%0 Journal Article %J New Phytologist %D 2023 %T Endophytic ancestors of modern leaf miners may have evolved in the Late Carboniferous %A Richard J. Knecht %A Anshuman Swain %A Jacob S. Benner %A Steve L. Emma %A Naomi E. Pierce %A Conrad C. Labandeira %X %B New Phytologist %V 240 %P 2050-2057 %G eng %U https://nph.onlinelibrary.wiley.com/doi/abs/10.1111/nph.19266 %N 5 %0 Journal Article %J Environmental Microbiology %D 2023 %T Carnivorous Nepenthes Pitchers with Less Acidic Fluid House Nitrogen-Fixing Bacteria %A Leonora S. Bittleston %A Charles J. Wolock %A Junko Maeda %A Valentina Infante %A Jean-Michel Ané %A Naomi E. Pierce %A Anne Pringle %X Carnivorous pitcher plants are uniquely adapted to nitrogen limitation, using pitfall traps to acquire nutrients from insect prey. Pitcher plants in the genus Sarracenia may also use nitrogen fixed by bacteria inhabiting the aquatic microcosms of their pitchers. Here, we investigated whether species of a convergently evolved pitcher plant genus, Nepenthes, might also use bacterial nitrogen fixation as an alternative strategy for nitrogen capture. First, we constructed predicted metagenomes of pitcher organisms from three species of Singaporean Nepenthes using 16S rRNA sequence data and correlated predicted nifH abundances with metadata. Second, we used gene-specific primers to amplify and quantify the presence or absence of nifH directly from 102 environmental samples and identified potential diazotrophs with significant differential abundance in samples that also had positive nifH PCR tests. Third, we analyzed nifH in eight shotgun metagenomes from four additional Bornean Nepenthes species. Finally, we conducted an acetylene reduction assay using greenhouse-grown Nepenthes pitcher fluids to confirm nitrogen fixation is indeed possible within the pitcher habitat. Results show active acetylene reduction can occur in Nepenthes pitcher fluid. Variation in nifH from wild samples correlates with Nepenthes host species identity and pitcher fluid acidity. Nitrogen-fixing bacteria are associated with more neutral fluid pH, while endogenous Nepenthes digestive enzymes are most active at low fluid pH. We hypothesize Nepenthes species experience a trade-off in nitrogen acquisition; when fluids are acidic, nitrogen is primarily acquired via plant enzymatic degradation of insects, but when fluids are neutral, Nepenthes plants take up more nitrogen via bacterial nitrogen fixation. %B Environmental Microbiology %V 89 %P e00812-23 %G eng %U https://journals.asm.org/doi/abs/10.1128/aem.00812-23 %N 7 %0 Journal Article %J Ecology and Evolution %D 2023 %T Cycad phylogeny predicts host plant use of Eumaeus butterflies %A Sierra-Botero, Laura %A Calonje, Michael %A Robbins, Robert K. %A Rosser, Neil %A Pierce, Naomi E. %A López-Gallego, Cristina %A Valencia-Montoya, Wendy A. %K butterfly phylogeny %K co-evolution %K gymnosperm herbivory %K phylogenetic tracking %K plant–insect interactions %X Abstract Eumaeus butterflies are obligate herbivores of Zamia, the most diverse neotropical genus of cycads. Eumaeus?Zamia interactions have been characterized mainly for species distributed in North and Central America. However, larval host plant use by the southern Eumaeus clade remains largely unknown, precluding a comprehensive study of co-evolution between the genera. Here, we combine fieldwork with museum and literature surveys to expand herbivory records for Eumaeus from 21 to 38 Zamia species. We inferred a time-calibrated phylogeny of Eumaeus to test for distinct macroevolutionary scenarios of larval host plant conservatism and co-evolution. We found a remarkable coincidence between Eumaeus and Zamia diversification, with the butterfly stem group diverging at the same time as the most recent radiation of Zamia in the Miocene. Cophylogenetic reconciliation analyses show a strong cophylogenetic signal between cycads and their butterfly herbivores. Bipartite model-based approaches indicate that this is because closely related Zamia species are used by the same Eumaeus species, suggesting larval host plant resource tracking by the butterfly herbivores. Our results highlight a case of tight evolution between Eumaeus butterflies and cycads, pointing to the generality of correlated evolution and phylogenetic tracking in plant?herbivore interactions across seed plants. %B Ecology and Evolution %I John Wiley & Sons, Ltd %V 13 %P e9978 %8 2023/04/01 %@ 2045-7758 %G eng %U https://doi.org/10.1002/ece3.9978 %N 4 %! Ecology and Evolution %0 