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Breaking bilateral symmetry in bivalves. A Cretaceous rudist Radiolites angeiodes showing disparate left (now upper) and right (now lower) valve (after Skelton, 1979, used by permission). B Eocene Caestocorbula praeviator, living within sediment with larger and more heavily ornamented left valve downward (from Beu & Raine, 2009). C extant scallop Cyclopecten hoskynsi, left valve above, right valve below (from Dijkstra et al., 2009). D Coiled Jurassic oyster Gryphaea arcuata, living with left valve downward on soft seafloor (from Seilacher, 1984). E Helically coiled Cretaceous oyster Ilymatogyra arietina, living with left valve downward on soft seafloor (from Roemer, 1862)
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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 cl...
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Citations
... Gene family expansion and convergent evolution in arctic-alpine lineages Ancient WGDs are often associated with trait innovations and climatic shifts (te Beest et al., 2012;Moghe and Shiu, 2014;Brockington et al., 2015;Smith et al., 2018b;Baniaga et al., 2020;Wu et al., 2020). One of the mechanisms by which ancient WGDs contribute to trait innovations is gene family expansion Wang et al., 2019;Jablonski, 2022). Smith et al. (2018b) tested climatic shifts in the Caryophyllales with increased sampling using Sangersequencing-based markers from GenBank and inferred a shift into colder climates (lower mean annual temperature) at the node corresponding to WGD1 (Figure 3). ...
... Punctuated evolution resembles slow-fast dynamics, where certain components within a system change at different rates and the system's dynamics are driven by the dependence on frequency [40], heterogeneous networks [33], and the spatial structure [41]. Eldredge and Gould [1] proposed that ephemeral local adaptations contribute to long-term trends only when 'fixed' by speciation as a threshold-like event. ...
For five decades, paleontologists, paleobiologists, and ecologists have investigated patterns of punctuated equilibria in biology. Here, we step outside those fields and summarize recent advances in the theory of and evidence for punctuated equilibria, gathered from contemporary observations in geology, molecular biology, genetics, anthropology, and sociotechnology. Taken in the aggregate, these observations lead to a more general theory that we refer to as punctuated evolution. The quality of recent datasets is beginning to illustrate the mechanics of punctuated evolution in a way that can be modeled across a vast range of phenomena, from mass extinctions hundreds of millions of years ago to the possible future ahead in the Anthropocene. We expect the study of punctuated evolution to be applicable beyond biological scenarios.
... There is also extensive debate about whether evolvability should be conceptualized as a cause or a product of evolution (Payne & Wagner, 2019). Some authors advocate viewing the ability for an organism to evolve as a selectable trait analogous to adaptation, while others argue it is more an outcome of various genetic and developmental properties (Jablonski, 2022). ...
This article reviews the edited volume “Evolvability: A Unifying Concept in Evolutionary Biology?” through biological and philosophical lenses. The book provides diverse angles on evolvability, which is affected by various hierarchical levels, timescales, and types of variation, thus moving beyond a purely genomics perspective. Evolvability is important to biosemiotics because understanding the dynamics of topological genotype spaces could help one better comprehend the phenotypic spaces of meaning, as developmental codes and interrelations can influence the emergence of biological novelty over time. This book is successful in provoking thought on evolvability's role as both a product and a driver of evolutionary innovations and explores the philosophical implications of how evolution has the capacity to generate novel forms of meaning.
... Since then, different mechanisms have been proposed to explain the origin and evolution of deep-sea biodiversity. For example, the onshore-to-offshore hypothesis suggests that disturbances including waves and storms at shelf depths (0-250 m) creates habitats that are challenging for species to inhabit, which results in diversification of species to occupy these habitats [9,10]. Compared with the shelf, the slope (250-3000 m) is exposed to fewer environmental perturbations, with littleto-no effect from waves and storms [11]. ...
... octocorals) to invade the shelf [29,62]. Most black corals in Clade A 1 have since diversified in the more dynamic shelf environment [9,10] potentially driving a rapid radiation in the group. We do not explicitly test rates of diversification here due to the limited sampling of species; however, there is evidence of elevated diversity on the slope: over 20 extant black coral genera occur on the slope, including the genus Antipathes that currently contains 75 of 300 currently accepted species in the order [51]. ...
