generally lower extinction rates. Physiological theory provides us with a possible causal link between these two observations-predicting that the synergistic impacts of oxygen and temperature on aerobic respiration would have made marine animals more vulnerable to ocean warming events during periods of limited surface oxygenation. Here, we evaluate the hypothesis that changes in surface oxygenation exerted a first-order control on extinction rates through the Phanerozoic using a combined Earth system and ecophysiological modeling approach. We find that although continental configuration, the efficiency of the biological carbon pump in the ocean, and initial climate state all impact the magnitude of modeled biodiversity loss across simulated warming events, atmospheric oxygen is the dominant predictor of extinction vulnerability, with metabolic habitat viability and global ecophysiotype extinction exhibiting inflection points around 40\% of present atmospheric oxygen. Given this is the broad upper limit for estimates of early Paleozoic oxygen levels, our results are consistent with the relative frequency of high-magnitude extinction events (particularly those not included in the canonical big five mass extinctions) early in the Phanerozoic being a direct consequence of limited early Paleozoic oxygenation and temperature-dependent hypoxia responses.",
    url = "https://doi.org/10.1073/pnas.2101900118",
    doi = "10.1073/pnas.2101900118",
    openalex = "W3204432017",
    references = "close2020the, doi101007bf00151270, doi101016jepsl200702018, doi1010292009gc002788, doi101038ncomms14845, doi101126science1155814, doi101126science1163156, doi101126science21545391501, doi101126scienceaaa1605, doi10120197802031805703, doi101242jeb037523"
}

Paleogeography is the study of the changing surface of Earth through time. Driven by plate tectonics, the configuration of the continents and ocean basins has been in constant flux. Plate tectonics pushes the land surface upward or pulls it apart, causing its collapse. All the while, the unrelenting forces of climate and weather slowly reduce mountains to sand and mud and redistribute these sediments to the sea. This article reviews the changing paleogeography of the past 750 million years. It describes the broad patterns of Phanerozoic paleogeography as well as many of the specific paleogeographic events that have shaped the modern continents and ocean basins. The focus is on the changing latitudinal distribution of the continents, fluctuations in sea level, the opening and closing of oceanic seaways, mountain building, and how these paleogeographic changes have affected global climate, ocean circulation, and the evolution of life. This review presents an atlas of 114 paleogeographic maps that illustrate how Earth's surface has evolved during the past 750 million years. During that time interval, Earth has witnessed the formation and breakup of two supercontinents: Pannotia and Pangea. The continents have been transformed from low-lying flooded platforms to high-standing land areas crisscrossed by the scars of past continental collisions. Oceans have opened and closed, and then opened again in a seemingly never-ending cycle. ▪ The changing configuration of the continents and ocean basins during the past 750 million years is illustrated in 114 paleogeographic maps. ▪ These maps describe how the surface of Earth has been continually modified by mountain building and erosion. ▪ The changing paleogeography has affected global climate, ocean circulation, and the evolution of life. ▪ The data and methods used to produce the maps are described in detail.

@article{doi101146annurevearth081320064052,
    author = "Scotese, Christopher R.",
    title = "An Atlas of Phanerozoic Paleogeographic Maps: The Seas Come In and the Seas Go Out",
    year = "2021",
    journal = "Annual Review of Earth and Planetary Sciences",
    abstract = "Paleogeography is the study of the changing surface of Earth through time. Driven by plate tectonics, the configuration of the continents and ocean basins has been in constant flux. Plate tectonics pushes the land surface upward or pulls it apart, causing its collapse. All the while, the unrelenting forces of climate and weather slowly reduce mountains to sand and mud and redistribute these sediments to the sea. This article reviews the changing paleogeography of the past 750 million years. It describes the broad patterns of Phanerozoic paleogeography as well as many of the specific paleogeographic events that have shaped the modern continents and ocean basins. The focus is on the changing latitudinal distribution of the continents, fluctuations in sea level, the opening and closing of oceanic seaways, mountain building, and how these paleogeographic changes have affected global climate, ocean circulation, and the evolution of life. This review presents an atlas of 114 paleogeographic maps that illustrate how Earth's surface has evolved during the past 750 million years. During that time interval, Earth has witnessed the formation and breakup of two supercontinents: Pannotia and Pangea. The continents have been transformed from low-lying flooded platforms to high-standing land areas crisscrossed by the scars of past continental collisions. Oceans have opened and closed, and then opened again in a seemingly never-ending cycle. ▪ The changing configuration of the continents and ocean basins during the past 750 million years is illustrated in 114 paleogeographic maps. ▪ These maps describe how the surface of Earth has been continually modified by mountain building and erosion. ▪ The changing paleogeography has affected global climate, ocean circulation, and the evolution of life. ▪ The data and methods used to produce the maps are described in detail.",
    url = "https://doi.org/10.1146/annurev-earth-081320-064052",
    doi = "10.1146/annurev-earth-081320-064052",
    openalex = "W3139147700",
    references = "doi101016jearscirev201203002, doi101016jearscirev2020103463, doi101016s0012821x0100588x, doi101029jb082i005p00803, doi101038267399a0, doi101038359117a0, doi101086628416, doi101126science1116412, doi101126science1156963, doi101126science1894201419, doi101126science23547931156, doi101126science27753341956, doi101126science28153811342, doi101146annurevearth32082503144359, doi101306m26490"
}