Journal Article %D 2023 %T One in five butterfly species sold online across borders %A Wang, Zhengyang %A Chan, Wei-Ping %A Pham, Nhat Tan %A Zeng, Jimmy %A Pierce, Naomi E. %A Lohman, David J. %A Meng, Weilin %K Aesthetics %K eBay %K Endangered insects %K Global wildlife trade %K Insect conservation %K Web scraping %X Humankind's appreciation for butterflies spans cultures and millennia, including the practice of assembling butterfly collections. We monitored the global e-commerce platform eBay.com for one year and obtained 50,555 time-stamped transactions of 3767 species (739 genera) of butterflies. This is nearly 20% of all butterfly species on Earth. A total of 552 sellers were based in 44 countries across five continents. At least 96% of the traded species required transportation of the specimen from its country of origin to its seller, usually from the Global South to the United States and Europe. To our knowledge, this is the most spatially and temporally detailed record of trans-boundary wildlife movement of any taxonomic group. We quantified the aesthetics of butterflies deemed desirable (e.g., size, shape, and color) and showed that while endangered species command higher prices, a butterfly's aesthetic ranking, not its range, abundance, nor phylogenetic status, best predicts its trade volume. These results emphasize the complicated interplay between wildlife market economics and human aesthetic appreciation. %V 283 %P 110092 %8 2023/07/01/ %@ 0006-3207 %G eng %U https://www.sciencedirect.com/science/article/pii/S0006320723001933 %! Biological Conservation %0 Journal Article %J bioRxiv %D 2023 %T Convergent evolution of noxious heat sensing by TRPA5, a novel class of heat sensor in Rhodnius prolixus %A Marjorie A. Lienard %A David Baez-Nieto %A Cheng-Chia Tsai %A Wendy A. Valencia-Montoya %A Balder Werin %A Urban Johanson %A Jean-Marc Lassance %A Jen Q. Pan %A Nanfang Yu %A Naomi E. Pierce %X Insects are ectotherms, and as such, rely on a diverse repertoire of thermoreceptors to monitor environmental temperature and control behavioral thermoregulation. Here, we use structural, phylogenetic, genetic, and functional analyses to show that TRPA5 genes, widespread across numerous insect orders, encode a novel class of noxious heat receptors. We show that in the triatomine bug Rhodnius prolixus, the primary vector of Chagas disease, Rp-TRPA52 differs biophysically and structurally from noxious thermoTRPAs previously described in insects. This includes key changes in the ankyrin repeat domain and the selectivity filter of the channel. In vitro, we find evidence that the homo-tetrameric channel is not activated by voltage, but displays high thermosensitivity with an enthalpy change (ΔH) of 72 kcal/mol associated with the channel activation, with a Q10 = 25 and T°half = 58.6°C. Structural analyses reveal parallels in the overall ion channel architecture between fruit fly TRPA1 and Rp-TRPA52; however, functional properties and expression patterns indicate that the role of Rp-TRPA52 is more similar to that of Pyrexia noxious heat receptors found in fruit flies. Pyrexia genes have been lost in true bugs, and our findings suggest that the rapidly evolving insect TRPA gene family has given rise to an independent evolutionary origin of a molecular transducer that is responsive to noxious thermal stimuli.Competing Interest StatementThe authors have declared no competing interest. %B bioRxiv %P 2023.05.26.542450 %G eng %U https://www.biorxiv.org/content/biorxiv/early/2023/05/28/2023.05.26.542450.full.pdf %R 10.1101/2023.05.26.542450 %0 Journal Article %J Journal of Experimental Biology %D 2023 %T A hypothesis for robust polarization vision: an example from the Australian imperial blue butterfly, Jalmenus evagoras %A Childers, Richard A. Rabideau %A Bernard, Gary D. %A Huang, Heqing %A Tsai, Cheng-Chia %A Stoddard, Mary Caswell %A Hogan, Benedict G. %A Greenwood, Joel S. F. %A Soucy, Edward R. %A Cornwall, Mark %A Lim, Matthew Lek Min %A Liénard, Marjorie A. %A Yu, Nanfang %A Pierce, Naomi E. %X The Australian lycaenid butterfly Jalmenus evagoras has iridescent wings that are sexually dimorphic, spectrally and in their degree of polarization, suggesting that these properties are likely to be important in mate recognition. We first describe the results of a field experiment showing that free-flying individuals of J. evagoras discriminate between visual stimuli that vary in polarization content in blue wavelengths but not in others. We then present detailed