Global biogeography of warning coloration in the butterfly Danaus chrysippus

Warning coloration provides a textbook example of natural selection, but the frequent observation of polymorphism in aposematic species presents an evolutionary puzzle. We investigated biogeography and polymorphism of warning patterns in the widespread butterfly Danaus chrysippus using records from citizen science (n = 5467), museums (n = 8864) and fieldwork (n = 2586). We find that polymorphism in three traits controlled by known mendelian loci is extensive. Broad allele frequency clines, hundreds of kilometres wide, suggest a balance between long-range dispersal and predation of unfamiliar morphs. Mismatched clines for the white hindwing and forewing tip in East Africa are consistent with a previous finding that the black wingtip allele has spread recently in the region through hitchhiking with a heritable endosymbiont. Light/dark background coloration shows more extensive polymorphism. The darker genotype is more common in cooler regions, possibly reflecting a trade-off between thermoregulation and predator warning. Overall, our findings show how studying local adaptation at the global scale provides a more complete picture of the evolutionary forces involved

Arthropod associates of Kenyan buffelgrass (Cenchrus ciliaris): a field survey for biological control candidates of a globally important invasive grass

Invasive species are the second largest contributor to biodiversity loss and drivers of ecosystem change. Buffelgrass is a C4, perennial grass native to Africa and Asia that was widely introduced across tropical and subtropical rangelands as livestock forage. Buffelgrass reduces native biodiversity and ecosystem stability in its introduced range when it escapes livestock pastures. Understanding the ecology of insects associated with buffelgrass in its native range may provide an understanding of invasion processes and biological control opportunities where buffelgrass has become an invasive challenge. Here, we present the results of a six-year survey of herbivorous arthropods of a native buffelgrass population from Kenya. Buffelgrass was examined for externally and internally feeding insects of vegetative and reproductive tissues. We also categorised buffelgrass detritivores and parasitoids that may use buffelgrass herbivores as hosts. The samples were photographed and Sanger sequenced to identify them to the lowest possible taxonomic ranking. We collected information on sample abundances, phenologies, tissues consumed, and putative diet breadths. We identified 25 morphospecies representing seven orders and 16 families. The putative host plant specialisation was as high as 67% for Diptera. Phenological variation in herbivore presence correlated with seasonal rainfall and provided a guide for when to conduct follow-up biological control agent searches. The most abundant herbivore was a gall midge (Diptera: Cecidomyiidae) that inhabits buffelgrass culms. Additional research should focus on bringing this species into containment where host choice trials can be conducted to determine if it is truly monophagous and assess its impact on buffelgrass growth

Systematic reduction of natural enemies and competition across variable precipitation approximates buffelgrass invasiveness ( Cenchrus ciliaris ) in its native range

Invasive grasses cause devastating losses to biodiversity and ecosystem function directly and indirectly by altering ecosystem processes. Escape from natural enemies, plant–plant competition, and variable resource availability provide frameworks for understanding invasion. However, we lack a clear understanding of how natural stressors interact in their native range to regulate invasiveness. In this study, we reduced diverse guilds of natural enemies and plant competitors of the highly invasive buffelgrass across a precipitation gradient throughout major climatic shifts in Laikipia, Kenya. To do this, we used a long‐term ungulate exclosure experiment design across a precipitation gradient with nested treatments that (1) reduced plant competition through clipping, (2) reduced insects through systemic insecticide, and (3) reduced fungal associates through fungicide application. Additionally, we measured the interaction of ungulates on two stem‐boring insect species feeding on buffelgrass. Finally, we measured a multiyear smut fungus outbreak. Our findings suggest that buffelgrass exhibits invasive qualities when released from a diverse group of natural stressors in its native range. We show natural enemies interact with precipitation to alter buffelgrass productivity patterns. In addition, interspecific plant competition decreased the basal area of buffelgrass, suggesting that biotic resistance mediates buffelgrass dominance in the home range. Surprisingly, systemic insecticides and fungicides did not impact buffelgrass production or reproduction, perhaps because other guilds filled the niche space in these highly diverse systems. For example, in the absence of ungulates, we showed an increase in host‐specific stem‐galling insects, where these insects compensated for reduced ungulate use. Finally, we documented a smut outbreak in 2020 and 2021, corresponding to highly variable precipitation patterns caused by a shifting Indian Ocean Dipole. In conclusion, we observed how reducing natural enemies and competitors and certain interactions increased properties related to buffelgrass invasiveness

The Afrotropical breeding grounds of the Palearctic-African migratory painted lady butterflies (Vanessa cardui)