During the past 541 million years, marine animals underwent three intervals of diversification (early Cambrian, Ordovician, Cretaceous–Cenozoic) separated by nondirectional fluctuation, suggesting diversity-dependent dynamics with the equilibrium diversity shifting through time. Changes in factors such as shallow-marine habitat area and climate appear to have modulated the nondirectional fluctuations. Directional increases in diversity are best explained by evolutionary innovations in marine animals and primary producers coupled with stepwise increases in the availability of food and oxygen. Increasing intensity of biotic interactions such as predation and disturbance may have led to positive feedbacks on diversification as ecosystems became more complex. Important areas for further research include improving the geographic coverage and temporal resolution of paleontological data sets, as well as deepening our understanding of Earth system evolution and the physiological and ecological traits that modulated organismal responses to environmental change.

@article{doi101146annurevecolsys012021035131,
    author = "Bush, Andrew M. and Payne, Jonathan L.",
    title = "Biotic and Abiotic Controls on the Phanerozoic History of Marine Animal Biodiversity",
    year = "2021",
    journal = "Annual Review of Ecology Evolution and Systematics",
    abstract = "During the past 541 million years, marine animals underwent three intervals of diversification (early Cambrian, Ordovician, Cretaceous–Cenozoic) separated by nondirectional fluctuation, suggesting diversity-dependent dynamics with the equilibrium diversity shifting through time. Changes in factors such as shallow-marine habitat area and climate appear to have modulated the nondirectional fluctuations. Directional increases in diversity are best explained by evolutionary innovations in marine animals and primary producers coupled with stepwise increases in the availability of food and oxygen. Increasing intensity of biotic interactions such as predation and disturbance may have led to positive feedbacks on diversification as ecosystems became more complex. Important areas for further research include improving the geographic coverage and temporal resolution of paleontological data sets, as well as deepening our understanding of Earth system evolution and the physiological and ecological traits that modulated organismal responses to environmental change.",
    url = "https://doi.org/10.1146/annurev-ecolsys-012021-035131",
    doi = "10.1146/annurev-ecolsys-012021-035131",
    openalex = "W3194305746",
    references = "hofmann2019diversity"
}

Body size is an important species trait, correlating with life span, fecundity, and other ecological factors. Over Earth's geological history, climate shifts have occurred, potentially shaping body size evolution in many clades. General rules attempting to summarize body size evolution include Bergmann's rule, which states that species reach larger sizes in cooler environments and smaller sizes in warmer environments, and Cope's rule, which poses that lineages tend to increase in size over evolutionary time. Tetraodontiform fishes (including pufferfishes, boxfishes, and ocean sunfishes) provide an extraordinary clade to test these rules in ectotherms owing to their exemplary fossil record and the great disparity in body size observed among extant and fossil species. We examined Bergmann's and Cope's rules in this group by combining phylogenomic data (1,103 exon loci from 185 extant species) with 210 anatomical characters coded from both fossil and extant species. We aggregated data layers on paleoclimate and body size from the species examined, and inferred a set of time-calibrated phylogenies using tip-dating approaches for downstream comparative analyses of body size evolution by implementing models that incorporate paleoclimatic information. We found strong support for a temperature-driven model in which increasing body size over time is correlated with decreasing oceanic temperatures. On average, extant tetraodontiforms are two to three times larger than their fossil counterparts, which otherwise evolved during periods of warmer ocean temperatures. These results provide strong support for both Bergmann's and Cope's rules, trends that are less studied in marine fishes compared to terrestrial vertebrates and marine invertebrates.