reflectance spectrophotometry measurements of the polarization content of male and female wings, showing that female wings exhibit blue-shifted reflectance, with a lower degree of polarization relative to male wings. Finally, we describe a novel method for measuring alignment of ommatidial arrays: by measuring variation of depolarized eyeshine intensity from patches of ommatidia as a function of eye rotation, we show that (a) individual rhabdoms contain mutually perpendicular microvilli; (b) many rhabdoms in the array have their microvilli misaligned with respect to neighboring rhabdoms by as much as 45 deg; and (c) the misaligned ommatidia are useful for robust polarization detection. By mapping the distribution of the ommatidial misalignments in eye patches of J. evagoras, we show that males and females exhibit differences in the extent to which ommatidia are aligned. Both the number of misaligned ommatidia suitable for robust polarization detection and the number of aligned ommatidia suitable for edge detection vary with respect to both sex and eye patch elevation. Thus, J. evagoras exhibits finely tuned ommatidial arrays suitable for perception of polarized signals, likely to match sex-specific life history differences in the utility of polarized signals. %B Journal of Experimental Biology %V 226 %@ 0022-0949 %G eng %U https://doi.org/10.1242/jeb.244515 %N 7 %1 jeb244515 %R 10.1242/jeb.244515 %0 Journal Article %J Proceedings of the National Academy of Sciences %D 2023 %T The Afrotropical breeding grounds of the Palearctic-African migratory painted lady butterflies (Vanessa cardui) %A Talavera, Gerard %A García-Berro, Aurora %A Talla, Valery N. K. %A Ng’iru, Ivy %A Bahleman, Farid %A Kébé, Khadim %A Nzala, Kelvin M. %A Plasencia, Dulce %A Marafi, Mohammad A. J. %A Kassie, Abeje %A Goudégnon, Eude O. A. %A Kiki, Martial %A Benyamini, Dubi %A Reich, Megan S. %A López-Mañas, Roger %A Benetello, Fulvia %A Collins, Steve C. %A Bataille, Clément P. %A Pierce, Naomi E. %A Martins, Dino J. %A Suchan, Tomasz %A Menchetti, Mattia %A Vila, Roger %X 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. %B Proceedings of the National Academy of Sciences %V 120 %P e2218280120 %G eng %U https://www.pnas.org/doi/abs/10.1073/pnas.2218280120 %N 16 %R doi:10.1073/pnas.2218280120 %0 Journal Article %D 2023 %T A global phylogeny of butterflies reveals their evolutionary history, ancestral hosts and biogeographic origins %A Kawahara, Akito Y. %A Storer, Caroline %A Carvalho, Ana Paula S. %A Plotkin, David M. %A Condamine, Fabien L. %A Braga, Mariana P. %A Ellis, Emily A. %A St Laurent, Ryan A. %A Li, Xuankun %A Barve, Vijay %A Cai, Liming %A Earl, Chandra %A Frandsen, Paul B. %A Owens, Hannah L. %A Valencia-Montoya, Wendy A. %A Aduse-Poku, Kwaku %A Toussaint, Emmanuel F. A. %A Dexter, Kelly M. %A Doleck, Tenzing %A Markee, Amanda %A Messcher, Rebeccah %A Nguyen, Y-Lan %A Badon, Jade Aster T. %A Benítez, Hugo A. %A Braby, Michael F. %A Buenavente, Perry A. C. %A Chan, Wei-Ping %A Collins, Steve C. %A Rabideau Childers, Richard A. %A Dankowicz, Even %A Eastwood, Rod %A Fric, Zdenek F. %A Gott, Riley J. %A Hall, Jason P. W. %A Hallwachs, Winnie %A Hardy, Nate B. %A Sipe, Rachel L. Hawkins %A Heath, Alan %A Hinolan, Jomar D. %A Homziak, Nicholas T. %A Hsu, Yu-Feng %A Inayoshi, Yutaka %A Itliong, Micael G. A. %A Janzen, Daniel H. %A Kitching, Ian J. %A Kunte, Krushnamegh %A Lamas, Gerardo %A Landis, Michael J. %A Larsen, Elise A. %A Larsen, Torben B. %A Leong, Jing V. %A Lukhtanov, Vladimir %A Maier, Crystal A. %A Martinez, Jose I. %A Martins, Dino J. %A Maruyama, Kiyoshi %A Maunsell, Sarah C. %A Mega, Nicolás Oliveira %A Monastyrskii, Alexander %A Morais, Ana B. B. %A Müller, Chris J. %A Naive, Mark Arcebal K. %A Nielsen, Gregory %A Padrón, Pablo Sebastián %A Peggie, Djunijanti %A Romanowski, Helena Piccoli %A Sáfián, Szabolcs %A Saito, Motoki %A Schröder, Stefan %A Shirey, Vaughn %A Soltis, Doug %A Soltis, Pamela %A Sourakov, Andrei %A Talavera, Gerard %A Vila, Roger %A Vlasanek, Petr %A Wang, Houshuai %A Warren, Andrew D. %A Willmott, Keith R. %A Yago, Masaya %A Jetz, Walter %A Jarzyna, Marta A. %A Breinholt, Jesse W. %A Espeland, Marianne %A Ries, Leslie %A Guralnick, Robert P. %A Pierce, Naomi E. %A Lohman, David J. %X Butterflies are a diverse and charismatic insect group that are thought to have evolved with plants and dispersed throughout the world in response to key geological events. However, these hypotheses have not been extensively tested because a comprehensive