Deep-sea lineages are generally thought to arise from shallow-water ancestors, but this hypothesis is based on a relatively small number of taxonomic groups. Anthozoans, which include corals and sea anemones, are significant contributors to the faunal diversity of the deep sea, but the timing and mechanisms of their invasion into this biome remain elusive. Here, we reconstruct a fully resolved, time-calibrated phylogeny of 83 species in the order Antipatharia (black coral) to investigate their bathymetric evolutionary history. Our reconstruction indicates that extant black coral lineages first diversified in continental slope depths (∼250–3000 m) during the early Silurian (∼437 millions of years ago (Ma)) and subsequently radiated into, and diversified within, both continental shelf (less than 250 m) and abyssal (greater than 3000 m) habitats. Ancestral state reconstruction analysis suggests that the appearance of morphological features that enhanced the ability of black corals to acquire nutrients coincided with their invasion of novel depths. Our findings have important conservation implications for anthozoan lineages, as the loss of ‘source’ slope lineages could threaten millions of years of evolutionary history and confound future invasion events, thereby warranting protection.
... The latter determines the evolvability. This biological emerging property is the propensity to evolve in response to opportunity 4 and has become a major topic in evolutionary studies 4,5 . Studying evolutionary patterns associated with similar ecological conditions (i.e., opportunity), we are able to factor out extrinsic drivers 4 . ...
... The latter determines the evolvability. This biological emerging property is the propensity to evolve in response to opportunity 4 and has become a major topic in evolutionary studies 4,5 . Studying evolutionary patterns associated with similar ecological conditions (i.e., opportunity), we are able to factor out extrinsic drivers 4 . ...
... This biological emerging property is the propensity to evolve in response to opportunity 4 and has become a major topic in evolutionary studies 4,5 . Studying evolutionary patterns associated with similar ecological conditions (i.e., opportunity), we are able to factor out extrinsic drivers 4 . The widespread ecomorphological convergence, i.e., independently evolved morphological similarity associated with convergent ecologies, reproduces this experimental design and was compared to a common-garden experiment by Jablonski 4 . ...
Eco-morphological convergence, i.e., similar phenotypes evolved in ecologically convergent taxa, naturally reproduces a common-garden experiment since it allows researchers to keep ecological factors constant, studying intrinsic evolutionary drivers. The latter may result in differential evolvability that, among individual anatomical parts, causes mosaic evolution. Reconstructing the evolutionary morphology of the humerus and femur of slow arboreal mammals, we addressed mosaicism at different bone anatomical spatial scales. We compared convergence strength, using it as indicator of evolvability, between bone external shape and inner structure, with the former expected to be less evolvable and less involved in convergent evolution, due to anatomical constraints. We identify several convergent inner structural traits, while external shape only loosely follows this trend, and we find confirmation for our assumption in measures of convergence magnitude. We suggest that future macroevolutionary reconstructions based on bone morphology should include structural traits to better detect ecological effects on vertebrate diversification.
... The onset of their Ordovician diversification coincided with an increase in nutrients and ocean ventilation [45] and the transformation of the seafloor from surfaces stabilized by microbial mats to softer, bioturbated sediments [15,29], with bivalves expanding into both on-and offshore habitats with soft-and hard-ground substrata [43]. Thus, bivalves may be another instance of a clade whose diversification required a chain of derived characters, rather than a single "key innovation" [46,47]. ...
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.
... The term evolvability bears various meanings in the current evolutionary biology (e.g., Brown 2014; Crother and Murray 2019;Feiner et al. 2021;Jablonski 2022;Love et al. 2022;Riederer et al. 2022). In the present paper, the use of this term is restricted to certain mathematical aspects of the G matrix and response vectors-the Hansen-Houle evolvability-unless stated otherwise. ...