@article{doi101073pnas2122486119,
    author = "Troyer, Emily M. and Betancur‐R, Ricardo and Hughes, Lily C. and Westneat, Mark W. and Carnevale, Giorgio and White, William T. and Pogonoski, John J. and Tyler, James C. and Baldwin, Carole C. and Ortı́, Guillermo and Brinkworth, Andrew and Clavel, Julien and Arcila, Dahiana",
    title = "The impact of paleoclimatic changes on body size evolution in marine fishes",
    year = "2022",
    journal = "Proceedings of the National Academy of Sciences",
    abstract = "Body size is an important species trait, correlating with life span, fecundity, and other ecological factors. Over Earth's geological history, climate shifts have occurred, potentially shaping body size evolution in many clades. General rules attempting to summarize body size evolution include Bergmann's rule, which states that species reach larger sizes in cooler environments and smaller sizes in warmer environments, and Cope's rule, which poses that lineages tend to increase in size over evolutionary time. Tetraodontiform fishes (including pufferfishes, boxfishes, and ocean sunfishes) provide an extraordinary clade to test these rules in ectotherms owing to their exemplary fossil record and the great disparity in body size observed among extant and fossil species. We examined Bergmann's and Cope's rules in this group by combining phylogenomic data (1,103 exon loci from 185 extant species) with 210 anatomical characters coded from both fossil and extant species. We aggregated data layers on paleoclimate and body size from the species examined, and inferred a set of time-calibrated phylogenies using tip-dating approaches for downstream comparative analyses of body size evolution by implementing models that incorporate paleoclimatic information. We found strong support for a temperature-driven model in which increasing body size over time is correlated with decreasing oceanic temperatures. On average, extant tetraodontiforms are two to three times larger than their fossil counterparts, which otherwise evolved during periods of warmer ocean temperatures. These results provide strong support for both Bergmann's and Cope's rules, trends that are less studied in marine fishes compared to terrestrial vertebrates and marine invertebrates.",
    url = "https://doi.org/10.1073/pnas.2122486119",
    doi = "10.1073/pnas.2122486119",
    openalex = "W4285012276",
    references = "doi101016jcub202107071"
}

Abstract Owing to the increasing availability of data for many fossil groups and a generally accepted palaeogeographical configuration, palaeontologists have been able to develop progressively more robust palaeobiogeographical scenarios for the spatial distributions of Ordovician marine faunas. However, most research in Early Paleozoic palaeobiogeography centres on data derived from extensively studied localities in North America and Europe. Thus, clear patterns are emerging of regional biogeography for these areas. However, the fragmentary nature of data from other regions hinders the development of a detailed understanding of palaeogeographical schemes of many clades at the global level. Provincial patterns are now available for several fossil groups, but the global coverage remains generally fragmentary. Palaeobiogeographical investigations were traditionally focused on better understanding of palaeogeographical scenarios and often employed quantitative analyses of faunal similarity. More recently palaeobiogeographical analyses have expanded to investigate questions such as the location and pace of speciation and macroevolution together with macroecological change. For example, studies on the evolution of speciation levels in the frame of the taxonomic radiation of the Great Ordovician Biodiversification are now available. Future investigations, including modelling, will provide more integrative, global patterns of provincialism, including the location of Ordovician biodiversity hotspots and the recognition of latitudinal diversity gradients.

@article{doi101144sp5322022168,
    author = "Servais, Thomas and Harper, David A. T. and Kröger, Björn and Scotese, Christopher R. and Stigall, Alycia L. and Zhen, Yong Yi",
    title = "Changing palaeobiogeography during the Ordovician Period",
    year = "2022",
    journal = "Geological Society London Special Publications",
    abstract = "Abstract Owing to the increasing availability of data for many fossil groups and a generally accepted palaeogeographical configuration, palaeontologists have been able to develop progressively more robust palaeobiogeographical scenarios for the spatial distributions of Ordovician marine faunas. However, most research in Early Paleozoic palaeobiogeography centres on data derived from extensively studied localities in North America and Europe. Thus, clear patterns are emerging of regional biogeography for these areas. However, the fragmentary nature of data from other regions hinders the development of a detailed understanding of palaeogeographical schemes of many clades at the global level. Provincial patterns are now available for several fossil groups, but the global coverage remains generally fragmentary. Palaeobiogeographical investigations were traditionally focused on better understanding of palaeogeographical scenarios and often employed quantitative analyses of faunal similarity. More recently palaeobiogeographical analyses have expanded to investigate questions such as the location and pace of speciation and macroevolution together with macroecological change. For example, studies on the evolution of speciation levels in the frame of the taxonomic radiation of the Great Ordovician Biodiversification are now available. Future investigations, including modelling, will provide more integrative, global patterns of provincialism, including the location of Ordovician biodiversity hotspots and the recognition of latitudinal diversity gradients.",
    url = "https://doi.org/10.1144/sp532-2022-168",
    doi = "10.1144/sp532-2022-168",
    openalex = "W4311053344",
    references = "doi101111pala12401, hofmann2019diversity"
}

