Ecological structure of diversity-dependent diversification in Phanerozoic marine bivalves.

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.

Riverine flow rate drives widespread convergence in the shell morphology of imperiled freshwater mussels.

Frequent and strong morphological convergence suggests that determinism tends to supersede historical contingencies in evolutionary radiations. For many lineages living within the water column of rivers and streams, hydrodynamic forces drive widespread morphological convergence. Living below the sediment-water interface may release organisms from these hydrodynamic pressures, permitting a broad array of morphologies, and thus less convergence. However, we show here that the semi-infaunal freshwater mussels have environmentally determined convergence in shell morphology. Using 3D morphometric data from 715 individuals among 164 Nearctic species, we find that species occurring in rivers with high flow rates have evolved traits that resist dislodgement from their burrowed position in the streambed: thicker shells for their body size, with the thickest sector of the shell being the most deeply buried. Species occurring in low flow environments have evolved thinner and more uniformly thickened shells, corresponding to an alternative adaptation to dislodgement: increased burrowing efficiency. Within species, individuals also show increased shell thickness for their body size at higher flow rates, suggesting that ecophenotypy may, in part, be an important mechanism for establishing populations in new environments and thus evolutionary divergence in this highly imperiledinvertebrate group.

Investigations into 3D-printed nautiloid-inspired pressure housings.

The shell of the chambered nautilus is one of the few examples in nature of a biologically derived one-atmosphere pressure housing, which the animal uses to maintain neutral buoyancy via a series of sealed chambers. Extant species such asNautilus pompiliuslive at depths from 200 to 800 m, and similar depth ranges have been hypothesized for their hyper diverse but extinct relatives, the ammonoids. Given the evolutionary success of these molluscan clades, their complex shell morphologies may reveal pressure-tolerant geometries comparable to the 'ideal' ones currently used in deep-sea marine robotics: simple spheres and cylinders, which have minimized surface area to volume ratio and easier manufacturability. We modeled and empirically tested 3D-printed bioinspired pressure housings for deep-sea applications using high resolution stereolithography 3D printing. These designs were modeled on the shells ofN. pompiliusand were compared to conventional 3D-printed spheres with similar wall thicknesses and implodable volumes. Two nautilus-inspired models with internal supports designed after their septal walls (one concave, one convex) had a higher-pressure tolerance compared to hollow models, but none outperformed spherical models with the same outer-wall thickness. Although spheres outperform the nautilus-inspired housings, the methods developed here show that pressure housings with complex geometries can be printed by additive manufacturing and empirically tested. From a biological perspective, this method can be a new tool for empirically testing viable depth tolerances for extinct coiled cephalopod morphologies.

Diversity, distribution and intrinsic extinction vulnerability of exploited marine bivalves.

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.

Trends in botanical exploration in Nigeria forecast over 1000 yet undescribed vascular plant species.

BACKGROUND AND AIMS: Taxonomists are primary actors of biodiversity assessment. At the same time, there is awareness by the taxonomic community at large that the field is going through a crisis, sometimes referred to as the "taxonomic impediment". Coupled with the ongoing biodiversity crisis, or 6 th mass extinction, this biodiversity impedance puts at risk the target set in the Convention on Biological Diversity's Global Biodiversity Framework vision 2050, which calls for urgent action to "…put biodiversity on a path to recovery by 2030 for the benefit of planet and people". This risk is particularly pronounced in tropical African countries where taxonomic studies are done on an ad hoc basis. In this study, our aim is to investigate the historical trends in botanical exploration of vascular plants in Nigeria and forecast the near-term (50 year) description of presently unknown species, which we use to discuss scenarios of taxonomic effort that may be necessary for a comprehensive biodiversity assessment in the country. METHODS: The study is based on a dataset from the World Checklist of Vascular Plants (WCVP), containing all vascular plant species reported to occur in Nigeria. We fit nested Bayesian time series regressions to estimate the long-term trend in the rate of description of vascular plant species in Nigeria. From these models, we use an ensemble forecast to estimate the number of species descriptions by the year 2070, and then evaluate the description rates per taxonomist required to meet this estimate under different totals of active taxonomists. KEY RESULTS: We find a striking difference in species description between Nigerian botanists and their foreign counterparts, with the former contributing relatively small numbers. Additionally, only a fraction of the authors involved in describing Nigeria's vascular plants are of indigenous origin. Our study reveals that the number of new species described annually exhibits a long-term increasing trend, with an average of 19.5 species described per year. However, after taking into account year-to-year variability and the number of taxonomists active in a given year, the long-term trend in species descriptions credibly declines over time. While the number of authors involved in describing species has generally increased over time, it remained stable since the 1950s. Predictions for the number of new species descriptions by 2070 vary by model, with an ensemble prediction estimating 1140 species descriptions, but ranging from 1004 to 2239 between individual models. CONCLUSIONS: The study estimates that current levels of taxonomic activity should lead to a 20% increase in known species of vascular plants in Nigeria over the next 50 years, which is still likely an underestimate of the true, unknown species richness. Urgent action is needed to address the taxonomic impediment so that local taxonomic studies in tropical African countries can achieve the CBD's Global Biodiversity Framework vision 2050. Here, we outline some key pathways to achieving this goal.

