Adams and Collyer (2019) evaluated the statistical performance of several approaches for quantifying morphological modularity and found that EMMLi had inflated type I error rates and a bias towards more complex models co...Adams and Collyer (2019) evaluated the statistical performance of several approaches for quantifying morphological modularity and found that EMMLi had inflated type I error rates and a bias towards more complex models compared to the Covariance Ratio (CR) approach. They suggested that this may have been at least partly driven by the fact that AICc values from EMMLi do not incorporate trait numbers, but this was not verified. Here I present a performance analysis of a trait-number corrected EMMLi approach ("EMMLip"), showing that this ameliorates rates of false discovery and produces conservative results that favor less complex models. The corrected EMMLi approach was effective at differentiating models of modularity with varying between- and within-module covariation especially when effect size or dataset size were sufficiently large. While CR tests remained more effective at specifically detecting overall modularity, I found that CR tests are sensitive to varying within/between module covariation, and in some cases had inflated model misspecification between 2- and 3-module hypotheses. With this minor correction (albeit incomplete), the combination of EMMLip and CR tests becomes the best available toolkit for detecting and contrasting modularity hypotheses. This toolkit is however still imperfect, and I discuss future avenues for improvements.
Sexually selected weapons can function as both combat tools and agonistic signals, depending on whether and how males assess rivals. We investigated the function of the enlarged male hindlegs in the New Guinean thorny de...Sexually selected weapons can function as both combat tools and agonistic signals, depending on whether and how males assess rivals. We investigated the function of the enlarged male hindlegs in the New Guinean thorny devil stick insect, Eurycantha calcarata in male-male and male-female interactions. Field and laboratory observations showed that larger males with proportionally larger hindlegs were more likely to win fights and subsequently mate. Behavioral sequence analyses and contest cost predictors indicated that males likely use a mutual assessment strategy. Surprisingly, males did not appear to use their hindlegs as signals of fighting ability. Rather, rival assessment may be mediated by tactile and chemical cues, as suggested by frequent antennation observed during contests. Hindlegs were employed mainly to deliver powerful squeezes in rare, escalated fights. During copulation, males also used hindlegs to stabilize their position, but females did not appear to resist, providing no evidence for a coercive function. These findings suggest that enlarged male hindlegs in E. calcarata serve purely as force-delivering combat tools rather than signaling structures. These results highlight how understanding the specific functions and contexts of weapon use provides critical insight into the diversification of sexually selected traits.
Theoretical work on the evolution of simple phenotypes with discrete states, such as RNA and protein structures, has shown that the most evolvable phenotypes have intermediate levels of mutational robustness, so long as...Theoretical work on the evolution of simple phenotypes with discrete states, such as RNA and protein structures, has shown that the most evolvable phenotypes have intermediate levels of mutational robustness, so long as we let neutral mutations cause the diversification of a population's mutationally-accessible phenotypic neighborhood. But whether this applies to more complex phenotypes is unclear. Here, I predict that for quantitative traits, intermediate levels of mutational robustness should boost evolvability whenever (i) the adaptive landscape changes after a period of stabilizing selection, and (ii) the phenotypic neighborhood of a genotype is determined in large part by the accumulation of cryptic alleles, the effects of which can be exposed to selection by epistatic mutations. I present evolutionary simulation models that support these predictions, provided that cryptic allele effects are not too large. Genetic diversity can also be concealed and exposed by incomplete environmental robustness (i.e., plasticity). With additional simulations, I show that evolvability can also be maximized with intermediate levels of environmental robustness, although in a more restrictive parameter space. So, as for discrete traits, quantitative trait robustness can be good for evolvability.
Host population dynamics influence viral evolution and host specialization. In a recent study, Visher et al. (2026) investigate the role of spatially structured host genotypes in the evolution of generalist and specialis...Host population dynamics influence viral evolution and host specialization. In a recent study, Visher et al. (2026) investigate the role of spatially structured host genotypes in the evolution of generalist and specialist viruses. Through experimentally evolving Plodia interpunctella granulovirus (PiGV) in inbred hosts of spatially clustered genotypes, the authors demonstrate that, even in heterogeneous populations, population clusters result in viral specialization and inhibit the rise of generalist viruses.