Abstract Siliceous marine ecosystems play a critical role in shaping the Earth's climate system by influencing rates of organic carbon burial and marine authigenic clay formation (i.e., reverse weathering). The ecological demise of silicifying organisms associated with the Permian‐Triassic mass extinction is postulated to have elevated marine authigenic clay formation rates, resulting in a prolonged greenhouse climate during the Early Triassic. Yet, our understanding of the response of siliceous marine organisms during this critical interval is poor. Whilst radiolarians experienced the strongest diversity loss in their evolutionary history and perhaps also the greatest population decline of silica‐secreting organisms during this event, only a small number of Griesbachian (post‐extinction) localities that record siliceous organisms are known. Here, we report newly discovered latest Changhsingian to early Griesbachian (Clarkina meishanensis ‐ Hindeodus parvus Zone) radiolarians and siliceous sponge spicules from Svalbard. This fauna documents the survival of a low‐diversity radiolarian assemblage alongside stem‐group hexactinellid sponges making this the first described account of post‐extinction silica‐secreting organisms from the Permian/Triassic boundary in a shallow marine shelf environment and a mid‐northern paleolatitudinal setting. Our findings indicate that latitudinal diversity gradients for silica‐secreting organisms following the mass extinction were significantly altered, and that silica productivity was restricted to high latitude and deep water thermal refugia. This result has potential to further shape our understanding of changes in marine dissolved silica levels and in turn rates of reverse weathering, with implications for our understanding of carbon cycle dynamics during this interval.

@article{doi1010292023pa004766,
    author = "Foster, William J. and Asatryan, G. and Rauzi, Sofia and Botting, Joseph P. and Buchwald, Stella Z. and Lazarus, David and Isson, Terry T. and Renaudie, Johan and Kiessling, Wolfgang",
    title = "Response of Siliceous Marine Organisms to the Permian‐Triassic Climate Crisis Based on New Findings From Central Spitsbergen, Svalbard",
    year = "2023",
    journal = "Paleoceanography and Paleoclimatology",
    abstract = "Abstract Siliceous marine ecosystems play a critical role in shaping the Earth's climate system by influencing rates of organic carbon burial and marine authigenic clay formation (i.e., reverse weathering). The ecological demise of silicifying organisms associated with the Permian‐Triassic mass extinction is postulated to have elevated marine authigenic clay formation rates, resulting in a prolonged greenhouse climate during the Early Triassic. Yet, our understanding of the response of siliceous marine organisms during this critical interval is poor. Whilst radiolarians experienced the strongest diversity loss in their evolutionary history and perhaps also the greatest population decline of silica‐secreting organisms during this event, only a small number of Griesbachian (post‐extinction) localities that record siliceous organisms are known. Here, we report newly discovered latest Changhsingian to early Griesbachian (Clarkina meishanensis ‐ Hindeodus parvus Zone) radiolarians and siliceous sponge spicules from Svalbard. This fauna documents the survival of a low‐diversity radiolarian assemblage alongside stem‐group hexactinellid sponges making this the first described account of post‐extinction silica‐secreting organisms from the Permian/Triassic boundary in a shallow marine shelf environment and a mid‐northern paleolatitudinal setting. Our findings indicate that latitudinal diversity gradients for silica‐secreting organisms following the mass extinction were significantly altered, and that silica productivity was restricted to high latitude and deep water thermal refugia. This result has potential to further shape our understanding of changes in marine dissolved silica levels and in turn rates of reverse weathering, with implications for our understanding of carbon cycle dynamics during this interval.",
    url = "https://doi.org/10.1029/2023pa004766",
    doi = "10.1029/2023pa004766",
    openalex = "W4389219389",
    references = "doi101111gcb16333"
}

