The function of prolonged copulations in Enchenopa treehoppers (Hemiptera: Membracidae).

Copulations are very brief in many species, sometimes taking only seconds, but in other species, they can be quite prolonged. Potential explanations for prolonged copulations include time requirements for the transfer of sperm and/or other ejaculate substances. Ejaculate substances could function to regulate female receptivity to subsequent matings, provide nutritional nuptial gifts, or hasten egg oviposition at a potential survival cost to the female. We investigated prolonged copulation in a member of the Enchenopa binotata complex of treehoppers (Hemiptera: Membracidae), in which females rarely remate and copulation can last several hours. We assigned females to treatments in which we interrupted copulation at different times. We also included a control where copulation was not interrupted. We found that females that experienced shorter copulations were more likely to be subsequently receptive to an attractive male. We also found that few females produced offspring when they engaged in short copulations compared to those with longer copulations. We did not find any differences in female survival. Our results support the sperm transfer and receptivity regulation hypotheses. We discuss potential reasons for why these processes should take so long in a species with low female remating.

On the function of a female-like signal type in the vibrational repertoire of Enchenopa male treehoppers (Hemiptera: Membracidae).

Animals often mimic the behaviours or signals of conspecifics of the opposite sex while courting. We explored the potential functions of a novel female-like signal type in the courtship displays of male Enchenopa treehoppers. In these plant-feeding insects, males produce plant-borne vibrational advertisement signals, to which females respond with their own duetting signals. Males also produce a signal type that resembles the female duetting responses. We experimentally tested whether this signal modifies the behaviour of receivers. First, we tested whether the female-like signal would increase the likelihood of a female response. However, females were as likely to respond to playbacks with or without them. Second, we tested whether the female-like signal would inhibit competing males, but males were as likely to produce displays after playbacks with or without them. Hence, we found no evidence that this signal has an adaptive function, despite its presence in the courtship display, where sexual selection affects signal features. Given these findings, we also explored whether the behavioural and morphological factors of the males were associated with the production of the female-like signal. Males that produced this signal had higher signalling effort (longer and more frequent signals) than males that did not produce it, despite being in worse body condition. Lastly, most males were consistent over time in producing the female-like signal or not. These findings suggest that condition-dependent or motivational factors explain the presence of the female-like signal. Alternatively, this signal might not bear an adaptive function, and it could be a way for males to warm up or practice signalling, or even be a by-product of how signals are transmitted through the plant. We suggest further work that might explain our puzzling finding that a signal in the reproductive context might not have an adaptive function.

Social Plasticity Enhances Signal-Preference Codivergence.

AbstractThe social environment is often the most dynamic and fitness-relevant environment animals experience. Here we tested whether plasticity arising from variation in social environments can promote signal-preference divergence-a key prediction of recent speciation theory but one that has proven difficult to test in natural systems. Interactions in mixed social aggregations could reduce, create, or enhance signal-preference differences. In the latter case, social plasticity could establish or increase assortative mating. We tested this by rearing two recently diverged species of Enchenopa treehoppers-sap-feeding insects that communicate with plant-borne vibrational signals-in treatments consisting of mixed-species versus own-species aggregations. Social experience with heterospecifics (in the mixed-species treatment) resulted in enhanced signal-preference species differences. For one of the two species, we tested but found no differences in the plastic response between sympatric and allopatric sites, suggesting the absence of reinforcement in the signals and preferences and their plastic response. Our results support the hypothesis that social plasticity can create or enhance signal-preference differences and that this might occur in the absence of long-term selection against hybridization on plastic responses themselves. Such social plasticity may facilitate rapid bursts of diversification.

Seasonal shift in diet affects female reproductive anatomy but not mating behavior.

Females experience considerable environmental variability when breeding seasons are long. Adverse nutritional conditions can result in a reduction in mating and reproduction. However, a return to good nutrition may help animals resume high reproductive investment. I tested the silver spoon hypothesis in which females raised under poor conditions are reproductively limited compared to those raised under good conditions regardless of their adult environment. I used a specialist herbivore, Narnia femorata (Hemiptera: Coreidae), that lives on seasonally changing cacti. I provided juveniles and adults with a cactus pad with fruit (good diet), without fruit (restricted diet), or an improved adult diet (no fruit as juveniles, fruit at adulthood) to simulate a seasonal change in their diets near the end of the breeding season. I found that both ovary size and egg presence were reduced for females fed the restricted diet compared to those fed the good diet. Females fed the improved diet grew large ovaries like those fed the good diet, but few produced any eggs. Interestingly, female mating behavior did not change but females were less attractive to males when fed restricted diets. My results support the silver spoon hypothesis for compensatory growth and suggest that tradeoffs may occur between early survival and future reproduction when females experience a poor early life diet.

Extreme variation in testes size in an insect is linked to recent mating activity.

Ample sperm production is essential for successful male reproduction in many species. The amount of sperm a male can produce is typically constrained by the size of his testes, which can be energetically expensive to grow and maintain. Although the economics of ejaculate allocation has been the focus of much theoretical and empirical literature, relatively little attention has been paid to individual adult variation and plasticity at the source of sperm production, the testes themselves. We experimentally address this issue using the insect Narnia femorata Stål (Hemiptera: Coreidae). We established the metabolic cost of testicular tissue and then quantified variation in individual testes mass in response to multiple mate quality and quantity treatments. We uncovered extreme variation across individuals and considerable short-term effects of mating activity on testes dry mass. Importantly, the observed variation in testes mass was associated with notable fitness consequences; females paired with males with larger testes had greater hatching success. Overall, pairing with a female resulted in a 11% reduction in dry testes mass. Despite this apparent considerable mating investment, we found no evidence of strategic allocation to higher quality females or longer-term changes in testes mass. The dynamic nature of testes mass and its metabolic cost is vital to consider in the context of re-mating rates, polyandry benefits and general mating system dynamics both in this species and more broadly.

