During the Ordovician, trilobites of the order Harpetida and the superfamily Trinucleioidea evolved unusual cephalic brims, quite unlike any structure known in modern arthropods. Brimmed trilobites were diverse and widespread, but we still do not fully understand why the harpiform brim evolved or what role it played in their success. Many authors have speculated about the brim's function, generating many untested hypotheses. We tested the hypothesis that the brim evolved to prevent sinking in soft sediments by calculating the depth to which brimless trilobites would sink and showed this explanation to be untenable. We then three-dimensionally printed model cephala with various brim shapes and moved them through natural sediments, testing the hypothesis that the brim evolved as a sediment plough. We found that increased horizontal brim width allowed a cephalon to efficiently displace more sediment, while increased brim height merely impeded the cephalon's progress. However, there is no clear evolutionary trend among harpetids or trinucleids towards wider or flatter brims, indicating that ploughing did not exert a strong selective pressure on brimmed trilobites. This work also serves as a case study in researching taxa without modern biomechanical analogues, demonstrating that many functional hypotheses can be adequately tested by straightforward experimental methods.

The physical, biogeochemical, and ecological properties of the modern seafloor are extensively shaped by the activities of burrowing and sediment-mixing animals, processes collectively known as bioturbation. Bioturbation is primarily recorded by homogenized sediments of the seafloor mixed layer and the underlying transition layer of discrete burrows. Although these two zones can be readily measured today, there has been limited understanding of how the mixed and transition layers evolved over the Phanerozoic since animals first began to extensively colonize the seafloor. Here, we provide a record for the depths of the sedimentary mixed and transition layers through the Phanerozoic. We find that although deepening of the sediment mixed layer spanned hundreds of millions of years, a deep transition layer was established as early as the Cambrian and did not further deepen until the Mesozoic-trajectories reflecting evolutionary radiations, changes in nutrient cycling, and alleviation of oxygen stress.

Both natural and human-induced stressors cause reef erosion, resulting in reef rubble formation. When consolidated, the rubble can facilitate reef recovery, sparking interest in artificial rubble stabilization as a method for reef restoration. However, our understanding of the natural processes governing coral reef regeneration within rubble beds is limited. This study examines the regeneration processes within ancient rubble frameworks in a Late Triassic carbonate platform. Results show that Late Triassic rubble environments exhibit successional trajectories similar to contemporary rubble environments. Key organisms such as sponges, calcareous red algae, bryozoans, microbes and scleractinian corals, which are instrumental in the consolidation of modern reef rubble, appear to have played comparable roles during the Late Triassic. The similarities between Late Triassic and modern reef rubble consolidation highlight enduring ecological mechanisms important for reef regeneration. This study deepens our understanding of reef dynamics and offers valuable insights for improving current reef restoration strategies, grounded in time-tested natural processes.

The Paleozoic Era was an interval of profound changes in metazoan reef ecosystems, which became magnificent in scale and diversity, owing to the evolution of various skeletal reef builders especially corals and sponges. This special issue comprises 16 papers that improve our understanding on the complex evolution and paleoecology of corals and metazoan reef ecosystems, and their response to environmental changes through the Cambrian to Permian. The case studies present detailed evidence that helps to reconstruct the complicated interactions among reef organisms, the succession of metazoan reefs, and their relations to paleoenvironments. Some key findings address (i) Cambrian reefs and microbe-metazoan reef transition to the Ordovician, (ii) reef component and paleoecological features of Siluro-Devonian coral-stromatoporoid reefs, and (iii) growth characteristics, paleogeography, and reef formation of Late Paleozoic corals and sponges. As such, this special issue will stimulate further work on reef studies and be of interest to a broad community of paleoecologists, to scholars in various fields of the geosciences as well as to reef biologists.