Abstract Ancient changes in the biosphere, from organismic traits to wholesale ecosystem changes, can be aligned with climate forcing across the Phanerozoic. Clear examples of abrupt climate warming causing biodiversity crises are primarily found between the Permian and Paleogene periods. During these times, catastrophic events occurred, resembling the extreme climate scenarios projected for the near future. The paleobiologic literature around these events generally supports the hypothesis that abrupt climate change was a dominant trigger of extinction and/or ecological crisis. When climate change and climate history are considered, virtually all post-Paleozoic global biotic events can be confidently attributed to climatic change, with abrupt warming (hyperthermal events) leaving the most consistent fingerprint. The combined stress of deoxygenation and warming are sufficient to explain marine extinction patterns across most hyperthermal events. Although ocean acidification may have contributed, the direct role of pH on the extinction toll of organisms is not consistently demonstrated. Future research can enhance the correspondence between the magnitudes of climatic changes and their biological impacts, even though observed rates of change cannot currently be compared across different timescales. Mimicking multi-scale approaches in modern ecology, paleontological approaches to climate impact research will benefit from specifically targeting scaling relationships.

@article{doi101017pab202420,
    author = "Kiessling, Wolfgang and Reddin, Carl J. and Dowding, Elizabeth M. and Dimitrijević, Danijela and Raja, Nussaïbah B. and Kocsis, Ádám T.",
    title = "Marine biological responses to abrupt climate change in deep time",
    year = "2024",
    journal = "Paleobiology",
    abstract = "Abstract Ancient changes in the biosphere, from organismic traits to wholesale ecosystem changes, can be aligned with climate forcing across the Phanerozoic. Clear examples of abrupt climate warming causing biodiversity crises are primarily found between the Permian and Paleogene periods. During these times, catastrophic events occurred, resembling the extreme climate scenarios projected for the near future. The paleobiologic literature around these events generally supports the hypothesis that abrupt climate change was a dominant trigger of extinction and/or ecological crisis. When climate change and climate history are considered, virtually all post-Paleozoic global biotic events can be confidently attributed to climatic change, with abrupt warming (hyperthermal events) leaving the most consistent fingerprint. The combined stress of deoxygenation and warming are sufficient to explain marine extinction patterns across most hyperthermal events. Although ocean acidification may have contributed, the direct role of pH on the extinction toll of organisms is not consistently demonstrated. Future research can enhance the correspondence between the magnitudes of climatic changes and their biological impacts, even though observed rates of change cannot currently be compared across different timescales. Mimicking multi-scale approaches in modern ecology, paleontological approaches to climate impact research will benefit from specifically targeting scaling relationships.",
    url = "https://doi.org/10.1017/pab.2024.20",
    doi = "10.1017/pab.2024.20",
    openalex = "W4405279981",
    references = "doi1010179781316711644, doi101111gcb16333"
}

The latitudinal biodiversity gradient (LBG) is a fundamental biological pattern seen across taxa and ecosystems today, but its drivers remain uncertain despite intense study. Palaeontological data may add valuable evidence from diversity distributions during intervals with different Earth system configurations, including potential analogues of future climate regimes. However, accurately characterizing these distributions is challenging because the geographic scope of fossil record coverage varies through time, introducing biases that have not been quantified by most previous studies. Here, we attempt a comprehensive documentation of latitudinal biodiversity distributions of marine invertebrates through the past 540 million years, explicitly accounting for regional variation in diversity and sampling. We demonstrate large uncertainties when using current fossil data at this scale. Nevertheless, some signals are detectable. We show that marine animal biodiversity declined with increasing palaeolatitude and with decreasing temperature during at least some intervals from the Permian onwards (298.9 Ma). Additionally, we find that the LBG was shallower on average when Earth's climate was hotter, although this signal is weak. We also document a strong, systematic bias due to intense sampling of the fossil record in North America and especially Europe, which may have led previous studies to incorrectly infer a mid‐latitude diversity peak during warm intervals of Earth history. Our results provide a baseline for what current fossil databases might tell us about Phanerozoic LBGs of marine animals, and suggests that quantitative evaluation of uncertainties and systematic bias will be central to advancing knowledge of geographic variation in diversity through Earth's history.