Theory predicts that the additive genetic covariance (\documentclass[12pt]{minimal} \usepackage{amsmath} \usepackage{wasysym} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{amsbsy} \usepackage{mathrsfs} \usepackage{upgreek} \setlength{\oddsidemargin}{-69pt} \begin{document}$$\textbf{G}$$\end{document}G) matrix determines a population’s short-term (in)ability to respond to directional selection—evolvability in the Hansen–Houle sense—which is typically quantified and compared via certain scalar indices called evolvability measures. Often, interest is in obtaining the averages of these measures across all possible selection gradients, but explicit formulae for most of these average measures have not been known. Previous authors relied either on approximations by the delta method, whose accuracy is generally unknown, or Monte Carlo evaluations (including the random skewers analysis), which necessarily involve random fluctuations. This study presents new, exact expressions for the average conditional evolvability, average autonomy, average respondability, average flexibility, average response difference, and average response correlation, utilizing their mathematical structures as ratios of quadratic forms. The new expressions are infinite series involving top-order zonal and invariant polynomials of matrix arguments, and can be numerically evaluated as their partial sums with, for some measures, known error bounds. Whenever these partial sums numerically converge within reasonable computational time and memory, they will replace the previous approximate methods. In addition, new expressions are derived for the average measures under a general normal distribution for the selection gradient, extending the applicability of these measures into a substantially broader class of selection regimes.
Supplementary Information
The online version contains supplementary material available at 10.1007/s00285-023-01930-8.
... We use evolvability, as defined by Houle (1992), to determine whether aggregate traits in colonies can evolve. Evolvability is a measure of trait variance that is scaled by the trait mean, and can be interpreted as the maximum possible rate of evolutionary change that can occur across a single generation under direct selection (Houle, 1992;Hansen & Houle, 2008;Love et al., 2021;Jablonski, 2022). ...
The study of trait evolution in modular animals is more complicated than that in solitary animals, because a single genotype of a modular colony can express an enormous range of phenotypic variation. Furthermore, traits can occur either at the module level or at the colony level. However, it is unclear how the traits at the colony level evolve. We test whether colony-level aggregate traits, defined as the summary statistics of a phenotypic distribution, can evolve. To quantify this evolutionary potential, we use parent-offspring pairs in two sister species of the bryozoan Stylopoma , grown and bred in a common garden breeding experiment. We find that the medians of phenotypic distributions are evolvable between generations of colonies. We also find that the structure of this evolutionary potential differs between these two species. Ancestral species align more closely with the direction of species divergence than the descendent species. This result indicates that aggregate trait evolvability can itself evolve.
... After all, there have been studies on the effect of temperature on rates of spontaneous mutation that could drive speciation (Husby et al, 2011;Waldvogel and Pfenninger, 2021;Z Lu et al, 2021;Gillman and Wright, 2014;Chu et al, 2018). What factors drive mutation and the divergence of species have been a recurring experimental and theoretical question (Jablonski, 2022;Livnat, 2017;Smith et al, 2014). In this paper, the effect of changing temperatures as it perturbs the balance in nature is investigated from an analytical framework. ...
The impact of climate change on biodiversity needs to be understood from a multidisciplinary approach. Using an analytical framework, we investigate the species response to rising temperatures. Common traits and characteristics among species that allow classification at different taxonomic levels imply an underlying symmetry that gives rise to invariances behind the biodiversity observed in nature. Changing temperatures that go beyond a critical limit break this underlying symmetry which could lead to enhanced speciation.
Objectives: Modular architecture of traits in complex organisms can be important for morphological evolution at micro- and sometimes macroevolutionary scales as it may influence the tempo and direction of changes to groups of traits that are essential for particular functions, including food acquisition and processing. We tested several distinct hypotheses about craniofacial modularity in the hominine skull in relation to feeding biomechanics.
Materials and Methods: First, we formulated hypothesized functional modules for craniofacial traits reflecting specific demands of feeding biomechanics (e.g., masseter leverage/gape or tooth crown mechanics) in Homo sapiens, Pan troglodytes, and Gorilla gorilla. Then, the pattern and strength of modular signal was quantified by the covariance ratio coefficient and compared across groups using covariance ratio effect size. Hierarchical clustering analysis was then conducted to examine whether a priori-defined functional modules correspond to empirically recovered clusters.
Results: There was statistical support for most a priori-defined functional modules in the cranium and half of the functional modules in the mandible. Modularity signal was similar in the cranium and mandible, and across the three taxa. Despite a similar strength of modularity, the empirically recovered clusters do not map perfectly onto our priori functional modules, indicating that further work is needed to refine our hypothesized functional modules.
Conclusion: The results suggest that modular structure of traits in association with feeding biomechanics were mostly shared with humans and the two African apes. Thus, conserved patterns of functional modularity may have facilitated evolutionary changes to the skull during human evolution.