Cambrian origin but no early burst in functional disparity for Class Bivalvia.

Both the Cambrian explosion, more than half a billion years ago, and its Ordovician aftermath some 35 Myr later, are often framed as episodes of widespread ecological opportunity, but not all clades originating during this interval showed prolific rises in morphological or functional disparity. In a direct analysis of functional disparity, instead of the more commonly used proxy of morphological disparity, we find that ecological functions of Class Bivalvia arose concordantly with and even lagged behind taxonomic diversification, rather than the early-burst pattern expected for clades originating in supposedly open ecological landscapes. Unlike several other clades originating in the Cambrian explosion, the bivalves' belated acquisition of key anatomical novelties imposed a macroevolutionary lag, and even when those novelties evolved in the Early Ordovician, functional disparity never surpassed taxonomic diversity. Beyond this early period of animal evolution, the founding and subsequent diversification of new major clades and their functions might be expected to follow the pattern of the early bivalves-one where interactions between highly dynamic environmental and biotic landscapes and evolutionary contingencies need not promote prolific functional innovation.

Biologically driven isotopic fractionations in bivalves: from palaeoenvironmental problem to palaeophysiological proxy.

Traditional bulk stable isotope (δ18 O and δ13 C) and clumped isotope (Δ47 ) records from bivalve shells provide invaluable histories of Earth's local and global climate change. However, biologically driven isotopic fractionations (BioDIFs) can overprint primary environmental signals in the shell. Here, we explore how conventional measurements of δ18 O, δ13 C, and Δ47 in bivalve shells can be re-interpreted to investigate these physiological processes deliberately. Using intrashell Δ47 and δ18 O alignment as a proxy for equilibrium state, we separately examine fractionations and/or disequilibrium occurring in the two major stages of the biomineralisation process: the secretion of the extrapallial fluid (EPF) and the precipitation of shell material from the EPF. We measured δ18 O, δ13 C, and Δ47 in fossil shells representing five genera (Lahillia, Dozyia, Eselaevitrigonia, Nordenskjoldia, and Cucullaea) from the Maastrichtian age [66-69 million years ago (Ma)] López de Bertodano Formation on Seymour Island, Antarctica. Material was sampled from both the outer and inner shell layers (OSL and ISL, respectively), which precipitate from separate EPF reservoirs. We find consistent δ18 O values across the five taxa, indicating that the composition of the OSL can be a reliable palaeoclimate proxy. However, relative to the OSL baseline, ISLs of all taxa show BioDIFs in one or more isotopic parameters. We discuss/hypothesise potential origins of these BioDIFs by synthesising isotope systematics with the physiological processes underlying shell biomineralisation. We propose a generalised analytical and interpretive framework that maximises the amount of palaeoenvironmental and palaeobiological information that can be derived from the isotopic composition of fossil shell material, even in the presence of previously confounding 'vital effects'. Applying this framework in deep time can expand the utility of δ18 O, δ13 C, and Δ47 measurements from proxies of past environments to proxies for certain biomineralisation strategies across space, time, and phylogeny among Bivalvia and other calcifying organisms.

