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29 April 2025 Terrestrialization: toward a shared framework for ecosystem evolution
C. Kevin Boyce, Matthew P. Nelsen
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Abstract

The Paleozoic evolution of a complex terrestrial biota has been among the most important events in Earth history. Here, we synthesize paleontological and neontological information across the different threads of the biota—including microbial life, fungi, animals, and plants—addressing discrepancies between the fossil record and time-calibrated molecular phylogenies. Four fundamental patterns are emphasized: (1) Most terrestrial animal lineages consist of diminutive inhabitants of soil and litter, with the soil fauna exhibiting remarkable continuity between the Paleozoic and present. (2) Faunal evolution tracks the ecological opportunities afforded by the evolution of the land flora. Flora and fauna alike were initially confined to the thin interface between soil and air, but animals explored both flight and burrowing as vascular plant size increased to encompass tree stature and deep rooting. (3) Skewed nutrient ratios of land plants present a fundamental challenge for animals that are accommodated through contrasting size-based dietary strategies. Detritivory and cell-by-cell herbivory are the diets most readily available for primary consumers but impose limits on the largest possible body sizes; only with subsequent evolution of herbivory in insects and then vertebrates could the dramatic increases in size in the Permian and Mesozoic have been achieved. (4) A second pulse of animal terrestrializations is apparent in the Cretaceous and Cenozoic that might be attributed to increased terrestrial productivity associated with angiosperm evolution. However, environmental changes to nutrient availability earlier in the Mesozoic prevent an unambiguous causal attribution, and the pulse may just be an artifact of our modern vantage point.

The Paleozoic evolution of a complex terrestrial biota has been among the most important events in Earth history. Here, we address how to integrate information from fossil and extant life highlighting four fundamental patterns: (1) The soil fauna shows remarkable continuity over 400 million years through to the present. (2) The evolution of animal ecologies closely tracks the opportunities provided by plant evolution via flight and burrowing as plants evolved the canopy stature and rooting depth of trees. (3) The skewed nutrient ratios of plants are a challenge that can be avoided by means of small body size, but that then places limits on the maximum body sizes seen before the evolution of insect and vertebrate herbivores. (4) The most recent 150 million years suggest a second pulse of animal terrestrializations, perhaps related to the evolution of flowering plants, but this linkage is questionable.

C. Kevin Boyce and Matthew P. Nelsen "Terrestrialization: toward a shared framework for ecosystem evolution," Paleobiology 51(1), 174-194, (29 April 2025). https://doi.org/10.1017/pab.2024.15
Received: 4 December 2023; Accepted: 25 April 2024; Published: 29 April 2025
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