Mating interactions depend on traits expressed jointly by males and females, yet the extent to which each sex controls variation in these shared mating traits remains unclear. Because the expression of such traits (like...Mating interactions depend on traits expressed jointly by males and females, yet the extent to which each sex controls variation in these shared mating traits remains unclear. Because the expression of such traits (like mating latency, copulation duration, and sperm transfer) depends on both partners, their evolution is constrained by intersexual correlations yet facilitated by behavioral plasticity that allows each sex to adjust trait expression across environments. In this study we investigated whether shared mating traits are determined by male or female control or whether the observed outcomes result from interactions of the developmental environment of both partners. Drawing from the well-known mating system of the banded cricket, Gryllodes sigillatus, we used a fully factorial mating design using combinations of male and female partners reared at high or low same-sex density and tested how they shape shared pre- and post-copulatory traits. We found that female developmental density affected mating latency, with low-density females exhibiting longer latencies, suggesting female control and mate choice. In contrast, male developmental density affected sperm transfer and subsequent sperm storage, with males from high-density treatments transferring significantly more sperm consistent with adaptive ejaculate adjustment to sperm competition risk, as well as contributing to higher sperm storage in females. Copulation duration varied with female body mass but not density, indicating plastic responses to partner quality rather than social context. By partitioning environmental effects between the sexes, our study highlights how developmental context can be used to examine sex-specific contributions to shared mating traits.
Pigmentation patterns aid in camouflage and may also serve for communication. To understand the evolution of different types of within-feather pigmentation patterns, we studied the bird family Estrildidae, in which barre...Pigmentation patterns aid in camouflage and may also serve for communication. To understand the evolution of different types of within-feather pigmentation patterns, we studied the bird family Estrildidae, in which barred and spotted feathers are common. We found that gains and losses of barred or spotted plumage generally occurred simultaneously in both sexes. Sexual dimorphism evolved mostly via sex-specific losses of pigmentation patterns, rather than sex-specific gains, especially for spotted plumage. These results are consistent with evolution by genetic correlation between the sexes, followed by evolution of sex-specific pathways for trait reduction. Evolution by genetic correlation may be common for pigmentation patterns because they are not conspicuous to distant predators, and expressing them should rarely be detrimental. Previous work found correlated evolution of courtship dance and the overall complexity of estrildid pigmentation patterns, but we did not find this association for the extent of pigmentation patterns. Perhaps other properties of pigmentation patterns, and not their extent, evolve together with dance. Furthermore, the extent of barred or spotted plumage was not associated with colour ornamentation, nor with socioecological traits indicating need for camouflage or sexual signalling. This may reflect different and/or multiple functions for these within-feather pigmentation patterns in different species.
Intraguild predation, defined as predation between consumers of a shared resource, is widespread in nature and likely to be an important selective pressure shaping the evolution of defensive and foraging traits in many e...Intraguild predation, defined as predation between consumers of a shared resource, is widespread in nature and likely to be an important selective pressure shaping the evolution of defensive and foraging traits in many ecosystems. In many lakes of Southern Greenland, threespine stickleback (Gasterosteus aculeatus) can both compete with- and be preyed upon by Arctic charr (Salvelinus alpinus), meaning that stickleback trait evolution in this system may be influenced by intraguild predation. To test this, we analyse multivariate morphological variation within and across 36 stickleback populations, of which 18 sympatric with charr and 18 without charr. We find evidence of stickleback adaptation to intraguild predation particularly in defensive morphology, via longer spines and larger structural armour, though surprisingly not via more numerous lateral plates. Foraging morphology also responds to the presence of charr, with a shift towards littoral-adapted traits. These differences are revealed beyond single trait comparisons, as we discover diverging patterns of phenotypic (co)variance, with generally lower variation and dimensionality and stronger integration in populations with charr, indicative of selection for efficiency and erosion of variation. These findings corroborate the niche shift hypothesis for the evolutionary consequences of intraguild predation. Further, we find agreement between ecological theory of intraguild predation stability and our results on phenotypic change. Ultimately, our study provides a link between a multi-faceted ecological interaction-intraguild predation-and its high-dimensional evolutionary outcome-the multivariate phenotype-and showcases the power and importance of adopting a multivariate approach in evolutionary ecology studies.
Antagonistic trade-offs between early-life fitness and somatic maintenance are predicted to maintain variation in longevity, yet empirical support is scarce. Shephard et al. (2026) experimentally tested whether such trad...Antagonistic trade-offs between early-life fitness and somatic maintenance are predicted to maintain variation in longevity, yet empirical support is scarce. Shephard et al. (2026) experimentally tested whether such trade-offs are concealed under typical developmental conditions using Mexican spadefoot tadpoles (Spea multiplicata) reared on typical or atypical diets. The atypical diet revealed cryptic genetic variation in larval growth and a significant negative genetic correlation with telomere length, indicating a diet-dependent trade-off in somatic maintenance.