Migratory insects are key players in ecosystem functioning and services, but their spatiotemporal distributions are typically poorly known. Ecological niche modeling (ENM) may be used to predict species seasonal distributions, but the resulting hypotheses should eventually be validated by field data. The painted lady butterfly (Vanessa cardui) performs multigenerational migrations between Europe and Africa and has become a model species for insect movement ecology. While the annual migration cycle of this species is well understood for Europe and northernmost Africa, it is still unknown where most individuals spend the winter. Through ENM, we previously predicted suitable breeding grounds in the subhumid regions near the tropics between November and February. In this work, we assess the suitability of these predictions through i) extensive field surveys and ii) two-year monitoring in six countries: a large-scale monitoring scheme to study butterfly migration in Africa. We document new breeding locations, year-round phenological information, and hostplant use. Field observations were nearly always predicted with high probability by the previous ENM, and monitoring demonstrated the influence of the precipitation seasonality regime on migratory phenology. Using the updated dataset, we built a refined ENM for the Palearctic-African range of V. cardui. We confirm the relevance of the Afrotropical region and document the missing natural history pieces of the longest migratory cycle described in butterflies.This work was funded by the National Geographic Society (grant WW1-300R-18); by the British Ecological Society (grant LRB16/1015); by the Research and Conservation Projects of the Fundació Barcelona Zoo; by the grant PID2020-117739GA-I00/MCIN/AEI/10.13039/501100011033 of the Spanish Ministry of Science and Innovation and the Spanish State Research Agency to G.T.; by the grant LINKA20399 from the Spanish National Research Council iLink program to G.T., C.P.B., N.E.P., and R.V.; by fellowship FPU19/01593 of the program Formación de Profesorado Universitario (FPU) to A.G.-B.; by the Turkana Basin Institute, National Geographic Society, and Whitley Fund for Nature to D.J.M.; and by grant 2018-00738 of the New Frontiers in Research Fund (Government of Canada) to G.T. and C.P.B.Significance Abstract Results Field Surveys, Larval Hostplants, and Field-Based Model Validation Monitoring Results and Population Dynamics across Regions A Refined Model for the Afrotropical Region Discussion The Afrotropical Breeding Grounds of V. cardui: Multiple Generations Shift South Toward the Tropics Diversity and Phenology of Larval Hostplants in the Afrotropics The Ecological Relevance of Delimiting Spatiotemporal Distributions in Migratory Insects Conclusion Methods December-January Field Surveys and Year-Round Monitoring Spatiotemporal Ecological Niche Modeling Data, Materials, and Software Availability Acknowledgments Supporting Information Reference

Automated detection of an insect‐induced keystone vegetation phenotype using airborne LiDAR

Ecologists, foresters and conservation practitioners need ‘biodiversity scanners’ to effectively inventory biodiversity, audit conservation progress and track changes in ecosystem function. Quantifying biological diversity using remote sensing methods remains challenging, especially for small invertebrates. However, insect aggregations can drastically alter landscapes and vegetation, and these ‘extended phenotypes’ could serve as environmental landmarks of insect presence in remotely sensed data. To test the feasibility of this approach, we studied symbiotic ants that alter the canopy shape of whistling thorn acacias (Acacia [syn. Vachellia] drepanolobium), a keystone tree species of the black cotton soils of east African savannas. We demonstrate a protocol for using light detection and ranging (LiDAR) data to collect, prepare (including a customizable tree‐segmentation algorithm) and apply a convolutional neural network‐based classification for the detection of ant‐inhabited acacia tree phenotypic variations. Applying this protocol enabled us to effectively detect intra‐specific tree phenotypic variation induced by insects. Surveying ant occupancy across 16 ha and 9680 acacia trees took 1000 work hours, whereas surveyed patterns of ant distribution were replicated by our trained classifier using only an hour‐long airborne LiDAR collection time. We suggest that large‐scale surveys of insect occupancy (including insect‐vectored disease) can be automated through a combination of airborne LiDAR and machine learning

The invasiveness of Guinea grass (Megathyrsus maximus) is characterized by habitat and differing herbivore assemblages in its native and invaded range

