Weapon performance drives weapon evolution.

Many sexually selected traits function as weapons, and these weapons can be incredibly diverse. However, the factors underlying weapon diversity among species remain poorly understood, and a fundamental hypothesis to explain this diversity remains untested. Although weapons can serve multiple functions, an undeniably important function is their role in fights. Thus, a crucial hypothesis is that weapon diversification is driven by the evolution of weapon modifications that provide an advantage in combat (e.g. causing more damage). Here, we test this fighting-advantage hypothesis using data from 17 species of coreid bugs. We utilize the fact that male-male combat in coreids often results in detectable damage, allowing us to link different weapon morphologies to different levels of damage among species. We find that certain weapon morphologies inflict much more damage than others, strongly supporting the fighting-advantage hypothesis. Moreover, very different weapon morphologies can inflict similarly severe amounts of damage, leading to a weapon performance landscape with multiple performance peaks. This multi-peak pattern could potentially drive different lineages towards divergent weapon forms, further increasing weapon diversity among species. Overall, our results may help explain how sexually selected weapons have evolved into the diversity of forms seen today.

Trade-offs between weapons and testes do not manifest at high social densities.

Social conditions can alter the allocation of resources to reproductive traits. For example, an increase in social density during development is frequently associated with an increase in the testes mass of males. Sperm competition theory assumes that increased investment in testes should come at the expense of investing into precopulatory traits, such as sexually selected weaponry. However, much remains unknown about the role of the social context on the concurrent, relative investment in both testes and weapons. We found that the leaf-footed cactus bug, Narnia femorata (Hemiptera: Coreidae), grew nearly 20% larger testes when raised in high social densities. In addition to manipulating social density, we used autotomy (limb loss) to limit investment in their hindlimb weapon during development. At low densities, we found that those that lost a weapon during development grew larger testes by adulthood, supporting previous work demonstrating a weapons-testes trade-off. However, at high social densities, males that dropped a hindlimb did not grow larger testes, though testes were already large at this density. These results underscore the importance of the social context to resource allocation patterns within the individual.

A weapons-testes trade-off in males is amplified in female traits.

Sexually selected weapons are assumed to trade off with traits related to ejaculates, such as testes. However, remarkably little is known about what governs resource allocation and why trade-offs are found in some cases and not others. Often-used models depict competitive allocation occurring within the functional grouping of traits (e.g. reproduction); however, other factors including tissue expense and developmental timing may influence allocation. Experimental comparisons of investment across the sexes have the potential to illuminate allocation rules, because the sexes do not always use traits for the same functions. Here, we capitalize upon a species where females have weapons-testes homologues. We report that a documented trade-off in investment between hind-limb weapons and testes in leaf-footed cactus bugs, Narnia femorata, is even more pronounced in female hind limbs and ovaries. Female hind limbs in this species do not share the clear reproductive function of male hind limbs; therefore, this trade-off spans trait functional groups. Such patterns of investment suggest that future studies of reproductive trade-offs should consider factors such as tissue expense and developmental timing.

Social Information on Fear and Food Drives Animal Grouping and Fitness.

Empirical studies in select systems suggest that social information-the incidental or deliberate information produced by animals and available to other animals-can fundamentally shape animal grouping behavior. However, to understand the role of social information in animal behavior and fitness, we must establish general theory that quantifies effects of social information across ecological contexts and generates expectations that can be applied across systems. Here we used dynamic state variable modeling to isolate effects of social information about food and predators on grouping behavior and fitness. We characterized optimal behavior from a set of strategies that included grouping with different numbers of conspecifics or heterospecifics and the option to forage or be vigilant over the course of a day. We show that the use of social information alone increases grouping behavior but constrains group size to limit competition, ultimately increasing individual fitness substantially across various ecological contexts. We also found that across various contexts, foraging in mixed-species groups is generally better than foraging in conspecific groups, supporting recent theory on competition-information quality trade-offs. Our findings suggest that multiple forms of social information shape animal grouping and fitness, which are sensitive to resource availability and predation pressure that determine information usefulness.

Phasmid species that inhabit colder environments are less likely to have the ability to fly.

A vast majority of insects can fly, but some cannot. Flight generally increases how far an individual can travel to access mates, enables the exploitation of additional food resources, and aids in predator avoidance. Despite its functional significance, much remains unknown about the factors that influence the evolution of flight. Here, I use phylogenetic comparative methods to investigate whether average annual temperature or wind speed, two components of the flying environment, is correlated with the evolution of flight using data from 107 species of stick and leaf insects (Insecta: Phasmatodea). I find no association between wind speed and flying ability in this clade. However, I find that colder temperatures are associated with the lack of flying ability. This pattern may be explained by the additional metabolic costs required for insects to fly when it is cold. This finding contradicts previous patterns observed in other insect groups and supports the hypothesis that cold temperatures can influence the evolution of flight.

Selection on weapon allometry in the wild.

