Adaptive radiation without independent stages of trait evolution in a group of Caribbean anoles.

Adaptive radiation involves diversification along multiple trait axes, producing phenotypically diverse, species-rich lineages. Theory generally predicts that multi-trait evolution occurs via a 'stages' model, with some traits saturating early in a lineage's history, and others diversifying later. Despite its multidimensional nature, however, we know surprisingly little about how different suites of traits evolve during adaptive radiation. Here, we investigated the rate, pattern, and timing of morphological and physiological evolution in the anole lizard adaptive radiation from the Caribbean island of Hispaniola. Rates and patterns of morphological and physiological diversity are largely unaligned, corresponding to independent selective pressures associated with structural and thermal niches. Cold tolerance evolution reflects parapatric divergence across elevation, rather than niche partitioning within communities. Heat tolerance evolution and the preferred temperature evolve more slowly than cold tolerance, reflecting behavioral buffering, particularly in edge-habitat species (a pattern associated with the Bogert effect). In contrast to the nearby island of Puerto Rico, closely related anoles on Hispaniola do not sympatrically partition thermal niche space. Instead, allopatric and parapatric separation across biogeographic and environmental boundaries serves to keep morphologically similar close relatives apart. The phenotypic diversity of this island's adaptive radiation accumulated largely as a by-product of time, with surprisingly few exceptional pulses of trait evolution. A better understanding of the processes that guide multidimensional trait evolution (and nuance therein) will prove key in determining whether the stages model should be considered a common theme of adaptive radiation.

Phylogenetic inference of where species spread or split across barriers.

SignificanceGeography molds how species evolve in space. Strong geographical barriers to movement, for instance, both inhibit dispersal between regions and allow isolated populations to diverge as new species. Weak barriers, by contrast, permit species range expansion and persistence. These factors present a conundrum: How strong must a barrier be before between-region speciation outpaces dispersal? We designed a phylogenetic model of dispersal, extinction, and speciation that allows regional features to influence rates of biogeographic change and applied it to the neotropical radiation of Anolis lizards. Separation by water induces a threefold steeper barrier to movement than equivalent distances over land. Our model will help biologists detect relationships between evolutionary processes and the spatial contexts in which they operate.

Functional Trade-Offs Asymmetrically Promote Phenotypic Evolution.

Trade-offs are thought to bias evolution and are core features of many anatomical systems. Therefore, trade-offs may have far-reaching macroevolutionary consequences, including patterns of morphological, functional, and ecological diversity. Jaws, like many complex anatomical systems, are comprised of elements involved in biomechanical trade-offs. We test the impact of a core mechanical trade-off, the transmission of velocity versus force (i.e., mechanical advantage), on rates of jaw evolution in Neotropical cichlids. Across 130 species representing a wide array of feeding ecologies, we find that the velocity-force trade-off impacts the evolution of the surrounding jaw system. Specifically, rates of jaw evolution are faster at functional extremes than in more functionally intermediate or unspecialized jaws. Yet, surprisingly, the effect on jaw evolution is uneven across the extremes of the velocity-force continuum. Rates of jaw evolution are 4- to 10-fold faster in velocity-modified jaws, whereas force-modified jaws are 7- to 18-fold faster, compared to unspecialized jaws, depending on the extent of specialization. Further, we find that a more extreme mechanical trade-off resulted in faster rates of jaw evolution. The velocity-force trade-off reflects a gradient from specialization on capture-intensive (e.g., evasive or buried) to processing-intensive prey (e.g., attached or shelled), respectively. The velocity extreme of the trade-off is characterized by large magnitudes of trait change leading to functionally divergent specialists and ecological stasis. By contrast, the force extreme of the trade-off is characterized by enhanced ecological lability made possible by phenotypes more readily co-opted for different feeding ecologies. This asymmetry of macroevolutionary outcomes along each extreme is likely the result of an enhanced utility of the pharyngeal jaw system as force-modified oral jaws are adapted for prey that requires intensive processing (e.g., algae, detritus, and mollusks). The velocity-force trade-off, a fundamental feature of many anatomical systems, promotes rapid phenotypic evolution of the surrounding jaw system in a canonical continental adaptive radiation. Considering that the velocity-force trade-off is an inherent feature of all jaw systems that involve a lower element that rotates at a joint, spanning the vast majority of vertebrates, our results may be widely applicable across the tree of life. [Adaptive radiation; constraint; decoupling; jaws; macroevolution; specialization.].