@article{benson2025marine,
    author = "Benson, Roger B.J. and Close, Roger A. and Antell, Gawain T. and Whittaker, Robert J. and Valdes, Paul and Farnsworth, Alex and Lunt, Daniel J. and Shen, Shuzhong and Fan, Junxuan and Saupe, Erin E.",
    title = "Marine animal diversity across latitudinal and temperature gradients during the Phanerozoic",
    year = "2025",
    journal = "Palaeontology",
    abstract = "The latitudinal biodiversity gradient (LBG) is a fundamental biological pattern seen across taxa and ecosystems today, but its drivers remain uncertain despite intense study. Palaeontological data may add valuable evidence from diversity distributions during intervals with different Earth system configurations, including potential analogues of future climate regimes. However, accurately characterizing these distributions is challenging because the geographic scope of fossil record coverage varies through time, introducing biases that have not been quantified by most previous studies. Here, we attempt a comprehensive documentation of latitudinal biodiversity distributions of marine invertebrates through the past 540 million years, explicitly accounting for regional variation in diversity and sampling. We demonstrate large uncertainties when using current fossil data at this scale. Nevertheless, some signals are detectable. We show that marine animal biodiversity declined with increasing palaeolatitude and with decreasing temperature during at least some intervals from the Permian onwards (298.9 Ma). Additionally, we find that the LBG was shallower on average when Earth's climate was hotter, although this signal is weak. We also document a strong, systematic bias due to intense sampling of the fossil record in North America and especially Europe, which may have led previous studies to incorrectly infer a mid‐latitude diversity peak during warm intervals of Earth history. Our results provide a baseline for what current fossil databases might tell us about Phanerozoic LBGs of marine animals, and suggests that quantitative evaluation of uncertainties and systematic bias will be central to advancing knowledge of geographic variation in diversity through Earth's history.",
    url = "https://doi.org/10.1111/pala.70006",
    doi = "10.1111/pala.70006",
    number = "3",
    volume = "68"
}

Reefs are important hotspots of marine biodiversity today and have acted as cradles of diversification in the geological past. However, we know little about how the diversity of reef-supporting regions varied through deep time, and how this differed from other regions. We quantified regional diversity patterns in reef-supporting and non-reef-supporting regions in the fossil record of Phanerozoic marine invertebrates. Diversity in reef-supporting regions is on average two- to threefold higher than in non-reef-supporting regions and has been remarkably stable over timescales of tens to hundreds of millions of years. This signal is present in both reefal and non-reefal facies within reef-supporting regions, suggesting that reefs enriched diversity in surrounding environments. Sepkoski's "Modern Fauna," an assemblage of higher taxa that includes gastropods, bivalves, and echinoids, has been a key component of reef-supporting regions since the Paleozoic, contrasting with its later rise to dominance in non-reef-supporting regions during the later Mesozoic-Cenozoic.

@article{doi101126sciadvadv9793,
    author = "Close, Roger A. and Benson, Roger and Kiessling, Wolfgang and Saupe, Erin E.",
    title = "Reefal regions were biodiversity hotspots throughout the Phanerozoic",
    year = "2025",
    journal = "Science Advances",
    abstract = {Reefs are important hotspots of marine biodiversity today and have acted as cradles of diversification in the geological past. However, we know little about how the diversity of reef-supporting regions varied through deep time, and how this differed from other regions. We quantified regional diversity patterns in reef-supporting and non-reef-supporting regions in the fossil record of Phanerozoic marine invertebrates. Diversity in reef-supporting regions is on average two- to threefold higher than in non-reef-supporting regions and has been remarkably stable over timescales of tens to hundreds of millions of years. This signal is present in both reefal and non-reefal facies within reef-supporting regions, suggesting that reefs enriched diversity in surrounding environments. Sepkoski's "Modern Fauna," an assemblage of higher taxa that includes gastropods, bivalves, and echinoids, has been a key component of reef-supporting regions since the Paleozoic, contrasting with its later rise to dominance in non-reef-supporting regions during the later Mesozoic-Cenozoic.},
    url = "https://doi.org/10.1126/sciadv.adv9793",
    doi = "10.1126/sciadv.adv9793",
    openalex = "W4415935258",
    references = "benson2025marine, doi101038nature21707, doi101038nature22901, doi101071mf99078, doi101093biomet4034237, doi10111114678721ep10768783, doi1011112041210x12613, doi101126science1059199, doi101126science1085706, doi101126science1152509, doi1018901119521"
}