Hosts have evolved a variety of innate immune responses to pathogens. In many cases, hosts directly detect pathogen-associated molecular patterns (PAMPs) or pathogen effectors to trigger an immune response. However, host...Hosts have evolved a variety of innate immune responses to pathogens. In many cases, hosts directly detect pathogen-associated molecular patterns (PAMPs) or pathogen effectors to trigger an immune response. However, hosts may also detect pathogens indirectly through 'guarding', whereby immune receptors ('guards') monitor the effects of pathogens (e.g., modification of target cells) rather than the pathogens themselves. Guarding poses a different evolutionary challenge for pathogens than direct recognition of PAMPs, as replication may necessitate the modification or disruption of guarded host proteins ('guardees'). Recently, self-guarding has been discovered, in which the host target functions as both guard and guardee. Self-guarding appears to present an intractable problem for pathogens: modification of the host target may benefit replication, but also triggers an immune response. If self-guarding creates an apparently inescapable detection mechanism, why has self-guarding only recently been discovered? Here, we use mathematical models of within-host pathogen and immune dynamics to compare guarding and self-guarding architectures. We show that self-guarding leads to a more rapid immune response and faster pathogen suppression, but is also more prone to false-positive immune responses, likely imposing greater costs through autoimmunity. We therefore hypothesise that the greater potential for false-positive immune responses may limit the conditions under which self-guarding evolves.
Dispersal strategies are critical life history traits influencing bio-geographical, ecological, and evolutionary patterns and processes including range-expansion and contractions, abiotic and biotic interactions, and rat...Dispersal strategies are critical life history traits influencing bio-geographical, ecological, and evolutionary patterns and processes including range-expansion and contractions, abiotic and biotic interactions, and rates of speciation. Yet, dispersal strategies are challenging to study because they are complex suites of morphological, physiological, and behavioral traits. Thus, our understanding of how and why dispersal strategies evolve remains incomplete. Wing dimorphisms have evolved repeatedly within nearly every major insect order and are characterized by the co-occurrence of fully winged dispersing and short- or wingless non-dispersing morphs within species or populations, making them ideal model systems for studying the evolution of alternative dispersal strategies. Forty years ago, Derek Roff proposed that wing dimorphisms are a threshold trait which evolves as an intermediate step on the trajectory toward complete loss of flight, driven by the high physiological and life history costs of dispersal. Here, we synthesize recent mechanistic studies of wing-morph determination along with new phylogenetic reconstructions of wing dimorphism in the well-developed model Gryllus field crickets to update and refine our evolutionary hypotheses and identify critical areas for future inquiry.
Rodents constitute a large proportion of extant mammal species and are mostly a uniform brown-gray coloration to avoid detection by predators. A minority, however, have longitudinal dorsal stripes, the function of which...Rodents constitute a large proportion of extant mammal species and are mostly a uniform brown-gray coloration to avoid detection by predators. A minority, however, have longitudinal dorsal stripes, the function of which is unknown. Using a comparative approach, we explored whether striping in rodents is a form of background matching, an example of dazzle coloration, a social signal, or a signal to avoid hybridization with sympatric congeners. We found some evidence that striping is associated with small species, diurnality, and raptor and marginally with owl predation, suggesting it could be a form of dazzle coloration interfering with accurate interception by aerial predators approaching from above. There was no evidence that stripes are used in communication between conspecifics or for avoidance of hybridization. Our study provides the beginning of a functional underpinning to developmental studies of pattern formation in mammals.
Although traditionally viewed as opposing divergence, hybridization can have diverse evolutionary outcomes. Yet its role in lineage divergence remains unclear. To investigate this process, we examined the evolutionary hi...Although traditionally viewed as opposing divergence, hybridization can have diverse evolutionary outcomes. Yet its role in lineage divergence remains unclear. To investigate this process, we examined the evolutionary history of the Black-cheeked Gnateater (Conopophaga melanops) from the Atlantic Forest of eastern Brazil. The central subspecies, C. m. perspicillata, exhibits plumage traits intermediate to northern and southern populations, raising the question of whether this pattern reflects isolation-by-distance, recent hybridization, or past admixture. Using genome-wide markers from sequencing of ultraconserved elements along with phenotypic data, we assessed genetic structure and trait variation across the species' range. Our analyses reveal four population genetic clusters, with the two central clusters exhibiting clear signatures of historical admixture. Despite this admixture, central populations are genetically differentiated from northern and southern lineages and sing a distinct song, suggesting divergence following admixture or transgressive segregation. We propose that past hybridization followed by geographic isolation contributed to the formation of a divergent, reticulate lineage within the Black-cheeked Gnateater. This system provides an opportunity to investigate how gene flow and allopatric divergence interact to shape lineage diversity and offers a natural framework for testing the conditions under which reticulate lineages may emerge, persist, and diverge.