Let's include the ladies: Do morphology-performance relationships vary between sexes in lizards?

An animal's morphology influences its ability to perform essential tasks, such as locomoting to obtain prey or escape predators. While morphology-performance relationships are well-studied in lizards, most conclusions have been based only on male study subjects, leaving unanswered questions about females. Sex-specific differences are important to understand because females carry the bulk of the physiological demands of reproduction. Consequently, their health and survival can determine the fate of the population as a whole. To address this knowledge gap, we sampled introduced populations of common wall lizards (Podarcis muralis) in Ohio, USA. We measured a complete suite of limb and body dimensions of both males and females, and we measured sprint speeds while following straight paths and curved paths on different substrates. Using a multivariate statistical approach, we identified that body dimensions relative to snout-to-vent length in males were much larger compared to females and that body dimensions of P. muralis have changed over time in both sexes. We found that sprint speed along curved paths increased with relative limb size in both males and females. When following straight paths, male speed similarly increased as body dimensions increased; conversely, female speed decreased as body dimensions increased. Female sprint speed was also found to have less variation than that of males and was less affected by changes in body size and hindfoot length compared to males. This study thus provides insights into how selective pressures might shape males and females differently and the functional implications of sexual dimorphism.

Parameterizing mechanistic niche models in biophysical ecology: a review of empirical approaches.

Mechanistic niche models are computational tools developed using biophysical principles to address grand challenges in ecology and evolution, such as the mechanisms that shape the fundamental niche and the adaptive significance of traits. Here, we review the empirical basis of mechanistic niche models in biophysical ecology, which are used to answer a broad array of questions in ecology, evolution and global change biology. We describe the experiments and observations that are frequently used to parameterize these models and how these empirical data are then incorporated into mechanistic niche models to predict performance, growth, survival and reproduction. We focus on the physiological, behavioral and morphological traits that are frequently measured and then integrated into these models. We also review the empirical approaches used to incorporate evolutionary processes, phenotypic plasticity and biotic interactions. We discuss the importance of validation experiments and observations in verifying underlying assumptions and complex processes. Despite the reliance of mechanistic niche models on biophysical theory, empirical data have and will continue to play an essential role in their development and implementation.

Ecological Opportunity from Innovation, not Islands, Drove the Anole Lizard Adaptive Radiation.

Islands are thought to facilitate adaptive radiation by providing release from competition and predation. Anole lizards are considered a classic example of this phenomenon: different ecological specialists ("ecomorphs") evolved in the Caribbean Greater Antilles (Cuba, Hispaniola, Jamaica, and Puerto Rico), resulting in convergent assemblages that are not observed in mainland Latin America. Yet, the role of islands in facilitating adaptive radiation is more often implied than directly tested, leaving uncertain the role of biogeography in stimulating diversification. Here, we assess the proposed "island effect" on anole diversification using Bayesian phylogenetic comparative methods that explicitly incorporate rate heterogeneity across the tree and demonstrate two cases of would be false positives. We discovered that rates of speciation and morphological evolution of island and mainland anoles are equivalent, implying that islands provide no special context for exceptionally rapid diversification. Likewise, rates of evolution were equivalent between island anoles that arose via in situ versus dispersal-based mechanisms, and we found no evidence for island-specific rates of speciation or morphological evolution. Nonetheless, the origin of Anolis is characterized by a speciation pulse that slowed over time-a classic signature of waning ecological opportunity. Our findings cast doubt on the notion that islands catalyzed the anole adaptive radiation and instead point to a key innovation, adhesive toe pads, which facilitated the exploitation of many arboreal niches sparsely utilized by other iguanian lizards. The selective pressures responsible for arboreal niche diversification differ between islands and the mainland, but the tempo of diversification driven by these discordant processes is indistinguishable. [Anolis; Caribbean; key innovation; morphological evolution; RevBayes; speciation.].

Thermal-metabolic phenotypes of the lizard Podarcis muralis differ across elevation, but converge in high-elevation hypoxia.

