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1. Sexual selection does not drive hindwing tail elaboration in a moon moth, Actias luna

   https://doi.org/10.1093/beheco/arad019  

   Rubin, Juliette J; Kawahara, Akito Y (April 2023, Behavioral Ecology)

   Jennions, MIchael D (Ed.)

   Abstract The most emblematic animal traits are often attributed to sexual selection. While this pressure is an important force, elaborated traits that have been driven solely by natural selection are less enumerated. Here, we test an elaborate trait in moths—hindwing tails—that has been studied in an anti-predator context, but that remains unstudied for its role in mating. We gave female Actias luna (Saturniidae) moths a choice between two males of differing hindwing tail treatments. In our primary experiment, males with intact tails garnered more matings than males with tails removed. This difference appears to result from damage incurred by tail removal, however, as demonstrated with additional experiments. We created a tail/no-tail experimental set where we removed tails from both males, then reglued tails to one and applied glue only to the hindwings of the other. We found no significant difference in mating success between these males. To ensure that this result was not due to the glue itself, we offered females two intact males, with glue added to the wings of one. This set also had equal mating success. We therefore do not find evidence that tails play a role in sexual selection. These results, in combination with previous research on bat-moth battles using A. luna, indicate that the non-sexually dimorphic hindwing tail was likely driven by natural selection. We suggest that future research testing multiple selective forces is needed to reveal the prevalence of natural versus sexual selection as the primary force driving trait elaboration in diverse animal taxa. 

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2. Testing bird-driven diurnal trade-offs of the moon moth's anti-bat tail

   https://doi.org/10.1098/rsbl.2022.0428  

   Rubin, Juliette J.; Martin, Nich W.; Sieving, Kathryn E.; Kawahara, Akito Y. (February 2023, Biology Letters)

   Traits are often caught in a dynamic tension of countervailing evolutionary pressures. Trade-offs can be imposed by predators evolutionarily curtailing the conspicuousness of a sexually selected trait, or acting in opposition to another natural selection pressure, for instance, a different predator with a divergent hunting strategy. Some moon moths (Saturniidae) have long hindwing tails that thwart echolocating bat attacks at night, allowing the moth to escape. These long tails may come at a cost, however, if they make the moth's roosting form more conspicuous to visually foraging predators during the day. To test this potential trade-off, we offered wild-caught Carolina wrens ( Thryothorus ludovicianus ) pastry dough models with real Actias luna wings that were either intact or had tails experimentally removed. We video recorded wrens foraging on models and found that moth models with tails did not experience increased detection and attack by birds. Thus, this elaborate trait, while obvious to human observers, does not seem to come at a cost of increased avian predator attention. The evolution of long hindwing tails, likely driven by echolocating predators at night, does not seem to be limited by opposing diurnal constraints. This study demonstrates the importance of testing presumed trade-offs and provides hypotheses for future testing. 

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3. Keystone predation: trait‐based or driven by extrinsic processes? Assessment using a comparative‐experimental approach

   https://doi.org/10.1002/ecm.1436  

   Menge, Bruce A.; Foley, Melissa M.; Robart, Matthew J.; Richmond, Erin; Noble, Mae; Chan, Francis (December 2020, Ecological Monographs)

   Abstract Keystone predation can be a determinant of community structure, including species diversity, but factors underlying “keystoneness” have been minimally explored. Using the system in which the original keystone, the sea starPisaster ochraceus, was discovered, we focused on two potential (but overlapping) determinants of keystoneness: intrinsic traits or state variables of the species (e.g., size, density), and extrinsic environmental parameters (e.g., prey productivity) that may provide conditions favorable for keystone predator evolution. Using a comparative‐experimental approach, with repeated field experiments at multiple sites across a variable coastal environment, we tested predation rates, or how quickly predators consumed prey, and predation effects, or community response to predator presence or absence. We tested five hypotheses: (H₁) predation rates and effects will vary in space but not time; (H₂) per population predation rates will vary primarily with individual traits and population variables; (HJHH₃) per capita predation rates will vary only with individual traits; (H₄) predation effects will vary with traits, variables, and external drivers; and (H₅) as predicted by the keystone predation hypothesis, diversity will vary unimodally with predation pressure. As hypothesized, predation rates differed among sites but not over time (H₁), and in caging exclusion experiments, predation effect varied with both intrinsic and extrinsic factors (H₄). Unexpectedly, predation rates varied with both intrinsic and extrinsic (H₂, per population), or only with extrinsic (H₃, per capita) factors. Further, in large‐plot exclusion experiments, predation effect was most closely associated with individual traits (contraH₄). Finally, taxon diversity varied unimodally with proxies of predation pressure (sessile prey abundance) and was sensitive to extrinsic factors (mussel growth, temperature, and upwelling,H₅). Hence, keystoneness depended on predator individual traits, predator population variables, and environmental parameters. However, temporal differences in caging experiments suggested that environmental characteristics underlying prey dynamics may be preeminent. Compared to prior experiments, predation was weaker with low prey input compared to periods with high prey input. Collectively, our results suggest that keystone predator evolution depends on the coalescence of species‐specific characteristics, and environmental parameters favoring high prey productivity. Our approach may be a model for future studies exploring the generality of keystoneness. 

