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Full-Text Articles in Evolution

Multilevel Selection In The Margins: A Review Of Its Representation In Undergraduate Biology Textbooks, C. A. Greene, S. J. Mcpeek, L. Mitchem, A. B. Clark, Vincent A. Formica Nov 2025

Multilevel Selection In The Margins: A Review Of Its Representation In Undergraduate Biology Textbooks, C. A. Greene, S. J. Mcpeek, L. Mitchem, A. B. Clark, Vincent A. Formica

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Multilevel selection (MLS) is a foundational framework that expands the concept of natural selection beyond the traditional focus on individuals or genes to larger units, like groups. Despite strong empirical support and applications in a broad range of biological fields, MLS, often referred to as group selection, has a controversial history, which has impeded its wide use and led to incorrect inferences on the sources and strength of selection. Here, we explore how MLS is currently presented to undergraduate biology students by analyzing the content of nine commonly used Evolutionary Biology and Animal Behavior textbooks. We ask whether and how …


Age-Dependent Changes In Reproductive Behaviour And Success In A Long-Lived Beetle (Bolitotherus Cornutus), C. A. Greene, L. D. Mitchem, Vitor Dos Anjos , '21, E. D. Brodie Iii, Vincent A. Formica Sep 2025

Age-Dependent Changes In Reproductive Behaviour And Success In A Long-Lived Beetle (Bolitotherus Cornutus), C. A. Greene, L. D. Mitchem, Vitor Dos Anjos , '21, E. D. Brodie Iii, Vincent A. Formica

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Age-related changes in behaviors often result in shifts in an individual’s competitive ability, reproductive success, and fitness. However, how mating behaviors, aggression, and mating success simultaneously change as individuals age in natural populations remains understudied. Investigating these age-dependent effects in wild populations could reveal how age-specific patterns shape social dynamics and influence natural selection. In this study, we ask how behaviors associated with mating and reproductive success vary across age in a wild metapopulation of a long-lived beetle, Bolitotherus cornutus. We used both cross-sectional and longitudinal data from a multi-year field study of individually marked beetles to investigate how …


Male Mating Preference For Larger Females Does Not Vary Among Age Classes In The Long-Lived Beetle Bolitotherus Cornutus, G. M. Jiron, C. A. Greene, E. D. Brodie Iii, Vincent A. Formica Aug 2025

Male Mating Preference For Larger Females Does Not Vary Among Age Classes In The Long-Lived Beetle Bolitotherus Cornutus, G. M. Jiron, C. A. Greene, E. D. Brodie Iii, Vincent A. Formica

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Theory and past experimental work suggest that as males age, the strength of their mate preference should decrease. However, the empirical work investigating this question has primarily been conducted in insects that have very short life spans and often live for just a single mating season. This leaves a gap in our understanding of the relationship between male mate preference and age across taxa, as age can conflate with other ecological changes in a single mating season. In this study, we ask how the strength of preference for large female body size changes as males age in a long-lived insect, …


​​Site Fidelity Is Associated With The Social Experience Of Forked Fungus Beetles (Bolitotherus Cornutus), Joo Yeon Kim , '22, C. W. Wood, E. D. Brodie Iii, Vincent A. Formica Jul 2025

​​Site Fidelity Is Associated With The Social Experience Of Forked Fungus Beetles (Bolitotherus Cornutus), Joo Yeon Kim , '22, C. W. Wood, E. D. Brodie Iii, Vincent A. Formica

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Consistently returning to or remaining within a specific location, a behavior known as site fidelity, is a useful metric for quantifying animal space use. When multiple individuals display similar site fidelity patterns, they are likely to interact, shaping the degree and quality of their social interactions. However, the link between the expression of site fidelity behavior and an individual’s social experience has not been thoroughly investigated. In this study we examined (1) how intrinsic individual properties (body size and sex) predicted variation in site fidelity and (2) whether site fidelity was linked to the social experience of individuals in wild …


The Influence Of Resource Patch Condition And Size On Social Interactions, Reproductive Behaviors, And Oviposition In A Fungivorous Beetle, Lillian J. Fornoff , '21, C. A. Stahlmann Roeder, E. D. Brodie Iii, Vincent A. Formica Jul 2024

The Influence Of Resource Patch Condition And Size On Social Interactions, Reproductive Behaviors, And Oviposition In A Fungivorous Beetle, Lillian J. Fornoff , '21, C. A. Stahlmann Roeder, E. D. Brodie Iii, Vincent A. Formica