A significant challenge of global change is the human-mediated movement of pasture grasses and their subsequent impact on ecosystem processes when they become invasive. We must understand invasive grass ecology and their natural enemies in native and introduced ranges to mitigate these impacts. Guinea grass (Megathyrsus maximus) is a pantropically introduced pasture grass that escapes intended areas and invades native ecosystems – threatening biodiversity and ecosystem function. The success of invasive plants has often been attributed to ecological release from stressors, including natural enemies and resource availability. Our objective was to assess Guinea grass functional traits across three different habitat types in native and invaded ranges by documenting ungulate and arthropod abundance, diversity, and feeding guilds. Guinea grass functional traits were assessed in three habitat types: grassland, riparian, and woody thickets around nitrogen-fixing Prosopis glandulosa in its introduced range in Texas, USA, and Senegalia mellifera in its native range in Kenya. We characterized Guinea grass functional traits by measuring plant height, cover, biomass, root-to-shoot ratios, and reproductive traits. We then examined the phytophagous arthropod and ungulate abundance and feeding guild diversity across the three habitat types. We hypothesized that functional trait expression related to invasiveness would be associated with Guinea grass in its introduced range. Also, we hypothesized that the abundance and diversity of phytophagous arthropods and ungulates would be lower in the invaded range. Finally, we hypothesized that Guinea grass functional traits would differ between the three habitat types, given the habitat types’ innate differences in resource availability. We found that Guinea grass was 2.5 times taller and 3.3 times more productive and covered 2.5 times more area in its invaded versus native ranges. Introduced Guinea grass had higher reproduction rates with 2.5 times more reproductive tillers, while habitat type drove vegetative reproduction with 15 times more stoloniferous establishment in wooded and riparian sites than grasslands. Texan ungulate communities were less species-rich, less functionally diverse, and less abundant than the Kenyan ungulate community. The phytophagous arthropod diversity on plants was twice as high on Kenyan Guinea grass than on Texan Guinea grass. Total arthropod family richness was nearly double, with 15 families represented in Kenya and 8 in Texas. These results suggest that Guinea grass has escaped a rich assemblage of arthropods and ungulates and likely explains some of its spread in introduced ranges. This study demonstrates how the invasive success of Guinea grass can be understood in terms of its competitive ability and interaction with natural enemies in the introduced and native ranges and may inform future biological control

Transposable element insertions are associated with Batesian mimicry in the pantropical butterfly Hypolimnas misippus.

Hypolimnas misippus is a Batesian mimic of the toxic African Queen butterfly (Danaus chrysippus). Female H. misippus butterflies use two major wing patterning loci (M and A) to imitate three colour morphs of D. chrysippus found in different regions of Africa. In this study, we examine the evolution of the M locus and identify it as an example of adaptive atavism. This phenomenon involves a morphological reversion to an ancestral character that results in an adaptive phenotype. We show that H. misippus has re-evolved an ancestral wing pattern present in other Hypolimnas species, repurposing it for Batesian mimicry of a D. chrysippus morph. Using haplotagging, a linked-read sequencing technology, and our new analytical tool, Wrath, we discover two large transposable element (TE) insertions located at the M locus and establish that these insertions are present in the dominant allele responsible for producing mimetic phenotype. By conducting a comparative analysis involving additional Hypolimnas species, we demonstrate that the dominant allele is derived. This suggests that, in the derived allele, the TEs disrupt a cis-regulatory element, leading to the reversion to an ancestral phenotype that is then utilized for Batesian mimicry of a distinct model, a different morph of D. chrysippus. Our findings present a compelling instance of convergent evolution and adaptive atavism, in which the same pattern element has independently evolved multiple times in Hypolimnas butterflies, repeatedly playing a role in Batesian mimicry of diverse model species

Transposable Element Insertions Are Associated with Batesian Mimicry in the Pantropical Butterfly Hypolimnas misippus.

Acknowledgements: This project would have not been possible without the help of Eunice Katili and Charo Ngumbao at Kipepeo Project in Malindi, Kenya, who were crucial in collecting samples. A.O. thanks Gergely Flamich for the numerous discussions about Wrath. A.O. thanks the Cambridge Trust and St. John's College for their support.Funder: Varley-Gradwell Travelling Fellowship in Insect Ecology 2021Funder: American Society of Naturalists Student Research AwardFunder: Lepidopterists’ Society Ron Leuschner Memorial Fund for ResearchFunder: Cambridge Trust; DOI: https://doi.org/10.13039/501100003343Funder: European Research ScholarshipFunder: St. John's College; DOI: https://doi.org/10.13039/100010937Funder: Benefactors’ ScholarshipHypolimnas misippus is a Batesian mimic of the toxic African Queen butterfly (Danaus chrysippus). Female H. misippus butterflies use two major wing patterning loci (M and A) to imitate three color morphs of D. chrysippus found in different regions of Africa. In this study, we examine the evolution of the M locus and identify it as an example of adaptive atavism. This phenomenon involves a morphological reversion to an ancestral character that results in an adaptive phenotype. We show that H. misippus has re-evolved an ancestral wing pattern present in other Hypolimnas species, repurposing it for Batesian mimicry of a D. chrysippus morph. Using haplotagging, a linked-read sequencing technology, and our new analytical tool, Wrath, we discover two large transposable element insertions located at the M locus and establish that these insertions are present in the dominant allele responsible for producing mimetic phenotype. By conducting a comparative analysis involving additional Hypolimnas species, we demonstrate that the dominant allele is derived. This suggests that, in the derived allele, the transposable elements disrupt a cis-regulatory element, leading to the reversion to an ancestral phenotype that is then utilized for Batesian mimicry of a distinct model, a different morph of D. chrysippus. Our findings present a compelling instance of convergent evolution and adaptive atavism, in which the same pattern element has independently evolved multiple times in Hypolimnas butterflies, repeatedly playing a role in Batesian mimicry of diverse model species