Organismal responses to changing thermal conditions can manifest in adaptive changes to numerous traits, but as yet, a systematic understanding of which traits are more labile than others is lacking. A recent study by Zi...Organismal responses to changing thermal conditions can manifest in adaptive changes to numerous traits, but as yet, a systematic understanding of which traits are more labile than others is lacking. A recent study by Zilio et al. (2026) demonstrates that physiological adaptation is sometimes more conserved than morphological and behavioral traits. Their study underscores the value of considering multiple trait types and comparative studies to understand phenotypic responses to novel selective pressures.
The evolution of different cranial kinesis types in modern birds has likely driven some spectacular adaptive radiations. However, in certain taxonomic groups, the functional role of new kinesis types remains unclear. Thi...The evolution of different cranial kinesis types in modern birds has likely driven some spectacular adaptive radiations. However, in certain taxonomic groups, the functional role of new kinesis types remains unclear. This is the case of furnariids, a family of neotropical passerines known for their stunning nest diversity and ecomorphological disparity. These birds exhibit two types of kinesis, each showing different degrees of craniofacial modularity and representing a distinct regime of evolutionary convergence in both beak and neurocranium. Nevertheless, the extrinsic factors shaping the traits associated with kinesis remain unknown. In this study, we assessed correlations between shape, kinesis type, and six ecological variables, alongside the role of kinesis in transitions toward domed nests and open habitats. We found that nest type and diet shaped the evolution of beak and cranial kinesis traits, while diet, foraging behavior, foraging strata, and primary habitat structure influenced neurocranium evolution. Specifically, the emergence of rhynchokinesis predicts the evolution of domed nests and the colonization of less forested environments. We discuss the role of modularity in the association between shape and ecology, the evolution of functional novelties in both kinetic groups, and the hierarchical order of ecomorphological diversity evolution through stages of the adaptive radiation.
Absence of phenotypic differentiation can challenge the study of diversification by concealing evolutionary divergence through morphological similarity. Such cryptic species diversity is found particularly often in extre...Absence of phenotypic differentiation can challenge the study of diversification by concealing evolutionary divergence through morphological similarity. Such cryptic species diversity is found particularly often in extreme environments, including arid mountain ranges. In the Hajar Mountains of southeastern Arabia, the Pristurus rupestris species complex comprises divergent lineages with notable phenotypic variation. Here, we investigate the evolutionary dynamics of this radiation by integrating geometric morphometric analyses with whole-genome sequencing and phylogenomic reconstructions. We analyzed five species-P. ali, P. assareen, P. feulneri, P. omanensis, and P. rupestris-including all previously identified genetic lineages within those species, and generated a new reference genome for an Old World sphaerodactylid gecko (P. rupestris sensu stricto). Morphometric analyses of comprehensive two- and three-dimensional morphological datasets indicate strong overlap in both body and head shape. Despite considerable phenotypic similarities, our results reveal deep interspecific divergences exceeding 16 million years and contrasting demographic histories. While we detected introgression restricted to specific lineage pairs, phenotypic crypticity persists across the entire complex, suggesting stabilizing selection in harsh environments. Our study highlights the P. rupestris species complex as an ideal model for investigating the role of environmental constraints in limiting morphological divergence and shaping diversification dynamics in arid mountain ecosystems.
The evolution of body size and its role in structuring communities has long been of interest. Here, we investigated the relationship between body size, phylogeny, and spatial patterns of distribution in three aquatic fro...The evolution of body size and its role in structuring communities has long been of interest. Here, we investigated the relationship between body size, phylogeny, and spatial patterns of distribution in three aquatic frog genera that have undergone extensive diversification in the Western Ghats of India. We used a comprehensive sampling protocol designed to uncover fine-scale divergence between allopatric and parapatric lineages. First, we found that body size differences between sympatrically distributed lineages were significantly greater than expected by chance in the stream-adapted genera, Nyctibatrachus and Micrixalus, with few assemblages across the entire Western Ghats containing two lineages of similar size. This pattern was not seen in Hylarana where sympatric species typically occupied different habitats, such as streams and ponds. Body size showed significant phylogenetic signal, but we found little overall evidence of phylogenetic clustering or overdispersion in all the genera. We also show that body size evolution in all three frog groups was not strongly affected by character displacement across closely related lineages. Taken together, this provides striking evidence for the potential role of species sorting, where competition leads to spatial assortment of body size, structuring aquatic frog assemblages in this tropical biodiversity hotspot.
