Teaching animal behavior online: A primer for the pandemic and beyond.

Behavior courses face numerous challenges when moving to an online environment, as has been made necessary by the COVID-19 pandemic. These challenges occur largely because behavior courses, like most organismal biology courses, often stress experiential learning through laboratories that involve live animals, as well as a lecture component that emphasizes formative assessment, discussion, and critical thinking. Although online behavior courses may be remote, they can still be interactive and social, and designed with inclusive pedagogy. Here, we discuss some of the key decisions that instructors should consider, provide recommendations, and point out new opportunities for student learning that stem directly from the move to online instruction. Specific topics include challenges related to generating an inclusive and engaging online learning environment, synchronous versus asynchronous formats, assignments that enhance student learning, testing format and execution, grade schemes, design of laboratory experiences including opportunities for community science, design of synthetic student projects, and workload balance for students and instructors. We designed this primer both for animal behavior instructors who need to quickly transition to online teaching in the midst of a pandemic, and for those facing such transitions in upcoming terms. Much of the manuscript's content should also be of general interest and value to instructors from all areas of organismal biology who are attempting to quickly transition to online teaching.

Do exaggerated chelicerae function as weapons or genitalia in a long-jawed spider? Functional allometric analysis yields an answer.

From the elongated neck of the giraffe to the elaborate train of the peacock, extreme traits can result from natural or sexual selection (or both). The extreme chelicerae of the long-jawed spiders (Tetragnatha) present a puzzle: do these exaggerated chelicerae function as weapons or genitalia? Bristowe first proposed that Tetragnatha chelicerae function as a holdfast because these spiders embrace chelicerae during mating. This hypothesis has remained untested until now. Here, we use functional allometry to examine how extreme chelicerae develop and perform in the long-jawed spider Tetragnatha elongata. Similar to other Tetragnatha species, chelicerae were longer in adult males than in adult females. Overall, we confirm Bristowe's hypothesis: elongation only occurred in the adult stage. However, we propose that chelicerae function as more than a holdfast in T. elongata. Male chelicerae exhibited positive allometry, which suggests scaling as weapons rather than genitalia. However, fieldwork revealed that the operational sex ratio is female-biased and both adult male-male competition and sexual cannibalism were rarely observed. Consequently, we propose that the positive allometry of male chelicerae may result from sexual selection to mechanically mesh with larger and more fecund females. Evidence for mechanical mesh includes multiple traits ranging from apophyses and grooves to guide teeth on the basal portion of the chelicerae. In contrast, we propose that chelicerae of females are analogous to the female peacock's tail: shortened by natural selection limiting the exaggeration of sexually selected traits. Indeed, females had increased foraging efficiency compared to males and exhibited negative cheliceral allometry. We discuss the implications for the evolution of elongated chelicerae in Tetragnatha.

Spiders in motion: demonstrating adaptation, structure-function relationships, and trade-offs in invertebrates.

Evolutionary history and structural considerations constrain all aspects of animal physiology. Constraints on invertebrate locomotion are especially straightforward for students to observe and understand. In this exercise, students use spiders to investigate the concepts of adaptation, structure-function relationships, and trade-offs. Students measure burst and endurance performance in several taxonomic families of spiders whose ecological niches have led to different locomotory adaptations. Based on observations of spiders in their natural habitat and prior background information, students make predictions about spider performance. Students then construct their own knowledge by performing a hands-on, inquiry-based scientific experiment where the results are not necessarily known. Depending on the specific families chosen, students can observe that web-dwelling spiders have more difficulty navigating complex terrestrial terrain than ground-dwelling spiders and that there is a trade-off between burst performance and endurance performance in spiders. Our inexpensive runway design allows for countless variations on this basic experiment; for example, we have successfully used runways to show students how the performance of heterothermic ectotherms varies with temperature. High levels of intra- and interindividual variation in performance underscore the importance of using multiple trials and statistical tests. Finally, this laboratory activity can be completely student driven or standardized, depending on the instructor's preference.

Indirect genetic effects and the lek paradox: inter-genotypic competition may strengthen genotype x environment interactions and conserve genetic variance.

Understanding the evolutionary mechanisms that maintain genetic variation in natural populations is one of the fundamental goals of evolutionary biology. There is growing evidence that genotype-by-environment interaction (G x E) can maintain additive genetic variance (V (A)), but we lack information on the relative performance of genotypes under the competitive situations encountered in the field. Competing genotypes may influence each other, and this interaction is also subject to selection through indirect genetic effects (IGE). Here, we explore how genotypes perform when interacting and evaluate IGE in order to understand its influence on V (A) for sexually-selected traits in the lesser waxmoth, Achroia grisella. We found that inter-genotype differences and crossover interactions under joint rearing are equal to or greater than values when reared separately. A focal genotype exhibited different performances when jointly reared with various genotypes-suggesting that IGE may be responsible for the increased levels of crossover and differences in performance observed. We suggest that some genotypes are superior competitors for food acquisition in the larval stage, and that these differences influence the development and evolution of other genotypes through IGE. We reaffirm the role of G x E in maintaining V (A) and note the general importance of IGE in studies of evolutionary mechanisms.