Living coral cover is rapidly decreasing due to global climate change and local human impacts. Response to the ongoing transformation of coral reefs requires a deep understanding of the mechanisms that regulate community assembly and sustain biodiversity in these systems. The goal of this study is to test the persistence of Devonian reef communities across 70 sea-level fluctuation cycles. Abundance data were collected for in situ coral and calcareous sponge taxa found along 164 transects across two field localities within the Canning Basin, Western Australia. The Bray-Curtis dissimilarity index was used to quantify the dissimilarity in taxonomic composition among transects. Despite the notable persistence of reef communities over numerous cycles (ranging from 22 to 29 cycles) spanning hundreds of thousands of years, significant taxonomic shifts unfolded over millions of years, primarily driven by the diversity and composition of coral assemblages. This indicates that while reef communities exhibit persistence over intermediate time scales, they undergo changes in taxonomic composition over extended periods. The observed shifts in taxonomic composition likely reflect the profound changes in Earth-surface systems that occurred during the Devonian (i.e., fluctuations in global sea-surface temperatures and sea-level). Therefore, sustained abiotic processes can act to disrupt community structure over extended time scales as the threshold for community persistence is exceeded. This study helps establish baseline community composition dynamics in systems devoid of human influence and thus deepens our understanding of naturally occurring phase shifts. Understanding the factors that contribute to the persistence of community assembly across multiple scales is crucial for conserving coral reef ecosystems in the face of ongoing environmental changes.

The geologic history of dissolved silica concentration in the ocean (DSi) is central to understanding the evolution of silica biomineralization, the interactions between the global carbon and silicon cycles, and their combined role controlling global climate over geologic time. However, the silica cycle in the geologic past is under-constrained, especially during major mass extinction events that impacted biosilicifiers and were associated with dramatic climate change. We measured the silicon isotope ratios (δ30Si) of 76 sponge spicules from the Panthalassic Ocean spanning the Triassic–Jurassic boundary (ca. 201 Ma) to constrain DSi concentrations during the mid-Mesozoic. Spicule measurements have mean δ30Si values of –0.25‰ ± 0.99‰. Our data, combined with constraints on seawater δ30Si from coeval radiolarians, suggest that mid-Mesozoic DSi was between 20–100 µM, a similar range to the modern ocean. Our results support increasing evidence that by the Mesozoic DSi had already decreased by orders of magnitude relative to the Precambrian. These results imply that radiolarians and sponges were drawing down DSi prior to diatom ecological dominance. Increasing sponge δ30Si values across the Triassic–Jurassic boundary, coupled with modeling evidence and previous palaeoecological observations, support that warming, increased weathering, and Si delivery before the end-Triassic extinction may have facilitated sponge expansion during the extinction recovery interval.

A three-dimensional tubular fabric known as "vermiform microstructure" in Phanerozoic and Neoproterozoic carbonate microbialites has been hypothesized to represent the body fossil of nonspicular keratose demosponges. If correct, this interpretation extends the sponge body fossil record and origin of animals to ~890 Ma. However, the veracity of the keratose sponge interpretation for vermiform microstructure remains in question, and the origin of the tubular fabric is enigmatic. Here we compare exceptionally well-preserved microbialite textures from the Upper Triassic to channel networks created by modern microbial biofilms. We demonstrate that anastomosing channel networks of similar size and geometries are produced by microbial biofilms in the absence of sponges, suggesting the origin for vermiform microstructure in ancient carbonates is not unique to sponges and perhaps best interpreted conservatively as likely microbial in origin. We present a taphonomic model of early biofilm lithification in seawater with anomalously high carbonate saturation necessary to preserve delicate microbial textures. This work has implications for the understanding of three-dimensional biofilm architecture that goes beyond the current micro-scale observations available from living biofilm experiments and suggests that biofilm channel networks have an extensive fossil record.

Harpetid and trinucleid trilobites share a similar and unusual morphology, the most striking feature of which is a wide, flattened cephalic brim with many pits or holes. This similarity was once interpreted as a sign that these two groups of trilobites were closely related, but in recent years it has instead been assumed that the 'harpiform' brim arose in both groups independently. However, relatedness and similarity can be difficult to disentangle in fossil taxa without close living relatives, and this assumption about the harpiform brim has never been explicitly tested. Our study re-evaluates the relationship between Harpetida and Trinucleioidea in order to test a longstanding assumption about trilobite relationships and as a case study in evaluating different kinds of morphological similarity in extinct groups. We inferred a new phylogenetic tree using parsimony methods and discrete morphological character data from a broad sampling of harpetids, trinucleids, and their relatives. Despite their gross morphological similarities, we found that harpetids and trinucleids were readily distinguished in our analyses, a result consistent with a hypothesis of multiple origins for the harpiform brim. By mapping brim-related characters across our new phylogeny, we identified a sequence of morphological innovations that arose in parallel in both groups and led ultimately in each case to the evolution of the harpiform brim. These results indicate that harpiform brims are a prime example of parallel evolution-the similar development of a morphological trait in distantly related taxa that nevertheless share a similar original morphology. In addition, our phylogeny supports the idea that trinucleids are specialized, harpiform asaphids, rather than an independent order of trilobites. We also provide new information on the relationships of the putative 'basal-most' members of Trinucleioidea, the Liostracinidae, and confirm recent assessments that this family is more distantly related to trinucleids.

