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Ecosystem structure—that is the species present, the functions they represent, and how those functions interact—is an important determinant of community stability. This in turn affects how ecosystems respond to natural and anthropogenic crises, and whether species or the ecological functions that they represent are able to persist. Here we use fossil data from museum collections, literature, and the Paleobiology Database to reconstruct trophic networks of Tethyan paleocommunities from the Anisian and Carnian (Triassic), Bathonian (Jurassic), and Aptian (Cretaceous) stages, and compare these to a previously reconstructed trophic network from a modern Jamaican reef community. We generated model food webs consistent with functional structure and taxon richnesses of communities, and compared distributions of guild level parameters among communities, to assess the effect of the Mesozoic Marine Revolution on ecosystem dynamics. We found that the trophic space of communities expanded from the Anisian to the Aptian, but this pattern was not monotonic. We also found that trophic position for a given guild was subject to variation depending on what other guilds were present in that stage. The Bathonian showed the lowest degree of trophic omnivory by top consumers among all Mesozoic networks, and was dominated by longer food chains. In contrast, the Aptian network displayed a greater degree of short food chains and trophic omnivory that we attribute to the presence of large predatory guilds, such as sharks and bony fish. Interestingly, the modern Jamaican community appeared to have a higher proportion of long chains, as was the case in the Bathonian. Overall, results indicate that trophic structure is highly dependent on the taxa and ecological functions present, primary production experienced by the community, and activity of top consumers. Results from this study point to a need to better understand trophic position when planning restoration activities because a community may be so altered by human activity that restoring a species or its interactions may no longer be possible, and alternatives must be considered to restore an important function. Further work may also focus on elucidating the precise roles of top consumers in moderating network structure and community stability. 

Predation traces found on fossilized prey remains can be used to quantify the evolutionary history of biotic interactions. Fossil mollusc shells bearing these types of traces provided key evidence for the rise of predation during the Mesozoic marine revolution (MMR), an event thought to have reorganized global marine ecosystems. However, predation pressure on prey groups other than molluscs has not been explored adequately. Consequently, the ubiquity, tempo and synchronicity of the MMR cannot be thoroughly assessed. Here, we expand the evolutionary record of biotic interactions by compiling and analysing a new comprehensively collected database on drilling predation in Meso-Cenozoic echinoids. Trends in drilling frequency reveal an Eocene rise in drilling predation that postdated echinoid infaunalization and the rise in mollusc-targeted drilling (an iconic MMR event) by approximately 100 Myr. The temporal lag between echinoid infaunalization and the rise in drilling frequencies suggests that the Eocene upsurge in predation did not elicit a coevolutionary or escalatory response. This is consistent with rarity of fossil samples that record high frequency of drilling predation and scarcity of fossil prey recording failed predation events. These results suggest that predation intensification associated with the MMR was asynchronous across marine invertebrate taxa and represented a long and complex process that consisted of multiple uncoordinated steps probably with variable coevolutionary responses. 

A thorough understanding of how communities respond to extreme changes, such as biotic invasions, is essential to manage ecosystems today. Here we constructed fossil food webs to identify changes in Late Ordovician (Katian) shallow-marine paleocommunity structure and functioning before and after the Richmondian invasion, a well-documented ancient invasion. Food webs were compared using descriptive metrics and cascading extinction on graphs models. Richness at intermediate trophic levels was underrepresented when using only data from the Paleobiology Database relative to museum collections, resulting in a spurious decrease in modeled paleocommunity stability. Therefore, museum collections and field sampling may provide more reliable sources of data for the reconstruction of trophic organization in comparison to online data repositories. The invasion resulted in several changes in ecosystem dynamics. Despite topological similarities between pre- and postinvasion food webs, species loss occurred corresponding to a minor decrease in functional groups. Invaders occupied all of the preinvasion functional guilds, with the exception of four incumbent guilds that were lost and one new guild, corroborating the notion that invaders replace incumbents and fill preexisting niche space. Overall, models exhibited strong resistance to secondary extinction, although the postinvasion community had a lower threshold of collapse and more variable response to perturbation. We interpret these changes in dynamics as a decrease in stability, despite similarities in overall structure. Changes in food web structure and functioning resulting from the invasion suggest that conservation efforts may need to focus on preserving functional diversity if more diverse ecosystems are not inherently more stable. 

ABSTRACT Interactions with predators and parasites can result in traces found on Recent and fossil echinoids. However, identifying specific trace makers, particularly on fossil echinoids, remains contentious. To document the range of trace morphologies present on echinoids and improve our ability to identify and quantify biotic interactions affecting echinoids, we characterized traces found on fossil echinoids using museum collections and field sampling spanning the Jurassic to Recent worldwide. Using light microscopy, 8,564 individual echinoid specimens were examined including 130 species, and 516 traces of potential biotic interactions identified. Morphological characteristics were recorded for each trace, including the shape of the trace outline, maximum diameter and cross-section profile. Based on shared morphological characteristics, it was possible to classify all traces into eight categories: circular, subcircular, elongated, irregular, rectangular, figure-eight, notched, and linear. Cross-section characteristics provided additional insights into the identity of potential trace makers. To further evaluate the proposed biotic origins of these traces, trace diversity was examined through time and compared with anticipated ecological trends associated with the diversification of echinoids, and their predators and parasites. Trace diversity increased over time, starting in the late Eocene, coincident with the proliferation of echinoid-drilling gastropods, an indication that biotic interactions intensified through evolutionary time, as predicted by several macroevolutionary hypotheses previously tested using mollusks. The morphological descriptions provided here enhance our understanding of biotic traces on fossil echinoids, and the potential to identify temporal trends in the intensity and diversity of biotic interactions that have affected echinoids throughout their evolutionary history.