phylogenetic framework and datasets for butterfly larval hosts and global distributions are lacking. We sequenced 391 genes from nearly 2,300 butterfly species, sampled from 90 countries and 28 specimen collections, to reconstruct a new phylogenomic tree of butterflies representing 92% of all genera. Our phylogeny has strong support for nearly all nodes and demonstrates that at least 36 butterfly tribes require reclassification. Divergence time analyses imply an origin ~100 million years ago for butterflies and indicate that all but one family were present before the K/Pg extinction event. We aggregated larval host datasets and global distribution records and found that butterflies are likely to have first fed on Fabaceae and originated in what is now the Americas. Soon after the Cretaceous Thermal Maximum, butterflies crossed Beringia and diversified in the Palaeotropics. Our results also reveal that most butterfly species are specialists that feed on only one larval host plant family. However, generalist butterflies that consume two or more plant families usually feed on closely related plants. %V 7 %P 903 - 913 %8 2023/06/01 %@ 2397-334X %G eng %U https://doi.org/10.1038/s41559-023-02041-9 %N 6 %! Nature Ecology & Evolution %0 Journal Article %J Ecology and Evolution %D 2023 %T Rapid radiation of ant parasitic butterflies during the Miocene aridification of Africa %A Espeland, Marianne %A Chazot, Nicolas %A Condamine, Fabian L. %A Lemmon, Alan R %A Lemmon, Emily M. %A Pringle, Ernest %A Heath, Alan %A Collins, Steve %A Tiren, Wilson %A Mutiso, Martha %A Lees, David C. %A Fisher, Stewart %A Murphy, Raymond %A Woodhall, Stephen %A Tropek, Robert %A Ahlborn, Svenja S. %A Cockburn, Kevin %A Dobson, Jeremy %A Bouyer, Thierry %A Kaliszewska, Zofia A. %A Baker, Christopher C.M. %A Talavera, Gerard %A Vila, Roger %A Gardiner, Alan J. %A Williams, Mark %A Martins, Dino J. %A Sáfián, Szabolcs %A Edge, David A. %A Pierce, Naomi E. %X Abstract Africa has undergone a progressive aridification during the last 20 My that presumably impacted organisms and fostered the evolution of life history adaptations. We test the hypothesis that shift to living in ant nests and feeding on ant brood by larvae of phyto-predaceous Lepidochrysops butterflies was an adaptive response to the aridification of Africa that facilitated the subsequent radiation of butterflies in this genus. Using anchored hybrid enrichment we constructed a time-calibrated phylogeny for Lepidochrysops and its closest, non-parasitic relatives in the Euchrysops section (Poloyommatini). We estimated ancestral areas across the phylogeny with process-based biogeographical models and diversification rates relying on time-variable and clade-heterogeneous birth-death models. The Euchrysops section originated with the emerging Miombo woodlands about 22 million years ago (Mya) and spread to drier biomes as they became available in the late Miocene. The diversification of the non-parasitic lineages decreased as aridification intensified around 10 Mya, culminating in diversity decline. In contrast, the diversification of the phyto-predaceous Lepidochrysops lineage proceeded rapidly from about 6.5 Mya when this unusual life history likely first evolved. The Miombo woodlands were the cradle for diversification of the Euchrysops section, and our findings are consistent with the hypothesis that aridification during the Miocene selected for a phyto-predaceous life history in species of Lepidochrysops, with ant nests likely providing caterpillars a safe refuge from fire and a source of food when vegetation was scarce. %B Ecology and Evolution %V 13 %P e10046 %@ 2045-7758 %G eng %U https://onlinelibrary.wiley.com/doi/abs/10.1002/ece3.10046 %N 5 %R https://doi.org/10.1002/ece3.10046 %0 Journal Article %J American Journal of Botany %D 2023 %T Herbarium specimens reveal herbivory patterns across the genus Cucurbita %A Jenny, Laura A %A Shapiro, Lori R %A Davis, Charles C %A Jonathan Davies, T %A Pierce, Naomi E %A Meineke, Emily %B American Journal of Botany %I Wiley Online Library %V 110 %P e16126 %@ 0002-9122 %G eng %U https://onlinelibrary.wiley.com/doi/10.1111/mec.16770 %N 2 %0 Journal Article %J Molecular Ecology %D 2023 %T Migratory behaviour is positively associated with genetic diversity in butterflies %A García‐Berro, Aurora %A Talla, Venkat %A Vila, Roger %A Wai, Hong Kar %A Shipilina, Daria %A Chan, Kok Gan %A Pierce, Naomi E %A Backström, Niclas %A Talavera, Gerard %B Molecular Ecology %I Wiley Online