Phanerozoic

@article{raup1972taxonomic,
    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",
    number = "4054",
    pages = "1065-1071",
    volume = "177"
}

@misc{cloud1977new1,
    author = "Cloud, P. E. and Morrison, K. and Lo, S. O",
    title = "New late pre-Phanerozoic and earliest Phanerozoic (?) microbiotas from Eastern Siberia",
    year = "1977",
    howpublished = "Geological Society of America, Abstracts with Programs, v. 9, no. 12",
    note = "talkorigins\_source = {true}; raw\_reference = {Cloud, P. E., Morrison, K., and Lo, S. O., 1977, New late pre-Phanerozoic and earliest Phanerozoic (?) microbiotas from Eastern Siberia: Geological Society of America, Abstracts with Programs, v. 9, no. 12.}"
}

@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{hallam1977secular2,
    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.}"
}

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"
}

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"
}

@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.}"
}

@incollection{sepkoski1992biotic,
    author = "Sepkoski, J. John and Schopf, J. William",
    title = "Biotic Diversity and Rates of Evolution During Proterozoic and Earliest Phanerozoic Time",
    year = "1992",
    booktitle = "The Proterozoic Biosphere",
    url = "https://doi.org/10.1017/cbo9780511601064.013",
    doi = "10.1017/cbo9780511601064.013",
    pages = "521-566"
}

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"
}

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"
}

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"
}