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4. Tiger beetles produce anti-bat ultrasound and are probable Batesian moth mimics

   https://doi.org/10.1098/rsbl.2023.0610  

   Gough, Harlan M; Rubin, Juliette J; Kawahara, Akito Y; Barber, Jesse R (May 2024, Biology Letters)

   Echolocating bats and their eared insect prey are in an acoustic evolutionary war. Moths produce anti-bat sounds that startle bat predators, signal noxiousness, mimic unpalatable models and jam bat sonar. Tiger beetles (Cicindelidae) also purportedly produce ultrasound in response to bat attacks. Here we tested 19 tiger beetle species from seven genera and showed that they produce anti-bat signals to playback of authentic bat echolocation. The dominant frequency of beetle sounds substantially overlaps the sonar calls of sympatric bats. As tiger beetles are known to produce defensive chemicals such as benzaldehyde and hydrogen cyanide, we hypothesized that tiger beetle sounds are acoustically advertising their unpalatability. We presented captive big brown bats (Eptesicus fuscus) with seven different tiger beetle species and found that 90 out of 94 beetles were completely consumed, indicating that these tiger beetle species are not aposematically signalling. Instead, we show that the primary temporal and spectral characteristics of beetle warning sounds overlap with sympatric unpalatable tiger moth (Arctinae) sounds and that tiger beetles are probably Batesian mimics of noxious moth models. We predict that many insect taxa produce anti-bat sounds and that the acoustic mimicry rings of the night sky are hyperdiverse. 

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5. Phylogenomics reveals the evolutionary timing and pattern of butterflies and moths

   https://doi.org/10.1073/pnas.1907847116  

   Kawahara, Akito Y.; Plotkin, David; Espeland, Marianne; Meusemann, Karen; Toussaint, Emmanuel F.; Donath, Alexander; Gimnich, France; Frandsen, Paul B.; Zwick, Andreas; dos Reis, Mario; et al (November 2019, Proceedings of the National Academy of Sciences)

   Butterflies and moths (Lepidoptera) are one of the major superradiations of insects, comprising nearly 160,000 described extant species. As herbivores, pollinators, and prey, Lepidoptera play a fundamental role in almost every terrestrial ecosystem. Lepidoptera are also indicators of environmental change and serve as models for research on mimicry and genetics. They have been central to the development of coevolutionary hypotheses, such as butterflies with flowering plants and moths’ evolutionary arms race with echolocating bats. However, these hypotheses have not been rigorously tested, because a robust lepidopteran phylogeny and timing of evolutionary novelties are lacking. To address these issues, we inferred a comprehensive phylogeny of Lepidoptera, using the largest dataset assembled for the order (2,098 orthologous protein-coding genes from transcriptomes of 186 species, representing nearly all superfamilies), and dated it with carefully evaluated synapomorphy-based fossils. The oldest members of the Lepidoptera crown group appeared in the Late Carboniferous (∼300 Ma) and fed on nonvascular land plants. Lepidoptera evolved the tube-like proboscis in the Middle Triassic (∼241 Ma), which allowed them to acquire nectar from flowering plants. This morphological innovation, along with other traits, likely promoted the extraordinary diversification of superfamily-level lepidopteran crown groups. The ancestor of butterflies was likely nocturnal, and our results indicate that butterflies became day-flying in the Late Cretaceous (∼98 Ma). Moth hearing organs arose multiple times before the evolutionary arms race between moths and bats, perhaps initially detecting a wide range of sound frequencies before being co-opted to specifically detect bat sonar. Our study provides an essential framework for future comparative studies on butterfly and moth evolution. 

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