Library %V 32 %P 560-574 %@ 0962-1083 %G eng %U https://onlinelibrary.wiley.com/doi/10.1111/mec.16770 %N 3 %0 Journal Article %D 2023 %T Complex hemolymph circulation patterns in grasshopper wings %A Salcedo, Mary K. %A Jun, Brian H. %A Socha, John J. %A Pierce, Naomi E. %A Vlachos, Pavlos P. %A Combes, Stacey A. %X An insect’s living systems—circulation, respiration, and a branching nervous system—extend from the body into the wing. Wing hemolymph circulation is critical for hydrating tissues and supplying nutrients to living systems such as sensory organs across the wing. Despite the critical role of hemolymph circulation in maintaining healthy wing function, wings are often considered “lifeless” cuticle, and flows remain largely unquantified. High-speed fluorescent microscopy and particle tracking of hemolymph in the wings and body of the grasshopper Schistocerca americana revealed dynamic flow in every vein of the fore- and hindwings. The global system forms a circuit, but local flow behavior is complex, exhibiting three distinct types: pulsatile, aperiodic, and “leaky” flow. Thoracic wing hearts pull hemolymph from the wing at slower frequencies than the dorsal vessel; however, the velocity of returning hemolymph (in the hindwing) is faster than in that of the dorsal vessel. To characterize the wing’s internal flow mechanics, we mapped dimensionless flow parameters across the wings, revealing viscous flow regimes. Wings sustain ecologically important insect behaviors such as pollination and migration. Analysis of the wing circulatory system provides a template for future studies investigating the critical hemodynamics necessary to sustaining wing health and insect flight. %V 6 %P 313 %8 2023/03/23 %@ 2399-3642 %G eng %U https://doi.org/10.1038/s42003-023-04651-2 %N 1 %! Communications Biology %0 Journal Article %J Systematic Entomology %D 2023 %T Phylogeny of the Poritiinae (Lepidoptera: Lycaenidae), butterflies with ant associations and unusual lichenivorous diets %A Boyle, John %A Espeland, Marianne %A Sáfián, Szabolcs %A Ducarme, Robert %A Gardiner, Alan %A Coleman, James %A Heath, Alan %A Fisher, Stewart %A Collins, Steve %A Martins, Dino %A Aduse-Poku, Kwaku %A Libert, Michel %A Dankowicz, Even %A Kawahara, Akito %A Lohman, David %A Pierce, Naomi %B Systematic Entomology %8 01 %G eng %U https://doi-org.ezp-prod1.hul.harvard.edu/10.1111/syen.12585 %R 10.1111/syen.12585 %0 Journal Article %J bioRxiv %D 2022 %T Diversification is correlated with temperature in white and sulfur butterflies %A Carvalho, Ana Paula S. %A Owens, Hannah L. %A Laurent, Ryan A. St %A Earl, Chandra %A Dexter, Kelly M. %A Messcher, Rebeccah L. %A Willmott, Keith R. %A Aduse-Poku, Kwaku %A Collins, Steve C. %A Homziak, Nicholas T. %A Hoshizaki, Sugihiko %A Hsu, Yu-Feng %A Kizhakke, Athulya G. %A Kunte, Krushnamegh %A Martins, Dino J. %A Mega, Nicolás O. %A Morinaka, Sadaharu %A Peggie, Djunijanti %A Romanowski, Helena P. %A Sáfián, Szabolcs %A Vila, Roger %A Wang, Houshuai %A Braby, Michael F. %A Espeland, Marianne %A Breinholt, Jesse W. %A Pierce, Naomi E. %A Kawahara, Akito Y. %A Lohman, David J. %X Temperature is thought to be a key variable explaining global patterns of species richness. However, to investigate this relationship carefully, it is necessary to study clades with broad geographic ranges that are comprised of species inhabiting diverse biomes with well- characterized species ranges. In the present study, we investigate the link between temperature and diversification in the butterfly family Pieridae (sulfurs and whites) by combining Next Generation sequences and published molecular data with fine-grained distribution information. After building the most comprehensive phylogeny of the group, with almost 600 species and all higher taxa (subfamilies, tribes and subtribes), we found strong support for the following relationships within the family: Dismorphiinae + (Coliadinae + (Pseudopontiinae + Pierinae)). With a curated distribution dataset of over 800,000 occurrences, we conducted multiple comparative phylogenetic analyses that provided strong evidence that species in environments with more stable daily temperatures or with cooler maximum temperatures in the warm seasons have higher diversification rates. We also found a positive correlation between speciation and extinction with paleotemperature: as global temperature decreased through geological time, so did diversification rates. Although many studies demonstrate higher diversity in the tropics, we have