Allometry is the scaling relationship between a trait and body size. This relationship can often explain considerable morphological variation within and among species. Nevertheless, much remains unknown about the factors that underlie allometric patterns. For example, when different allometric relationships are observed amongst closely related species, these differences are regularly considered to be products of selection. However, directional selection on allometry (particularly the slope) has rarely been tested and observed in natural populations. Here, we investigate selection on the scaling relationship between weapon size and body size (i.e., weapon allometry) in a wild population of giant mesquite bugs, Pachylis neocalifornicus (previously Thasus neocalifornicus). Males in this species use their weapons (enlarged femurs) to compete with one another over access to resources and females. We found that large males with relatively large weapons successfully secured access to mates. However, we also found that small males with relatively small weapons could access mates as well. These two patterns together can increase the allometric slope of the sexually selected weapon, suggesting a straightforward process by which the allometric slope can evolve.

Why are animals conspicuously colored? Evolution of sexual versus warning signals in land vertebrates.

Conspicuous colors (e.g., red, yellow, blue) have evolved numerous times across animals. But the function of this coloration can differ radically among species. Many species use this coloration as a sexual signal to conspecifics, whereas others use it as a warning signal to predators. Why do different species evolve conspicuous coloration in association with one function as opposed to the other? We address this question in terrestrial vertebrates (tetrapods) using phylogenetic approaches, and test whether day-night activities of species help determine these patterns. Using phylogenetic logistic regression, we found that conspicuous, sexually dimorphic coloration is significantly associated with diurnal lineages (e.g., many birds and lizards). By contrast, the evolution of warning signals was significantly associated with large-scale clades that were ancestrally nocturnal (e.g., snakes, amphibians), regardless of the current diel activity of species. Overall, we show that the evolution of conspicuous coloration as warning signals or sexual signals is influenced by the ecology of species, both recently and in the ancient past.

Do sexually selected weapons drive diversification?

Sexual selection is often thought to promote speciation. This expectation is largely driven by the fact that sexually selected traits can influence mating patterns and contribute to reproductive isolation. Indeed, some comparative studies have shown that clades with sexually selected traits have increased rates of speciation and diversification. However, these studies have almost exclusively focused on one mechanism of sexual selection: female choice. Another widespread mechanism is male-male competition. Few empirical studies (if any) have investigated the role of this alternative mechanism in driving diversification. Nevertheless, recent reviews have suggested that male-male competition can increase speciation rates. Here, we investigated whether traits associated with precopulatory male-male competition (i.e., sexually selected weapons) have promoted speciation and diversification in insects. We focused on three clades with both weapons and suitable phylogenies: leaf-footed and broad-headed bugs (Coreidae+Alydidae; ∼2850 species), stick insects and relatives (Phasmatodea; ∼3284 species), and scarab beetles (Scarabaeoidea; ∼39,717 species). We found no evidence that weapon-bearing lineages in these clades have higher rates of speciation or diversification than their weaponless relatives. Thus, our results suggest that precopulatory male-male competition may not have strong, general effects on speciation and diversification in insects, a group encompassing ∼60% of all described species.

Leaf-footed bugs possess multiple hidden contrasting color signals, but only one is associated with increased body size.

Antipredatory displays that incorporate hidden contrasting coloration are found in a variety of different animals. These displays are seen in organisms that have drab coloration at rest, but when disturbed reveal conspicuous coloration. Examples include the bright abdomens of mountain katydids and the colorful underwings of hawk moths. Such hidden displays can function as secondary defenses, enabling evasion of a pursuant predator. To begin to understand why some species have these displays while others do not, we conducted phylogenetic comparative analyses to investigate factors associated with the evolution of hidden contrasting coloration in leaf-footed bugs. First, we investigated whether hidden contrasting coloration was associated with body size because these displays are considered to be more effective in larger organisms. We then investigated whether hidden contrasting coloration was associated with an alternative antipredatory defense, in this case rapid autotomy. We found that leaf-footed bugs with hidden contrasting coloration tended to autotomize more slowly, but this result was not statistically significant. We also found that the presence of a body size association was dependent upon the form of the hidden color display. Leaf-footed bugs that reveal red/orange coloration were the same size, on average, as species without a hidden color display. However, species that reveal white patches on a black background were significantly larger than species without a hidden color display. These results highlight the diversity of forms that hidden contrasting color signal can take, upon which selection may act differently.

The evolution of autotomy in leaf-footed bugs.

Sacrificing body parts is one of many behaviors that animals use to escape predation. This trait, termed autotomy, is classically associated with lizards. However, several other taxa also autotomize, and this trait has independently evolved multiple times throughout Animalia. Despite having multiple origins and being an iconic antipredatory trait, much remains unknown about the evolution of autotomy. Here, we combine morphological, behavioral, and genomic data to investigate the evolution of autotomy within leaf-footed bugs and allies (Insecta: Hemiptera: Coreidae + Alydidae). We found that the ancestor of leaf-footed bugs autotomized and did so slowly; rapid autotomy (<2 min) then arose multiple times. The ancestor likely used slow autotomy to reduce the cost of injury or to escape nonpredatory entrapment but could not use autotomy to escape predation. This result suggests that autotomy to escape predation is a co-opted benefit (i.e., exaptation), revealing one way that sacrificing a limb to escape predation may arise. In addition to identifying the origins of rapid autotomy, we also show that across species variation in the rates of autotomy can be explained by body size, distance from the equator, and enlargement of the autotomizable appendage.