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Variation among resource patches can structure spatial patterns of social behavior. Individual preferences and requirements for heterogeneous resources can lead to differences in where behaviors take place and which kinds of individuals interact. In this study we examined whether properties of habitat patches predict presence of and interactions among wild forked fungus beetles (Bolitotherus cornutus). B. cornutus use shelf-like fruiting bodies of wood decaying fungi (brackets) as social arenas, courtship and mating sites, oviposition locations, and food sources. Brackets on a single log vary in size and condition and represent a dynamic resource as individual brackets decay over …


Random-Effects Substitution Models For Phylogenetics Via Scalable Gradient Approximations, A. F. Magee, A. J. Holbrook, J. E. Pekar, Itzue W. Caviedes-Solis, F. A. Matsen Iv, G. Baele, J. O. Wertheim, X. Ji, P. Lemey, M. A. Suchard May 2024

Random-Effects Substitution Models For Phylogenetics Via Scalable Gradient Approximations, A. F. Magee, A. J. Holbrook, J. E. Pekar, Itzue W. Caviedes-Solis, F. A. Matsen Iv, G. Baele, J. O. Wertheim, X. Ji, P. Lemey, M. A. Suchard

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Phylogenetic and discrete-trait evolutionary inference depend heavily on an appropriate characterization of the underlying character substitution process. In this paper, we present random-effects substitution models that extend common continuous-time Markov chain models into a richer class of processes capable of capturing a wider variety of substitution dynamics. As these random-effects substitution models often require many more parameters than their usual counterparts, inference can be both statistically and computationally challenging. Thus, we also propose an efficient approach to compute an approximation to the gradient of the data likelihood with respect to all unknown substitution model parameters. We demonstrate that this approximate …


Individual And Population Age Impact Social Behavior And Network Structure In A Long-Lived Insect, P. A. Cook, R. A. Costello, Vincent A. Formica, E. D. Brodie Iii Nov 2023

Individual And Population Age Impact Social Behavior And Network Structure In A Long-Lived Insect, P. A. Cook, R. A. Costello, Vincent A. Formica, E. D. Brodie Iii

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Social behaviors vary among individuals, and social networks vary among groups. Understanding the causes of such variation is important for predicting or altering ecological processes such as infectious disease outbreaks. Here, we ask whether age contributes to variation in social behavior at multiple levels of organization: within individuals over time, among individuals of different ages, among local social environments, and among populations. We used experimental manipulations of captive populations and a longitudinal dataset to test whether social behavior is associated with age across these levels in a long-lived insect, the forked fungus beetle (Bolitotherus cornutus). In cross-sectional analyses, …


Multilevel Selection On Social Network Traits Differs Between Sexes In Experimental Populations Of Forked Fungus Beetles, R. A. Costello, P. A. Cook, E. D. Brodie Iii, Vincent A. Formica Jan 2023

Multilevel Selection On Social Network Traits Differs Between Sexes In Experimental Populations Of Forked Fungus Beetles, R. A. Costello, P. A. Cook, E. D. Brodie Iii, Vincent A. Formica

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Both individual and group behavior can influence individual fitness, but multilevel selection is rarely quantified on social behaviors. Social networks provide a unique opportunity to study multilevel selection on social behaviors, as they describe complex social traits and patterns of interaction at both the individual and group levels. In this study, we used contextual analysis to measure the consequences of both individual network position and group network structure on individual fitness in experimental populations of forked fungus beetles (Bolitotherus cornutus) with two different resource distributions. We found that males with high individual connectivity (strength) and centrality (betweenness) had …


Nature, Data, And Power: How Hegemonies Shaped This Special Section, A. Kamath, B. Velocci, A. Wesner, N. Chen, Vincent A. Formica, B. Subramaniam, M. Rebolleda-Gómez Jul 2022

Nature, Data, And Power: How Hegemonies Shaped This Special Section, A. Kamath, B. Velocci, A. Wesner, N. Chen, Vincent A. Formica, B. Subramaniam, M. Rebolleda-Gómez

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Systems of oppression—racism, colonialism, misogyny, cissexism, ableism, heteronormativity, and more—have long shaped the content and practice of science. But opportunities to reckon with these influences are rarely found within academic science, even though such critiques are well developed in the social sciences and humanities. In this special section, we attempt to bring cross-disciplinary conversations among ecology, evolution, behavior, and genetics on the one hand and critical perspectives from the social sciences and humanities on the other into the pages—and in front of the readers—of a scientific journal. In this introduction to the special section, we recount and reflect on the …