In response to a warming climate, many montane species are shifting upslope to track the emergence of preferred temperatures. Characterizing patterns of variation in metabolic, physiological and thermal traits along an elevational gradient, and the plastic potential of these traits, is necessary to understand current and future responses to abiotic constraints at high elevations, including limited oxygen availability. We performed a transplant experiment with the upslope-colonizing common wall lizard (Podarcis muralis) in which we measured nine aspects of thermal physiology and aerobic capacity in lizards from replicate low- (400 m above sea level, ASL) and high-elevation (1700 m ASL) populations. We first measured traits at their elevation of origin and then transplanted half of each group to extreme high elevation (2900 m ASL; above the current elevational range limit of this species), where oxygen availability is reduced by ∼25% relative to sea level. After 3 weeks of acclimation, we again measured these traits in both the transplanted and control groups. The multivariate thermal-metabolic phenotypes of lizards originating from different elevations differed clearly when measured at the elevation of origin. For example, high-elevation lizards are more heat tolerant than their low-elevation counterparts (counter-gradient variation). Yet, these phenotypes converged after exposure to reduced oxygen availability at extreme high elevation, suggesting limited plastic responses under this novel constraint. Our results suggest that high-elevation populations are well suited to their oxygen environments, but that plasticity in the thermal-metabolic phenotype does not pre-adapt these populations to colonize more hypoxic environments at higher elevations.

Interactions between thermoregulatory behavior and physiological acclimatization in a wild lizard population.

Although the importance of thermoregulation and plasticity as compensatory mechanisms for climate change has long been recognized, they have largely been studied independently. Thus, we know comparatively little about how they interact to shape physiological variation in natural populations. Here, we test the hypothesis that behavioral thermoregulation and thermal acclimatization interact to shape physiological phenotypes in a natural population of the diurnal lizard, Sceloporus torquatus. Every month for one year we examined thermoregulatory effectiveness and changes in the population mean in three physiological parameters: cold tolerance (Ctmin), heat tolerance (Ctmax), and the preferred body temperature (Tpref), to indirectly assess thermal acclimatization in population means. We discovered that S. torquatus is an active thermoregulator throughout the year, with body temperature varying little despite strong seasonal temperature shifts. Although we did not observe a strong signal of acclimatization in Ctmax, we did find that Ctmin shifts in parallel with nighttime temperatures throughout the year. This likely occurs, at least in part, because thermoregulation is substantially less effective at buffering organisms from selection on lower physiological limits than upper physiological limits. Active thermoregulation is effective at limiting exposure to extreme temperatures during the day, but is less effective at night, potentially contributing to greater plasticity in Ctmin than Ctmax. Importantly, however, Tpref tracked seasonal changes in temperature, which is one the factors contributing to highly effective thermoregulation throughout the year. Thus, behavior and physiological plasticity do not always operate independently, which could impact how organisms can respond to rising temperatures.

Thermoregulatory Behavior Simultaneously Promotes and Forestalls Evolution in a Tropical Lizard.

The role of behavior in evolution has long been discussed, with some arguing that behavior promotes evolution by exposing organisms to selection (behavioral drive) and others proposing that it inhibits evolution by shielding organisms from environmental variation (behavioral inertia). However, this discussion has generally focused on the effects of behavior along a single axis without considering that behavior simultaneously influences selection in various niche dimensions. By examining evolutionary change along two distinct niche axes-structural and thermal-we propose that behavior simultaneously drives and impedes evolution in a group of Anolis lizards from the Caribbean island of Hispaniola. Specifically, a behavioral shift in microhabitat to boulders at high altitude enables thermoregulation, thus forestalling physiological evolution in spite of colder environments. This same behavioral shift drives skull and limb evolution to boulder use. Our results emphasize the multidimensional effects of behavior in evolution. These findings reveal how, rather than being diametrically opposed, niche conservatism and niche lability can occur simultaneously. Furthermore, patterns of niche evolution may vary at different geographic scales: because of thermoregulatory behavior, lizards at high and low elevation share similar microclimatic niches (consistent with niche conservatism) while inhabiting distinct macroclimatic environments (consistent with niche divergence). Together, our results suggest that behavior can connect patterns of niche divergence and conservatism at different geographic scales and among traits.

Mechanical sensitivity and the dynamics of evolutionary rate shifts in biomechanical systems.