been able to identify specific climate variables associated with changes in diversification, while also inferring the most robust and well sampled phylogenomic framework for Pieridae to date.Competing Interest StatementThe authors have declared no competing interest. %B bioRxiv %I Cold Spring Harbor Laboratory %G eng %U https://www.biorxiv.org/content/early/2022/09/23/2022.09.22.509088 %R 10.1101/2022.09.22.509088 %0 Journal Article %J bioRxiv %D 2022 %T Specialized metabolic convergence in the gut microbiomes of cycad-feeding insects tolerant to β-methylamino-L-alanine (BMAA) %A Gutiérrez-García, Karina %A Whitaker, Melissa R.L. %A Bustos-Díaz, Edder D. %A Salzman, Shayla %A Ramos-Aboites, Hilda E. %A Reitz, Zachary L. %A Pierce, Naomi E. %A Cibrián-Jaramillo, Angélica %A Barona-Gómez, Francisco %X Ingestion of the cycad toxins β-methylamino-L-alanine (BMAA) and azoxyglycosides is harmful to diverse organisms. However, some insects are specialized to feed on toxin-rich cycads with apparent immunity. Some cycad-feeding insects possess a common set of gut bacteria, which might play a role in detoxifying cycad toxins. Here, we investigated the composition of gut microbiota from a worldwide sample of cycadivorous insects and characterized the biosynthetic potential of bacteria isolated as putative keystone taxa. Cycadivorous insects shared a core gut microbiome consisting of six bacterial taxa, mainly belonging to the Proteobacteria, which we were able to isolate. To further investigate these potential keystone taxa from diverging lineages, we performed shotgun metagenomic sequencing of co-cultured bacterial sub-communities. We postulate and characterize four putative keystone bacteria from Serratia, Pantoea, and two different Stenotrophomonas lineages. The biosynthetic potential of these microorganisms includes a suite of biosynthetic gene clusters notably rich in siderophores and carotenoid-like aryl polyene pathways. Siderophore semi-untargeted metabolomics revealed a broad range of chemically related yet diverse iron-chelating metabolites, indicating a complex evolutionary landscape in which siderophores may have converged within the guts of cycadivorous insects. Among these, we provide evidence of the occurrence of an unprecedent desferrioxamine-like biosynthetic pathway that remains to be identified. These results provide a foundation for future investigations into how cycadivorous insects tolerate diets rich in azoxyglycosides, BMAA, and other cycad toxins, and highlight convergent evolution underlying chemical diversity.Competing Interest StatementThe authors have declared no competing interest. %B bioRxiv %I Cold Spring Harbor Laboratory %G eng %U https://www.biorxiv.org/content/early/2022/12/01/2022.12.01.518742 %R 10.1101/2022.12.01.518742 %0 Journal Article %J Philosophical Transactions of the Royal Society B: Biological Sciences %D 2022 %T Molecular advances to study the function, evolution and spectral tuning of arthropod visual opsins %A Liénard, Marjorie A. %A Valencia-Montoya, Wendy A. %A Pierce, Naomi E. %X Visual opsins of vertebrates and invertebrates diversified independently and converged to detect ultraviolet to long wavelengths (LW) of green or red light. In both groups, colour vision largely derives from opsin number, expression patterns and changes in amino acids interacting with the chromophore. Functional insights regarding invertebrate opsin evolution have lagged behind those for vertebrates because of the disparity in genomic resources and the lack of robust in vitro systems to characterize spectral sensitivities. Here, we review bioinformatic approaches to identify and model functional variation in opsins as well as recently developed assays to measure spectral phenotypes. In particular, we discuss how transgenic lines, cAMP-spectroscopy and sensitive heterologous expression platforms are starting to decouple genotype–phenotype relationships of LW opsins to complement the classical physiological-behavioural-phylogenetic toolbox of invertebrate visual sensory studies. We illustrate the use of one heterologous method by characterizing novel LW Gq opsins from 10 species, including diurnal and nocturnal Lepidoptera, a terrestrial dragonfly and an aquatic crustacean, expressing them in HEK293T cells, and showing that their maximum absorbance spectra (λmax) range from 518 to 611 nm. We discuss the advantages of molecular approaches for arthropods with complications such as restricted availability, lateral filters, specialized