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Fossil deposits with exceptional preservation ("lagerstätten") provide important details not typically preserved in the fossil record, such that they hold an outsized influence on our understanding of biodiversity and evolution. In particular, the potential bias imparted by this so-called "lagerstätten effect" remains a critical, but underexplored aspect of reconstructing evolutionary relationships. Here, we quantify the amount of phylogenetic information available in the global fossil records of 1,327 species of non-avian theropod dinosaurs, Mesozoic birds, and fossil squamates (e.g., lizards, snakes, mosasaurs), and then compare the influence of lagerstätten deposits on phylogenetic information content and taxon selection in phylogenetic analyses to other fossil-bearing deposits. We find that groups that preserve a high amount of phylogenetic information in their global fossil record (e.g., non-avian theropods) are less vulnerable to a "lagerstätten effect" that leads to disproportionate representation of fossil taxa from one geologic unit in an evolutionary tree. Additionally, for each taxonomic group, we find comparable amounts of phylogenetic information in lagerstätten deposits, even though corresponding morphological character datasets vary greatly. Finally, we unexpectedly find that ancient sand dune deposits of the Late Cretaceous Gobi Desert of Mongolia and China exert an anomalously large influence on the phylogenetic information available in the squamate fossil record, suggesting a "lagerstätten effect" can be present in units not traditionally considered lagerstätten. These results offer a phylogenetics-based lens through which to examine the effects of exceptional fossil preservation on biological patterns through time and space, and invites further quantification of evolutionary information in the rock record.

The defining feature of a mass extinction and its associated recovery is the global amount of extinction that occurred[1]. However, there is finer-scale detail to the story, as seen in the ongoing modern mass extinction（the sixth mass extinction）, which exhibits variable effects among marine ecosystems and environments, and hence geographic areas [1].

Modern coral reefs and associated biodiversity are severely threatened by increasing terrestrial runoff. Similar scenarios could be suspected for geological times, but reef coral resilience is still an enigma. In late Visean-Serpukhovian (Mississippian foraminiferal zones/MFZ 14-16) times, a major glaciation phase of the late Palaeozoic Ice Age (LPIA) associated with enhanced terrestrial weathering and runoff coincides with a biodiversity crisis and coral reef decline. In this study, the impact of enhanced terrestrial runoff is tested on size variations of colonial corals Aulina rotiformis and Lithostrotion decipiens along a gradient of contemporaneous (Serpukhovian) open marine carbonate to near-shore siliciclastic facies in South China. Along this gradient, their sizes decrease from carbonate, through intermediate carbonate-siliciclastic, to siliciclastic facies. This is consistent with increasing abundance of terrestrial materials of high silicon, aluminium and phosphorus values. On a larger million-year-long interval (MFZ14-16) and for several palaeocontinents, size data of Lithostrotion decipiens and Siphonodendron pauciradiale show a distinct decline in late Visean, when enhanced terrestrial weathering occurred commonly with palaeosols developed during regression. This suggests that terrestrial sediment and nutrient input may have mainly controlled phenotypic plasticity in Mississippian reef corals, with a decrease in size as a component of resilience across the LPIA onset.

Mass extinctions have fundamentally altered the structure of the biosphere throughout Earth's history. The ecological severity of mass extinctions is well studied in marine ecosystems by categorizing marine taxa into functional groups based on ‘ecospace’ approaches, but the ecological response of terrestrial ecosystems to mass extinctions is less well understood due to the lack of a comparable methodology. Here, we present a new terrestrial ecospace framework that categorizes fauna into functional groups as defined by tiering, motility and feeding traits. We applied the new terrestrial and traditional marine ecospace analyses to data from the Paleobiology Database across the end-Triassic mass extinction—a time of catastrophic global warming—to compare changes between the marine and terrestrial biospheres. We found that terrestrial functional groups experienced higher extinction severity, that taxonomic and functional richness are more tightly coupled in the terrestrial, and that the terrestrial realm continued to experience high ecological dissimilarity in the wake of the extinction. Although signals of extinction severity and ecological turnover are sensitive to the quality of the terrestrial fossil record, our findings suggest greater ecological pressure from the end-Triassic mass extinction on terrestrial ecosystems than marine ecosystems, contributing to more prolonged terrestrial ecological flux.