The influence of biophysical relationships on rates of morphological evolution is a cornerstone of evolutionary theory. Mechanical sensitivity-the correlation strength between mechanical output and the system's underlying morphological components-is thought to impact the evolutionary dynamics of form-function relationships, yet has rarely been examined. Here, we compare the evolutionary rates of the mechanical components of the four-bar linkage system in the raptorial appendage of mantis shrimp (Order Stomatopoda). This system's mechanical output (kinematic transmission (KT)) is highly sensitive to variation in its output link, and less sensitive to its input and coupler links. We found that differential mechanical sensitivity is associated with variation in evolutionary rate: KT and the output link exhibit faster rates of evolution than the input and coupler links to which KT is less sensitive. Furthermore, for KT and, to a lesser extent, the output link, rates of evolution were faster in 'spearing' stomatopods than 'smashers', indicating that mechanical sensitivity may influence trait-dependent diversification. Our results suggest that mechanical sensitivity can impact morphological evolution and guide the process of phenotypic diversification. The connection between mechanical sensitivity and evolutionary rates provides a window into the interaction between physical rules and the evolutionary dynamics of morphological diversification.

Untangling intra- and interspecific effects on body size clines reveals divergent processes structuring convergent patterns in Anolis lizards.

Bergmann's rule-the tendency for body size to increase in colder environments-remains controversial today, despite 150 years of research. Considerable debate has revolved around whether the rule applies within or among species. However, this debate has generally not considered that clade-level relationships are caused by both intra- and interspecific effects. In this article, we implement a novel approach that allows for the separation of intra- and interspecific components of trait-environment relationships. We apply this approach to body size clines in two Caribbean clades of Anolis lizards and discover that their similar body size gradients are constructed in very different ways. We find inverse Bergmann's clines-high-elevation lizards are smaller bodied-for both the cybotes clade on Hispaniola and the sagrei clade on Cuba. However, on Hispaniola, the inverse cline is driven by interspecific differences, whereas intraspecific variation is responsible for the inverse cline on Cuba. Our results suggest that similar body size clines can be constructed through differing evolutionary and ecological processes, namely, through local adaptation or phenotypic plasticity (intraspecific clines) and/or size-ordered spatial sorting (interspecific clines). We propose that our approach can help integrate a divided research program by focusing on how the combined effects of intra- and interspecific processes can enhance or erode clade-level relationships at large biogeographic scales.

Evolutionary stasis and lability in thermal physiology in a group of tropical lizards.

Understanding how quickly physiological traits evolve is a topic of great interest, particularly in the context of how organisms can adapt in response to climate warming. Adjustment to novel thermal habitats may occur either through behavioural adjustments, physiological adaptation or both. Here, we test whether rates of evolution differ among physiological traits in the cybotoids, a clade of tropical Anolis lizards distributed in markedly different thermal environments on the Caribbean island of Hispaniola. We find that cold tolerance evolves considerably faster than heat tolerance, a difference that results because behavioural thermoregulation more effectively shields these organisms from selection on upper than lower temperature tolerances. Specifically, because lizards in very different environments behaviourally thermoregulate during the day to similar body temperatures, divergent selection on body temperature and heat tolerance is precluded, whereas night-time temperatures can only be partially buffered by behaviour, thereby exposing organisms to selection on cold tolerance. We discuss how exposure to selection on physiology influences divergence among tropical organisms and its implications for adaptive evolutionary response to climate warming.

Amphibians exhibit extremely high hydric costs of respiration.

Terrestrial environments pose many challenges to organisms, but perhaps one of the greatest is the need to breathe while maintaining water balance. Breathing air requires thin, moist respiratory surfaces, and thus the conditions necessary for gas exchange are also responsible for high rates of water loss that lead to desiccation. Across the diversity of terrestrial life, water loss acts as a universal cost of gas exchange and thus imposes limits on respiration. Amphibians are known for being vulnerable to rapid desiccation, in part because they rely on thin, permeable skin for cutaneous respiration. Yet we have a limited understanding of the relationship between water loss and gas exchange within and among amphibian species. In this study, we evaluated the hydric costs of respiration in amphibians using the transpiration ratio, which is defined as the ratio of water loss (mol H2O d-1) to gas uptake (mol O2 d-1). A high ratio suggests greater hydric costs relative to the amount of gas uptake. We compared the transpiration ratio of amphibians with that of other terrestrial organisms to determine if amphibians had greater hydric costs of gas uptake relative to plants, insects, birds, and mammals. We also evaluated the effects of temperature, humidity, and body mass on the transpiration ratio both within and among amphibian species. We found that hydric costs of respiration in amphibians were two to four orders of magnitude higher than the hydric costs of plants, insects, birds, and mammals. We also discovered that larger amphibians had lower hydric costs than smaller amphibians, at both the species- and individual-level. Amphibians also reduced the hydric costs of respiration at warm temperatures, potentially reflecting adaptive strategies to avoid dehydration while also meeting the demands of higher metabolic rates. Our results suggest that cutaneous respiration is an inefficient mode of respiration that produces the highest hydric costs of respiration yet to be measured in terrestrial plants and animals. Yet, amphibians largely avoid these costs by selecting aquatic or moist environments, which may facilitate more independent evolution of water loss and gas exchange.