The leaf-footed cactus bug is not a cactus specialist: Narnia femorata feeds, fights, and mates on thistle.

Novel host plants are incorporated into the diets of phytophagous insects when females oviposit and juveniles feed and survive on them. A change in diet, however, can have morphological consequences. We recently found a population of the leaf-footed cactus bug, Narnia femorata (Hemiptera: Coreidae), a historical cactus specialist, living and feeding on Cirsium thistle. We also found adults breeding and males using their enlarged hind legs (i.e., weapons) in male-male combat on thistle. When we compared this thistle population with a population feeding on cactus, we found that both populations had similar body and weapon sizes as well as weapon composition. However, the population living on thistle had longer mouthparts than the population found on cactus, although this difference only occurred at larger body sizes. This difference in adult mouthpart size is likely a result of the juvenile rearing environment (i.e., thistle or cactus). However, genetic differences may also affect this trait. Our results provide some interesting avenues for future research (e.g., a reciprocal transplant experiment) in a species with a recent host range expansion.

High-quality host plant diets partially rescue female fecundity from a poor early start.

Nutrition is a dynamic environmental factor and compensatory growth may help animals handle seasonal fluctuations in their diets. Yet, how the dynamic changes in nutrition affect female reproduction is understudied. We took advantage of a specialist insect herbivore, Narnia femorata Stål (Hemiptera: Coreidae), that feeds and reproduces on cactus across three seasons. We first examined how cactus quality can affect female reproductive success. Then, we investigated the extent to which reproductive success can be improved by a switch in diet quality at adulthood. We placed N. femorata juveniles onto prickly pear cactus pads with early-season (low-quality) or late-season (high-quality) fruit and tracked survivorship and development time. A subset of the females raised on low-quality diets were provided with an improved adult diet to simulate a seasonal change in diet. Adult female survival and egg production were tracked over time. All fitness-related traits were lower for females fed low-quality diets compared with females fed high-quality diets. However, when females had access to an improved adult diet, egg production was partially rescued. These findings show that a seasonal improvement in diet can enhance reproduction, but juvenile nutrition still has lasting effects that females cannot overcome.

Long-term reproductive success is predicted by sexual behavior and impaired by temporary nutritional stress during sexual maturation.

Nutritional stress during the earliest stages of an animal's life can have long-term effects on its behavior and reproductive performance, but the effects of brief periods of nutritional stress later in life are less well-studied. We manipulated female diet in Narnia femorata (Hemiptera: Coreidae) and investigated to what extent nutritional stress during sexual maturation affects subsequent sexual behavior and long-term offspring production. We show that nutritional stress at this key point during early adulthood can have lasting effects on reproduction, impairing long-term offspring production despite the subsequent return of good nutrition. These results demonstrate that nutritional availability during late stages of development, in young adults, can be crucial to future fitness. We found no effect of temporary nutritional stress on female receptivity to mating or attractiveness to males; although females that were less receptive also produced fewer offspring in the next month. Overall, we demonstrate that even brief periods of nutritional deprivation late in development can have drastic long-term effects, apparently beyond compensation, and despite a good early nutritional environment.

Seasonal Effects on the Population, Morphology and Reproductive Behavior of Narnia femorata (Hemiptera: Coreidae).

Many insects are influenced by the phenology of their host plants. In North Central Florida, Narnia femorata (Hemiptera: Coreidae) spends its entire life cycle living and feeding on Opuntia mesacantha ssp. lata. This cactus begins producing flower buds in April that lead to unripe green fruit in June that ripen into red fruit through December. Many morphological and behavioral characteristics of N. femorata are known to be affected by cactus phenology in a controlled laboratory setting, including the degree of sexual dimorphism and mating behavior. Our goal with this study was to determine if similar phenotypic changes of N. femorata occurred over time in the wild, and the extent to which these changes were concordant with phenological changes in its host plant. Further, we investigate the length of the insect mouthparts (beak) over time. Ongoing work has suggested that beak length may change across cohorts of developing insects in response to feeding deep within cactus fruit where seed and pulp depth decrease as the fruit ripens. Our results revealed a drop in cactus fruit abundance between the months of July through October 2015 as cactus fruits turned red and ripened. Simultaneously, the average body size of both males and females of N. femorata declined at two sampled sites. Male hind femora (a sexually-selected weapon) decreased disproportionately in size over time so that males later in the year had relatively smaller hind femora for their body size. The sex-specific patterns of morphological change led to increased sexual-size dimorphism and decreased sexual dimorphism for hind femora later in the year. Further, we found that beak length decreased across cohorts of insects as cactus fruit ripened, suggesting phenotypic plasticity in mouthpart length. Behavioral studies revealed that female readiness to mate increased as the season progressed. In sum, we found pronounced changes in the phenotypes of these insects in the field. Although this study is far from comprehensive, it provides tantalizing patterns that suggest many directions for future research.

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.