The ecology and evolution of autotomy.

Autotomy, the self-induced loss of a body part, occurs throughout Animalia. A lizard dropping its tail to escape predation is an iconic example, however, autotomy occurs in a diversity of other organisms. Octopuses can release their arms, crabs can drop their claws, and bugs can amputate their legs. The diversity of organisms that can autotomize body parts has led to a wealth of research and several taxonomically focused reviews. These reviews have played a crucial role in advancing our understanding of autotomy within their respective groups. However, because of their taxonomic focus, these reviews are constrained in their ability to enhance our understanding of autotomy. Here, we aim to synthesize research on the ecology and evolution of autotomy throughout Animalia, building a unified framework on which future studies can expand. We found that the ability to drop an appendage has evolved multiple times throughout Animalia and that once autotomy has evolved, selection appears to act on the removable appendage to increase the efficacy and/or efficiency of autotomy. This could explain why some autotomizable body parts are so elaborate (e.g. brightly coloured). We also show that there are multiple benefits, and variable costs, associated with autotomy. Given this variation, we generate an economic theory of autotomy (modified from the economic theory of escape) which makes predictions about when an individual should resort to autotomy. Finally, we show that the loss of an autotomizable appendage can have numerous consequences on population and community dynamics. By taking this broad taxonomic approach, we identified patterns of autotomy that transcend specific lineages and highlight clear directions for future research.

Males that drop a sexually selected weapon grow larger testes.

Costly sexually selected weapons are predicted to trade off with postcopulatory traits, such as testes. Although weapons can be important for achieving access to females, individuals of some species can permanently drop (i.e. autotomize) their weapons, without regeneration, to escape danger. We capitalized on this natural behavior to experimentally address whether the loss of a sexually selected weapon leads to increased testes investment in the leaf-footed cactus bug, Narnia femorata Stål (Hemiptera: Coreidae). In a second experiment, we measured offspring production for males that lost a weapon during development. As predicted, males that dropped a hind limb during development grew significantly larger testes than the control treatments. Hind-limb autotomy did not result in the enlargement of other nearby traits. Our results are the first to experimentally demonstrate that males compensate for natural weapon loss by investing more in testes. In a second experiment we found that females paired with males that lost a hind limb had 40% lower egg hatching success than females paired with intact males, perhaps because of lower mating receptivity to males with a lost limb. Importantly, in those cases where viable offspring were produced, males missing a hind limb produced 42% more offspring than males with intact limbs. These results suggest that the loss of a hind-limb weapon can, in some cases, lead to greater fertilization success.

Cut your losses: self-amputation of injured limbs increases survival.

Autotomy, self-induced limb loss, is an extreme trait observed throughout the animal kingdom; lizards drop their tails, crickets release their legs, and crabs drop their claws. These repeated evolutionary origins suggest that autotomy is adaptive. Yet, we do not have a firm understanding of the selective pressures that promote and maintain this extreme trait. Although multiple adaptive hypotheses exist, research has generally focused on autotomy's adaptive value as a form of predator escape. However, autotomy could also be selected to reduce the cost of an injured limb, which we investigate here. Previously, this alternative hypothesis has been challenging to directly test because when an injury occurs on an autotomizable limb, that limb is almost always dropped (i.e., autotomy is behaviorally fixed within populations). Recently, however, we have identified a species, Narnia femorata (Insecta: Hemiptera: Coreidae), where some individuals autotomize limbs in response to injury, but some do not. This natural variation allowed us to investigate both the survival costs of retaining an injured limb and the benefits of autotomizing it. In this study, we find a positive association between autotomizing injured limbs and survival, thereby quantifying a new and likely widespread benefit of autotomy-reducing the cost of injury.

Broadening the voice of science: Promoting scientific communication in the undergraduate classroom.

Effective and accurate communication of scientific findings is essential. Unfortunately, scientists are not always well trained in how to best communicate their results with other scientists nor do all appreciate the importance of speaking with the public. Here, we provide an example of how the development of oral communication skills can be integrated with research experiences at the undergraduate level. We describe our experiences developing, running, and evaluating a course for undergraduates that complemented their existing undergraduate research experiences with instruction on the nature of science and intensive training on the development of science communication skills. Students delivered science talks, research monologues, and poster presentations about the ecological and evolutionary research in which they were involved. We evaluated the effectiveness of our approach using the CURE survey and a focus group. As expected, undergraduates reported strong benefits to communication skills and confidence. We provide guidance for college researchers, instructors, and administrators interested in motivating and equipping the next generation of scientists to be excellent science communicators.