Comparison of hydric and thermal physiology in an environmentally diverse clade of Caribbean anoles.

As the world becomes warmer and precipitation patterns less predictable, organisms will experience greater heat and water stress. It is crucial to understand the factors that predict variation in thermal and hydric physiology among species. This study focuses on investigating the relationships between thermal and hydric diversity, and their environmental predictors, in a clade of Hispaniolan anole lizards, which are part of a broader Caribbean adaptive radiation. This clade, the 'cybotoid' anoles, occupies a wide range of thermal habitats (from sea level to several kilometers above it) and hydric habitats (such as xeric scrub, broadleaf forest, and pine forest), setting up the possibility for ecophysiological specialization among species. Among the thermal traits only cold tolerance is correlated with environmental temperature, and none of our climate variables were correlated with hydric physiology. Nevertheless, we found a negative relationship between heat tolerance (critical thermal maximum) and evaporative water loss at higher temperatures such that more heat tolerant lizards are also more desiccation tolerant at higher temperatures. This finding hints at shared thermal and hydric specialization at higher temperatures, underscoring the importance of considering the interactive effects of temperature and water balance in ecophysiological studies. While ecophysiological differentiation is a core feature of the anole adaptive radiation, our results suggest that close relatives in this lineage do not diverge in hydric physiology and only diverge partially in thermal physiology.

Viviparity imparts a macroevolutionary signature of ecological opportunity in the body size of female Liolaemus lizards.

Viviparity evolved ~115 times across squamate reptiles, facilitating the colonization of cold habitats, where oviparous species are scarce or absent. Whether the ecological opportunity furnished by such colonization reconfigures phenotypic diversity and accelerates evolution is unclear. We investigated the association between viviparity and patterns and rates of body size evolution in female Liolaemus lizards, the most species-rich tetrapod genus from temperate regions. Here, we discover that viviparous species evolve ~20% larger optimal body sizes than their oviparous relatives, but exhibit similar rates of body size evolution. Through a causal modeling approach, we find that viviparity indirectly influences body size evolution through shifts in thermal environment. Accordingly, the colonization of cold habitats favors larger body sizes in viviparous species, reconfiguring body size diversity in Liolaemus. The catalyzing influence of viviparity on phenotypic evolution arises because it unlocks access to otherwise inaccessible sources of ecological opportunity, an outcome potentially repeated across the tree of life.

Divergence in coloration and ecological speciation in the Anolis marmoratus species complex.

Adaptive divergence in coloration is expected to produce reproductive isolation in species that use colourful signals in mate choice and species recognition. Indeed, many adaptive radiations are characterized by differentiation in colourful signals, suggesting that divergent selection acting on coloration may be an important component of speciation. Populations in the Anolis marmoratus species complex from the Caribbean island of Guadeloupe display striking divergence in the colour and pattern of adult males that occurs over small geographic distances, suggesting strong divergent selection. Here we test the hypothesis that divergence in coloration results in reduced gene flow among populations. We quantify variation in adult male coloration across a habitat gradient between mesic and xeric habitats, use a multilocus coalescent approach to infer historical demographic parameters of divergence, and examine gene flow and population structure using microsatellite variation. We find that colour variation evolved without geographic isolation and in the face of gene flow, consistent with strong divergent selection and that both ecological and sexual selection are implicated. However, we find no significant differentiation at microsatellite loci across populations, suggesting little reproductive isolation and high levels of contemporary gene exchange. Strong divergent selection on loci affecting coloration probably maintains clinal phenotypic variation despite high gene flow at neutral loci, supporting the notion of a porous genome in which adaptive portions of the genome remain fixed whereas neutral portions are homogenized by gene flow and recombination. We discuss the impact of these findings for studies of colour evolution and ecological speciation.

A latitudinal gradient of deep-sea invasions for marine fishes.