photochemistry and/or electrophysiological constraints. This article is part of the theme issue ‘Understanding colour vision: molecular, physiological, neuronal and behavioural studies in arthropods’. %B Philosophical Transactions of the Royal Society B: Biological Sciences %V 377 %P 20210279 %G eng %U https://royalsocietypublishing.org/doi/abs/10.1098/rstb.2021.0279 %R 10.1098/rstb.2021.0279 %0 Journal Article %J Frontiers in Ecology and Evolution %D 2022 %T Predictors of sequence capture in a large-scale anchored phylogenomics project %A Nunes, Renato %A Storer, Caroline %A Doleck, Tenzing %A Kawahara, Akito Y. %A Pierce, Naomi E. %A Lohman, David J. %X Next-generation sequencing (NGS) technologies have revolutionized phylogenomics by decreasing the cost and time required to generate sequence data from multiple markers or whole genomes. Further, the fragmented DNA of biological specimens collected decades ago can be sequenced with NGS, reducing the need for collecting fresh specimens. Sequence capture, also known as anchored hybrid enrichment, is a method to produce reduced representation libraries for NGS sequencing. The technique uses single-stranded oligonucleotide probes that hybridize with pre-selected regions of the genome that are sequenced via NGS, culminating in a dataset of numerous orthologous loci from multiple taxa. Phylogenetic analyses using these sequences have the potential to resolve deep and shallow phylogenetic relationships. Identifying the factors that affect sequence capture success could save time, money, and valuable specimens that might be destructively sampled despite low likelihood of sequencing success. We investigated the impacts of specimen age, preservation method, and DNA concentration on sequence capture (number of captured sequences and sequence quality) while accounting for taxonomy and extracted tissue type in a large-scale butterfly phylogenomics project. This project used two probe sets to extract 391 loci or a subset of 13 loci from over 6,000 butterfly specimens. We found that sequence capture is a resilient method capable of amplifying loci in samples of varying age (0–111 years), preservation method (alcohol, papered, pinned), and DNA concentration (0.020 ng/μl - 316 ng/ul). Regression analyses demonstrate that sequence capture is positively correlated with DNA concentration. However, sequence capture and DNA concentration are negatively correlated with sample age and preservation method. Our findings suggest that sequence capture projects should prioritize the use of alcohol-preserved samples younger than 20 years old when available. In the absence of such specimens, dried samples of any age can yield sequence data, albeit with returns that diminish with increasing age. %B Frontiers in Ecology and Evolution %V 10 %G eng %U https://www.frontiersin.org/articles/10.3389/fevo.2022.943361 %R 10.3389/fevo.2022.943361 %0 Journal Article %D 2022 %T LepTraits 1.0 A globally comprehensive dataset of butterfly traits %A Shirey, Vaughn %A Larsen, Elise %A Doherty, Andra %A Kim, Clifford A. %A Al-Sulaiman, Faisal T. %A Hinolan, Jomar D. %A Itliong, Micael Gabriel A. %A Naive, Mark Arcebal K. %A Ku, Minji %A Belitz, Michael %A Jeschke, Grace %A Barve, Vijay %A Lamas, Gerardo %A Kawahara, Akito Y. %A Guralnick, Robert %A Pierce, Naomi E. %A Lohman, David J. %A Ries, Leslie %X Here, we present the largest, global dataset of Lepidopteran traits, focusing initially on butterflies (ca. 12,500 species records). These traits are derived from field guides, taxonomic treatments, and other literature resources. We present traits on wing size, phenology,voltinism, diapause/overwintering stage, hostplant associations, and habitat affinities (canopy, edge, moisture, and disturbance). This dataset will facilitate comparative research on butterfly ecology and evolution and our goal is to inspire future research collaboration and the continued development of this dataset. %V 9 %P 382 %8 2022/07/06 %@ 2052-4463 %G eng %U https://doi.org/10.1038/s41597-022-01473-5 %N 1 %! Scientific Data %0 Generic %D 2022 %T Coevolution of Lepidoptera and their host plants: The Salient Aroma Hypothesis %A Po-An Lin %A Wei-Ping Chan %A Liming Cai %A Even Dankowicz %A Kadeem Gilbert %A Pierce, Naomi E. %A Gary Felton %G eng %U https://doi.org/10.21203/rs.3.rs-1700048/v1 %0 Journal Article %J bioRxiv %D 2022 %T Evolution and Diversification Dynamics of Butterflies %A Kawahara, Akito Y. %A Storer, Caroline %A Carvalho, Ana Paula S. %A Plotkin, David M. %A