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Abstract Relatively few clades have colonized the deep sea. Here, we analyse evolutionary pathways into this harsh environment, as a continuum defined by two potential endmembers—a ‘piecemeal model’, with exclusively deep-sea species (deep-sea endemics herein) derived from multiple, independent entries and an ‘in situ diversification model’ with one entry followed by species proliferation. We focus first on two ancient, distantly related subclades in Class Bivalvia, Mytilidae and Lucinidae, each with hundreds of species occurring globally from the intertidal to abyssal plains. Placing bathymetric ranges into newly inferred molecular phylogenies, we find that the deep-sea endemics within Lucinidae derive in piecemeal fashion, estimating up to 16 phylogenetically isolated entries and perhaps one modest in situ diversification. Mytilidae entered the deep sea just four times, with most endemics stemming from the prolific in situ diversification of Bathymodiolinae. We suggest that the contrasting phylogenetic patterns of entry and proliferation in these clades may be determined by differences in ancestral adult feeding modes. Across Bivalvia, we find that under half of extant families have deep-sea endemics, with the piecemeal model occurring more frequently. Taken together with other clades, we suggest that evolutionary pathways to deep-sea endemicity are more often shaped by multiple, independent events than by in situ diversification. 

Abstract Mass extinctions are natural experiments on the short- and long-term consequences of pushing biotas past breaking points, often with lasting effects on the structure and function of biodiversity. General properties of mass extinctions—exceptionally severe, taxonomically broad, global losses of taxa—are starting to come into focus through comparisons among dimensions of biodiversity, including morphological, functional, and phylogenetic diversity. Notably, functional diversity tends to persist despite severe losses of taxonomic diversity, whereas taxic and morphological losses may or may not be coupled. One of the biggest challenges in synthesizing and extracting general consequences of these events has been that they are often driven by multiple, interacting pressures, and the taxa and their traits vary among events, making it difficult to link single stressors to specific traits. Ongoing improvements in the taxonomic and stratigraphic resolution of these events for multiple clades will sharpen tests for selectivity and help to isolate hitchhiking effects, whereby organismal traits are carried by differential survival or extinction of taxa owing to other organismal or higher-level attributes, such as geographic-range size. Direct comparative analyses across multiple extinction events will also clarify the impacts of particular drivers on taxa, functional traits, and morphologies. It is not just the extinction filter that deserves attention, as the longer-term impact of extinctions derives in part from their ensuing rebounds. More work is needed to uncover the biotic and abiotic circumstances that spur some clades into re-diversification while relegating others to marginal shares of biodiversity. Combined insights from mass extinction filters and their rebounds bring a macroevolutionary view to approaching the biodiversity crisis in the Anthropocene, helping to pinpoint the clades, functional groups, and morphologies most vulnerable to extinction and failed rebounds. 

The end-Cretaceous (K-Pg) mass extinction shows how large-scale taxonomic loss affects functional diversity over short and long timeframes. In a macroevolutionary model system, we find that, despite losing ~60% of genera and ~20% of family-level diversity, marine bivalves lost only ~5% of their functional diversity, inconsistent with random extinction. Even with evolutionary opportunities presented by a disrupted ecosystem, low-diversity groups prior to the extinction or those originating in the Cenozoic rarely reach higher ranks today, implying long-term diversity ceilings to certain ecological roles. Clades that survived the extinction tend to dominate functions today, 66 million years post-extinction, but both relative richness and phylogenetic structure of those functional groups have been significantly shuffled. Thus, neither the composition of the pre-extinction biota nor the set of taxa that survived the extinction fully accounts for the functional and phylogenetic structure of today’s biota. The extinction disrupted Mesozoic biodiversity but did not fully determine the present-day configuration. 

Abstract Siphons in bivalves have been postulated as a key adaptive trait, enabling modes of life inaccessible to asiphonate lineages, that afford better protection from predation and dislodgement, thereby enhancing their taxonomic diversification. To test the impact of siphons on diversity, we compared two bivalve clades with similar shell forms and life positions that differ in the presence/absence of this supposed key trait: the asiphonate Archiheterodonta (origin ~ 420 Myr ago) and the siphonate Veneridae (origin ~ 170 Myr ago). We measured three characters relevant to burrowing (shell length, cross-sectional area, and proportional shell volume) in these two groups, finding that siphonate venerids occupy more modes of life than archiheterodonts because they can live at a greater range of distances from the sediment–water interface, with the thinnest shells occurring in the deepest-burrowing groups. Asiphonate taxa have thicker shells, perhaps as a compensatory adaptation in response to the potential for exposure and attack because they are limited to shallower depths of burial. The lack of siphons may have impeded morphologic and taxonomic diversification in archiheterodonts. In contrast, siphons are consistent with a key adaptive trait in the Veneridae, evidently enabling taxonomic diversification into a greater range of morphologies. 