Convergence and contingency in the evolution of a specialized mode of life: multiple origins and high disparity of rock-boring bivalves.

Evolutionary adaptation to novel, specialized modes of life is often associated with a close mapping of form to the new function, resulting in narrow morphological disparity. For bivalve molluscs, endolithy (rock-boring) has biomechanical requirements thought to diverge strongly from those of ancestral functions. However, endolithy in bivalves has originated at least eight times. Three-dimensional morphometric data representing 75 species from approximately 94% of extant endolithic genera and families, along with 310 non-endolithic species in those families, show that endolithy is evolutionarily accessible from many different morphological starting points. Although some endoliths appear to converge on certain shell morphologies, the range of endolith shell form is as broad as that belonging to any other bivalve substrate use. Nevertheless, endolithy is a taxon-poor function in Bivalvia today. This limited richness does not derive from origination within source clades having significantly low origination or high extinction rates, and today's endoliths are not confined to low-diversity biogeographic regions. Instead, endolithy may be limited by habitat availability. Both determinism (as reflected by convergence among distantly related taxa) and contingency (as reflected by the endoliths that remain close to the disparate morphologies of their source clades) underlie the occupation of endolith morphospace.

Specimen alignment with limited point-based homology: 3D morphometrics of disparate bivalve shells (Mollusca: Bivalvia).

Background: Comparative morphology fundamentally relies on the orientation and alignment of specimens. In the era of geometric morphometrics, point-based homologies are commonly deployed to register specimens and their landmarks in a shared coordinate system. However, the number of point-based homologies commonly diminishes with increasing phylogenetic breadth. These situations invite alternative, often conflicting, approaches to alignment. The bivalve shell (Mollusca: Bivalvia) exemplifies a homologous structure with few universally homologous points-only one can be identified across the Class, the shell 'beak'. Here, we develop an axis-based framework, grounded in the homology of shell features, to orient shells for landmark-based, comparative morphology. Methods: Using 3D scans of species that span the disparity of shell morphology across the Class, multiple modes of scaling, translation, and rotation were applied to test for differences in shell shape. Point-based homologies were used to define body axes, which were then standardized to facilitate specimen alignment via rotation. Resulting alignments were compared using pairwise distances between specimen shapes as defined by surface semilandmarks. Results: Analysis of 45 possible alignment schemes finds general conformity among the shape differences of 'typical' equilateral shells, but the shape differences among atypical shells can change considerably, particularly those with distinctive modes of growth. Each alignment corresponds to a hypothesis about the ecological, developmental, or evolutionary basis of morphological differences, but we suggest orientation via the hinge line for many analyses of shell shape across the Class, a formalization of the most common approach to morphometrics of shell form. This axis-based approach to aligning specimens facilitates the comparison of approximately continuous differences in shape among phylogenetically broad and morphologically disparate samples, not only within bivalves but across many other clades.

Evolutionary modularity, integration and disparity in an accretionary skeleton: analysis of venerid Bivalvia.

Modular evolution, the relatively independent evolution of body parts, may promote high morphological disparity in a clade. Conversely, integrated evolution via stronger covariation of parts may limit disparity. However, integration can also promote high disparity by channelling morphological evolution along lines of least resistance-a process that may be particularly important in the accumulation of disparity in the many invertebrate systems having accretionary growth. We use a time-calibrated phylogenetic hypothesis and high-density, three-dimensional semilandmarking to analyse the relationship between modularity, integration and disparity in the most diverse extant bivalve family: the Veneridae. In general, venerids have a simple, two-module parcellation of their body that is divided into features of the calcium carbonate shell and features of the internal soft anatomy. This division falls more along developmental than functional lines when placed in the context of bivalve anatomy and biomechanics. The venerid body is tightly integrated in absolute terms, but disparity appears to increase with modularity strength among subclades and ecologies. Thus, shifts towards more mosaic evolution beget higher morphological variance in this speciose family.