Investigations into the genetic basis of color polymorphism have played a key role in our understanding of genetic architecture and the evolution of mating systems. Sulfur butterflies (Colias) have been models in this fi...Investigations into the genetic basis of color polymorphism have played a key role in our understanding of genetic architecture and the evolution of mating systems. Sulfur butterflies (Colias) have been models in this field, but also contain unsolved puzzles with respect to species boundaries and hybridization. We surveyed genomic variation across 5 years in a location where phenotypic intermediates between Colias eurytheme and C. eriphyle are observed, but ancestry variation of potential hybrids has not been quantified. Our results reveal individuals with hybrid ancestry roughly in proportion to the frequency of individuals of intermediate phenotype recorded in the wild. Individuals with hybrid ancestry were predominantly those with intermediate morphologies, but morphologically intermediate individuals were not always of hybrid origin, which raises alternative possibilities for the origin and maintenance of color variation in the system. Genomic regions differentiated between species are largely located on the Z chromosome, and we find more candidates on autosomes than in another Colias contact zone. The dynamics of hybridization in this system are highly variable through time, suggesting fertile avenues for future study into the maintenance of species boundaries in the face of temporally variable, climatically influenced, and pervasive hybridization.
Understanding how form-function relationships scale across levels of biological organization is essential for uncovering the mechanisms driving morphological and performance diversity. We examined the association between...Understanding how form-function relationships scale across levels of biological organization is essential for uncovering the mechanisms driving morphological and performance diversity. We examined the association between head shape and bite force in lacertid lizards across three hierarchical levels: individuals within species, species within the genus Podarcis, and species across the family Lacertidae. Using geometric morphometrics of dorsal and lateral head shape combined with bite force measurements, we tested whether the strength and direction of the form-function relationship is conserved across scales and whether body size mediates these patterns. Our analyses revealed significant associations between head shape and bite force at all levels, with body size exerting a strong but not exclusive influence. Importantly, while the form-function link persisted after removing allometric effects, the evolutionary trajectories of this relationship were not aligned across scales: regression vectors differed randomly rather than following consistent directions. These results indicate that performance consistently constrains head morphology, yet the evolutionary pathways linking form and function vary across scales, reflecting a flexible interplay between selective pressures, developmental constraints, and phylogenetic history.
Upon infecting a bacterial cell, temperate phages make a decision between lysis and lysogeny. While research has previously explored how phages sense environmental information to make this choice, most studies have focus...Upon infecting a bacterial cell, temperate phages make a decision between lysis and lysogeny. While research has previously explored how phages sense environmental information to make this choice, most studies have focused on modelling known mechanisms that impact the decision. These mechanisms tell us what environmental information the phage does respond to, but not what it should respond to, as the signals sensed by the phage may serve as proxies for other sources of information. Here, using a mechanism-agnostic population dynamics model, we find that irreversible phage binding to lysogens protects sensitive host cells from infection. This results in lysogens being an additional environmental factor that the phage should sense while making its decision to undergo lysis or lysogeny. Using this model, we derive a responsive lysogeny probability for phages that respond to both cell and lysogen densities optimized towards invading phage-occupied systems, and show that it is more capable of invading and resisting invasion than phage with fixed lysogeny probabilities across different environmental conditions.
Species are confined within spatial boundaries, which may be determined by various factors. An understudied one is the genetic non-independence of traits, which can reduce the adaptive potential. We tested this hypothesi...Species are confined within spatial boundaries, which may be determined by various factors. An understudied one is the genetic non-independence of traits, which can reduce the adaptive potential. We tested this hypothesis on elevational gradients, where environmental change occurs over short geographic distances. We collected seeds of three montane and three alpine Brassicaceae species, each from its core and both the lower and upper range edges, and reared offspring plants of matrilines in the greenhouse under two thermal treatments: warm and control. We measured key growth and leaf traits that diverge across elevation in Brassicaceae and estimated the genetic covariance (G-)matrix for each population and growth treatment. Genetic variances were reduced in lower-edge populations of montane species, while those of alpine species did not differ over the elevational gradients but were consistently low. Genetic integration-the reduction in evolvability due to genetic covariances-differed little over the elevational gradients and was generally strong. Similarly, the evolutionary potential ("evolvability") varied little and was low across populations. Finally, populations had substantial genetic variation in the direction of divergence along elevational gradients. Overall, our findings suggest that substantial genetic integration restricts population divergence along certain trait axes and reduces the evolutionary potential, which may hinder the evolution of species' ranges.