Rigorous analysis of diversity-dependence—the hypothesis that the rate of proliferation of new species is inversely related to standing diversity—requires consideration of the ecology of the organisms in question. Differences between infaunal marine bivalves (living entirely within the sediment) and epifaunal forms (living partially or completely above the sediment–water interface) predict that these major ecological groups should have different diversity dynamics: epifaunal species may compete more intensely for space and be more susceptible to predation and physical disturbance. By comparing detrended standing diversity with rates of diversification, origination, and extinction in this exceptional fossil record, we find that epifaunal bivalves experienced significant, negative diversity-dependence in origination and net diversification, whereas infaunal forms show little appreciable relationship between diversity and evolutionary rates. This macroevolutionary contrast is robust to the time span over which dynamics are analysed, whether mass-extinction rebounds are included in the analysis, the treatment of stratigraphic ranges that are not maximally resolved, and the details of detrending. We also find that diversity-dependence persists over hundreds of millions of years, even though diversity itself rises nearly exponentially, belying the notion that diversity-dependence must imply equilibrial diversity dynamics. 

Abstract Marine bivalves are important components of ecosystems and exploited by humans for food across the world, but the intrinsic vulnerability of exploited bivalve species to global changes is poorly known. Here, we expand the list of shallow-marine bivalves known to be exploited worldwide, with 720 exploited bivalve species added beyond the 81 in the United Nations FAO Production Database, and investigate their diversity, distribution and extinction vulnerability using a metric based on ecological traits and evolutionary history. The added species shift the richness hotspot of exploited species from the northeast Atlantic to the west Pacific, with 55% of bivalve families being exploited, concentrated mostly in two major clades but all major body plans. We find that exploited species tend to be larger in size, occur in shallower waters, and have larger geographic and thermal ranges—the last two traits are known to confer extinction-resistance in marine bivalves. However, exploited bivalve species in certain regions such as the tropical east Atlantic and the temperate northeast and southeast Pacific, are among those with high intrinsic vulnerability and are a large fraction of regional faunal diversity. Our results pinpoint regional faunas and specific taxa of likely concern for management and conservation. 

Abstract Many of the most dramatic patterns in biological diversity are created by “Perfect Storms” —rare combinations of mutually reinforcing factors that push origination, extinction, or diversity accommodation to extremes. These patterns include the strongest diversification events (e.g. the Cambrian Explosion of animal body plans), the proliferation of hyperdiverse clades (e.g. insects, angiosperms), the richest biodiversity hotspots (e.g. the New World Tropical Montane regions and the ocean's greatest diversity pump, the tropical West Pacific), and the most severe extinction events (e.g. the Big Five mass extinctions of the Phanerozoic). Human impacts on the modern biota are also a Perfect Storm, and both mitigation and restoration strategies should be framed accordingly, drawing on biodiversity's responses to multi-driver processes in the geologic past. This approach necessarily weighs contributing factors, identifying their often non-linear and time-dependent interactions, instead of searching for unitary causes. 

The largest source of empirical data on the history of life largely derives from the marine invertebrates. Their rich fossil record is an important testing ground for macroecological and macroevolutionary theory, but much of this historical biodiversity remains locked away in consolidated sediments. Manually preparing invertebrate fossils out of their matrix can require weeks to months of careful excavation and cannot guarantee the recovery of important features on specimens. Micro-CT is greatly improving our access to the morphologies of these fossils, but it remains difficult to digitally separate specimens from sediments of similar compositions, e.g., calcareous shells in a carbonate rich matrix. Here we provide a workflow for using deep learning—a subset of machine learning based on artificial neural networks—to augment the segmentation of these difficult fossils. We also provide a guide for bulk scanning fossil and Recent shells, with sizes ranging from 1 mm to 20 cm, enabling the rapid acquisition of large-scale 3D datasets for macroevolutionary and macroecological analyses (300–500 shells in 8 hours of scanning). We then illustrate how these approaches have been used to access new dimensions of morphology, allowing rigorous statistical testing of spatial and temporal patterns in morphological evolution, which open novel research directions in the history of life. 

Abstract Evolvability is best addressed from a multi-level, macroevolutionary perspective through a comparative approach that tests for among-clade differences in phenotypic diversification in response to an opportunity, such as encountered after a mass extinction, entering a new adaptive zone, or entering a new geographic area. Analyzing the dynamics of clades under similar environmental conditions can (partially) factor out shared external drivers to recognize intrinsic differences in evolvability, aiming for a macroevolutionary analog of a common-garden experiment. Analyses will be most powerful when integrating neontological and paleontological data: determining differences among extant populations that can be hypothesized to generate large-scale, long-term contrasts in evolvability among clades; or observing large-scale differences among clade histories that can by hypothesized to reflect contrasts in genetics and development observed directly in extant populations. However, many comparative analyses can be informative on their own, as explored in this overview. Differences in clade-level evolvability can be visualized in diversity-disparity plots, which can quantify positive and negative departures of phenotypic productivity from stochastic expectations scaled to taxonomic diversification. Factors that evidently can promote evolvability include modularity—when selection aligns with modular structure or with morphological integration patterns; pronounced ontogenetic changes in morphology, as in allometry or multiphase life cycles; genome size; and a variety of evolutionary novelties, which can also be evaluated using macroevolutionary lags between the acquisition of a trait and phenotypic diversification, and dead-clade-walking patterns that may signal a loss of evolvability when extrinsic factors can be excluded. High speciation rates may indirectly foster phenotypic evolvability, and vice versa. Mechanisms are controversial, but clade evolvability may be higher in the Cambrian, and possibly early in the history of clades at other times; in the tropics; and, for marine organisms, in shallow-water disturbed habitats.