Mycophagous Beetle Females Do Not Behave Competitively During Intrasexual Interactions In Presence Of A Fungal Resource, L. D. Mitchem, Vincent A. Formica, R. Debray, Dana E. Homer , '21, E. D. Brodie Iii Jun 2022

Mycophagous Beetle Females Do Not Behave Competitively During Intrasexual Interactions In Presence Of A Fungal Resource, L. D. Mitchem, Vincent A. Formica, R. Debray, Dana E. Homer , '21, E. D. Brodie Iii

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Intrasexual interactions can determine which individuals within a population have access to limited resources. Despite their potential importance on fitness generally and mating success especially, female–female interactions are not often measured in the same species where male–male interactions are well-defined. In this study, we characterized female–female interactions in Bolitotherus cornutus, a mycophagous beetle species native to Northeastern North America. We used dyadic, behavioral assays to determine whether females perform directly aggressive or indirectly exclusionary competitive behaviors. Polypore shelf fungus, an important food and egg-laying resource for B. cornutus females, is patchily distributed and of variable quality, so we tested …


Group And Individual Social Network Metrics Are Robust To Changes In Resource Distribution In Experimental Populations Of Forked Fungus Beetles, R. A. Costello, P. A. Cook, Vincent A. Formica, E. D. Brodie Iii Apr 2022

Group And Individual Social Network Metrics Are Robust To Changes In Resource Distribution In Experimental Populations Of Forked Fungus Beetles, R. A. Costello, P. A. Cook, Vincent A. Formica, E. D. Brodie Iii

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1. Social interactions drive many important ecological and evolutionary processes. It is therefore essential to understand the intrinsic and extrinsic factors that underlie social patterns. A central tenet of the field of behavioural ecology is the expectation that the distribution of resources shapes patterns of social interactions. 2. We combined experimental manipulations with social network analyses to ask how patterns of resource distribution influence complex social interactions. 3. We experimentally manipulated the distribution of an essential food and reproductive resource in semi-natural populations of forked fungus beetles Bolitotherus cornutus. We aggregated resources into discrete clumps in half of the populations …


Male Competition Reverses Female Preference For Male Chemical Cues, Z. Vilella-Pacheco, L. D. Mitchem, Vincent A. Formica, E. D. Brodie Iii Mar 2021

Male Competition Reverses Female Preference For Male Chemical Cues, Z. Vilella-Pacheco, L. D. Mitchem, Vincent A. Formica, E. D. Brodie Iii

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Females must choose among potential mates with different phenotypes in a variety of social contexts. Many male traits are inherent and unchanging, but others are labile to social context. Competition, for example, can cause physiological changes that reflect recent wins and losses that fluctuate throughout time. We may expect females to respond differently to males depending on the outcome of their most recent fight. In Bolitotherus cornutus (forked fungus beetles), males compete for access to females, but copulation requires female cooperation. In this study, we use behavioral trials to determine whether females use chemical cues to differentiate between males and …


Actuarial Senescence In A Dimorphic Bird: Different Rates Of Ageing In Morphs With Discrete Reproductive Strategies, M. L. Grunst, A. S. Grunst, Vincent A. Formica, M. L. Korody, A. M. Betuel, M. Barcelo-Serra, R. A. Gonser, E. M. Tuttle Nov 2018

Actuarial Senescence In A Dimorphic Bird: Different Rates Of Ageing In Morphs With Discrete Reproductive Strategies, M. L. Grunst, A. S. Grunst, Vincent A. Formica, M. L. Korody, A. M. Betuel, M. Barcelo-Serra, R. A. Gonser, E. M. Tuttle

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It is often hypothesized that intra-sexual competition accelerates actuarial senescence, or the increase in mortality rates with age. However, an alternative hypothesis is that parental investment is more important to determining senescence rates. We used a unique model system, the white-throated sparrow (Zonotrichia albicollis), to study variation in actuarial senescence. In this species, genetically determined morphs display discrete mating strategies and disassortative pairing, providing an excellent opportunity to test the predictions of the above hypotheses. Compared to tan-striped males, white-striped males are more polygynous and aggressive, and less parental. Tan-striped females receive less parental support, and invest more into parental …


Morph-Specific Patterns Of Reproductive Senescence: Connections To Discrete Reproductive Strategies, A. S. Grunst, M. L. Grunst, Vincent A. Formica, M. L. Korody, A. M. Betuel, M. Barcelo-Serra, S. Ford, R. A. Gonser, E. M. Tuttle Jun 2018