Although the tropics harbor the greatest species richness globally, recent work has demonstrated that, for many taxa, speciation rates are faster at higher latitudes. Here, we explore lability in oceanic depth as a potential mechanism for this pattern in the most biodiverse vertebrates - fishes. We demonstrate that clades with the highest speciation rates also diversify more rapidly along the depth gradient, drawing a fundamental link between evolutionary and ecological processes on a global scale. Crucially, these same clades also inhabit higher latitudes, creating a prevailing latitudinal gradient of deep-sea invasions concentrated in poleward regions. We interpret these findings in the light of classic ecological theory, unifying the latitudinal variation of oceanic features and the physiological tolerances of the species living there. This work advances the understanding of how niche lability sculpts global patterns of species distributions and underscores the vulnerability of polar ecosystems to changing environmental conditions.

Phenotypic rate and state are decoupled in response to river-to-lake transitions in cichlid fishes.

Geographic access to isolated ecosystems is an important catalyst of adaptive radiation. Cichlid fishes repeatedly colonized rift, crater, and volcanic lakes from surrounding rivers. We test the "lake effect" on the phenotypic rate and state across 253 cichlid species. The rate of evolution was consistently higher (~10-fold) in lakes, and consistent across different dimensions of the phenotype. Rate shifts tended to occur coincident with or immediately following river-to-lake transitions, generally resulting in 2- to 5-fold faster rates than in the founding riverine lineage. By contrast, river- and lake-dwelling cichlids exhibit considerable overlap in phenotypes, generally with less disparity in lakes, but often different evolutionary optima. Taken together, these results suggest that lake radiations rapidly expand into niches largely already represented by ancestral riverine lineages, albeit in different frequencies. Lakes may provide ecological opportunity via ecological release (e.g., from predators/competitors) but need not be coupled with access to novel ecological niches.

Material properties of skin in the flying snake Chrysopelea ornata.

In snakes, the skin serves for protection, camouflage, visual signaling, locomotion, and its ability to stretch facilitates large prey ingestion. The flying snakes of the genus Chrysopelea are capable of jumping and gliding through the air, requiring additional functional demands: its skin must accommodate stretch in multiple directions during gliding and, perhaps more importantly, during high-speed, direct-impact landing. Is the skin of flying snakes specialized for gliding? Here, we characterized the material properties of the skin of Chrysopelea ornata and compared them with two nongliding species of colubrid snakes, Thamnophis sirtalis and Pantherophis guttatus, as well as with previously published values. The skin was examined using uniaxial tensile testing to measure stresses, and digital image correlation methods to determine strains, yielding metrics of strength, elastic modulus, strain energy, and extensibility. To test for loading orientation effects, specimens were tested from three orientations relative to the snake's long axis: lateral, circumferential, and ventral. Specimens were taken from two regions of the body, pre- and pos-tpyloric, to test for regional effects related to the ingestion of large prey. In comparison with T. sirtalis and P. guttatus, C. ornata exhibited higher post-pyloric and lower pre-pyloric extensibility in circumferential specimens. However, overall there were few differences in skin material properties of C. ornata compared to other species, both within and across studies, suggesting that the skin of flying snakes is not specialized for gliding locomotion. Surprisingly, circumferential specimens demonstrated lower strength and extensibility in pre-pyloric skin, suggesting less regional specialization related to large prey.

The Bogert effect, a factor in evolution.

Behavior is one of the major architects of evolution: by behaviorally modifying how they interact with their environments, organisms can influence natural selection, amplifying it in some cases and dampening it in others. In one of the earliest issues of Evolution, Charles Bogert proposed that regulatory behaviors (namely thermoregulation) shield organisms from selection and limit physiological evolution. Here, I trace the history surrounding the origin of this concept (now known as the "Bogert effect" or "behavioral inertia"), and its implications for physiological and evolutionary research throughout the 20th century. A key follow-up study in the early 21st century galvanized renewed interest in Bogert's classic ideas, and established a focus on slowdowns in the rate of evolution in response to regulatory behaviors. I illustrate recent progress on the Bogert effect in evolutionary research, and discuss the ecological variables that predict whether and how strongly the phenomenon unfolds. Based on these discoveries, I provide hypotheses for the Bogert effect across several scales: patterns of trait evolution within and among groups of species, spatial effects on the phenomenon, and its importance for speciation. I also discuss the inherent link between behavioral inertia and behavioral drive through an empirical case study linking the phenomena. Modern comparative approaches can help put the macroevolutionary implications of behavioral buffering to the test: I describe progress to date, and areas ripe for future investigation. Despite many advances, bridging microevolutionary processes with macroevolutionary patterns remains a persistent gap in our understanding of the Bogert effect, leaving wide open many avenues for deeper exploration.