Morph-Specific Patterns Of Reproductive Senescence: Connections To Discrete Reproductive Strategies, A. S. Grunst, M. L. Grunst, Vincent A. Formica, M. L. Korody, A. M. Betuel, M. Barcelo-Serra, S. Ford, R. A. Gonser, E. M. Tuttle

Biology Faculty Works

How reproductive strategies contribute to patterns of senescence in natural populations remains contentious. We studied reproductive senescence in the dimorphic white-throated sparrow, an excellent species for exploring this issue. Within both sexes the morphs use distinct reproductive strategies, and disassortative pairing by morph results in pair types with distinct parental systems. White morph birds are more colorful and aggressive than tan counterparts, and white males compete for extrapair matings, whereas tan males are more parental. Tan males and white females share parental care equally, whereas white males provide little parental support to tan females. We found morph-specific patterns of reproductive …


Multiple Signaling Functions Of Song In A Polymorphic Species With Alternative Reproductive Strategies, M. L. Grunst, A. S. Grunst, Vincent A. Formica, R. A. Gonser, E. M. Tuttle Jan 2018

Multiple Signaling Functions Of Song In A Polymorphic Species With Alternative Reproductive Strategies, M. L. Grunst, A. S. Grunst, Vincent A. Formica, R. A. Gonser, E. M. Tuttle

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Vocal traits can be sexually selected to reflect male quality, but may also evolve to serve additional signaling functions. We used a long-term dataset to examine the signaling potential of song in dimorphic white-throated sparrows (Zonotrichia albicollis). We investigated whether song conveys multifaceted information about the vocalizing individual, including fitness, species identity, individual identity, and morph. We also evaluated whether song traits correlate differently with fitness in the two morphs, as the more promiscuous strategy of white, relative to tan, morph males might impose stronger sexual selection. Males with high song rates achieved higher lifetime reproductive success, and this pattern …


No Effect Of Host Species On Phenoloxidase Activity In A Mycophagous Beetle, Vincent A. Formica, Amanda Kar-Men Chan , '16 Oct 2015

No Effect Of Host Species On Phenoloxidase Activity In A Mycophagous Beetle, Vincent A. Formica, Amanda Kar-Men Chan , '16

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Ecological immunology is an interdisciplinary field that helps elucidate interactions between the environment and immune response. The host species individuals experience have profound effects on immune response in many species of insects. However, this conclusion comes from studies of herbivorous insects even though species of mycophagous insects also inhabit many different host species. The goal of this study was to determine if fungal host species as well as individual, sex, body size, and host patch predict one aspect of immune function, phenoloxidase activity (PO). We sampled a metapopulation of Bolitotherus cornutus, a mycophagous beetle in southwestern Virginia. B. cornutus live …


Surprisingly Little Population Genetic Structure In A Fungus-Associated Beetle Despite Its Exploitation Of Multiple Hosts, Corlett W. Wood , '08, H. M. Donald, Vincent A. Formica, E. D. Brodie Iii Jun 2013

Surprisingly Little Population Genetic Structure In A Fungus-Associated Beetle Despite Its Exploitation Of Multiple Hosts, Corlett W. Wood , '08, H. M. Donald, Vincent A. Formica, E. D. Brodie Iii

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In heterogeneous environments, landscape features directly affect the structure of genetic variation among populations by functioning as barriers to gene flow. Resource-associated population genetic structure, in which populations that use different resources (e.g., host plants) are genetically distinct, is a well-studied example of how environmental heterogeneity structures populations. However, the pattern that emerges in a given landscape should depend on its particular combination of resources. If resources constitute barriers to gene flow, population differentiation should be lowest in homogeneous landscapes, and highest where resources exist in equal proportions. In this study, we tested whether host community diversity affects population genetic …


Morphological Correlates Of A Combat Performance Trait In The Forked Fungus Beetle, Bolitotherus Cornutus, K. M. Benowitz, E. D. Brodie Iii, Vincent A. Formica Aug 2012

Morphological Correlates Of A Combat Performance Trait In The Forked Fungus Beetle, Bolitotherus Cornutus, K. M. Benowitz, E. D. Brodie Iii, Vincent A. Formica

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Combat traits are thought to have arisen due to intense male-male competition for access to females. While large and elaborate weapons used in attacking other males have often been the focus of sexual selection studies, defensive traits (both morphological and performance) have received less attention. However, if defensive traits help males restrict access to females, their role in the process of sexual selection could also be important. Here we examine the morphological correlates of grip strength, a defensive combat trait involved in mate guarding, in the tenebrionid beetle Bolitotherus cornutus. We found that grip strength was repeatable and differed between …