Condamine, Fabien %A Braga, Mariana P. %A Ellis, Emily A. %A St Laurent, Ryan A. %A Li, Xuankun %A Barve, Vijay %A Cai, Liming %A Earl, Chandra %A Frandsen, Paul B. %A Owens, Hannah L. %A Valencia-Montoya, Wendy A. %A Aduse-Poku, Kwaku %A Toussaint, Emmanuel F. A. %A Dexter, Kelly M. %A Doleck, Tenzing %A Markee, Amanda %A Messcher, Rebeccah %A Nguyen, Y-Lan %A Badon, Jade Aster T. %A Benítez, Hugo A. %A Braby, Michael F. %A Buenavente, Perry A. C. %A Chan, Wei-Ping %A Collins, Steve C. %A Childers, Richard A. Rabideau %A Dankowicz, Even %A Eastwood, Rod %A Fric, Zdenek F. %A Gott, Riley J. %A Hall, Jason P. W. %A Hallwachs, Winnie %A Hardy, Nate B. %A Sipe, Rachel L. Hawkins %A Heath, Alan %A Hinolan, Jomar D. %A Homziak, Nicholas T. %A Hsu, Yu-Feng %A Inayoshi, Yutaka %A Itliong, Micael G.A. %A Janzen, Daniel H. %A Kitching, Ian J. %A Kunte, Krushnamegh %A Lamas, Gerardo %A Landis, Michael J. %A Larsen, Elise A. %A Larsen, Torben B. %A Leong, Jing V. %A Lukhtanov, Vladimir %A Maier, Crystal A. %A Martinez, Jose I. %A Martins, Dino J. %A Maruyama, Kiyoshi %A Maunsell, Sarah C. %A Mega, Nicolás Oliveira %A Monastyrskii, Alexander %A Morais, Ana B. B. %A Müller, Chris J. %A Naive, Mark Arcebal K. %A Nielsen, Gregory %A Padrón, Pablo Sebastián %A Peggie, Djunijanti %A Romanowski, Helena Piccoli %A Sáfián, Szabolcs %A Saito, Motoki %A Schröder, Stefan %A Shirey, Vaughn %A Soltis, Doug %A Soltis, Pamela %A Sourakov, Andrei %A Talavera, Gerard %A Vila, Roger %A Vlasanek, Petr %A Wang, Houshuai %A Warren, Andrew D. %A Willmott, Keith R. %A Yago, Masaya %A Jetz, Walter %A Jarzyna, Marta A. %A Breinholt, Jesse W. %A Espeland, Marianne %A Ries, Leslie %A Guralnick, Robert P. %A Pierce, Naomi E. %A Lohman, David J. %X Butterflies are a diverse and charismatic insect group that are thought to have diversified via coevolution with plants and in response to dispersals following key geological events. These hypotheses have been poorly tested at the macroevolutionary scale because a comprehensive phylogenetic framework and datasets on global distributions and larval hosts of butterflies are lacking. We sequenced 391 genes from nearly 2,000 butterfly species to construct a new, phylogenomic tree of butterflies representing 92% of all genera and aggregated global distribution records and larval host datasets. We found that butterflies likely originated in what is now the Americas, \~100 Ma, shortly before the Cretaceous Thermal Maximum, then crossed Beringia and diversified in the Paleotropics. The ancestor of modern butterflies likely fed on Fabaceae, and most extant families were present before the K/Pg extinction. The majority of butterfly dispersals occurred from the tropics (especially the Neotropics) to temperate zones, largely supporting a “cradle” pattern of diversification. Surprisingly, host breadth changes and shifts to novel host plants had only modest impacts.Competing Interest StatementThe authors have declared no competing interest. %B bioRxiv %I Cold Spring Harbor Laboratory %G eng %U https://www.biorxiv.org/content/early/2022/05/24/2022.05.17.491528 %R 10.1101/2022.05.17.491528 %0 Journal Article %J Journal of Plant Interactions %D 2022 %T A semi-detritivorous pitcher plant, Nepenthes ampullaria diverges in its regulation of pitcher fluid properties %A Kadeem J. Gilbert %A Thibaut Goldsborough %A Weng Ngai Lam %A Felicia Leong %A Naomi E. Pierce %B Journal of Plant Interactions %V 17 %P 956-966 %G eng %U https://www.tandfonline.com/doi/full/10.1080/17429145.2022.2123567 %N 1 %0 Journal Article %J Nature Communications Biology %D 2022 %T A high-throughput multispectral imaging system for museum specimens %A Wei-Ping Chan %A Richard Rabideau Childers %A Sorcha Ashe %A Cheng-Chia Tsai %A Caroline Elson %A Kirsten J. Keleher %A Rachel L. Hawkins Sipe %A Crystal A. Maier %A Andrei Sourakov %A Lawrence F. Gall %A Gary D. Bernard %A Edward R. Soucy %A Nanfang Yu %A Naomi E. Pierce %B Nature Communications Biology %V 5 %G eng %U https://doi.org/10.1038/s42003-022-04282-z %N 1318 %0 Journal Article %J Current Opinion in Insect Science %D 2022 %T Behavioral, ecological and evolutionary mechanisms underlying caterpillar-ant symbioses %A Naomi E Pierce %A Dankowicz, Even %B Current Opinion in Insect Science %V 52 %P 100898 %8 aug %G eng %U https://linkinghub.elsevier.com/retrieve/pii/S2214574522000335 %R 10.1016/j.cois.2022.100898 %0 Journal Article %J Systematic Biology %D 2022 %T DNA Barcodes Combined with Multilocus Data of Representative Taxa Can Generate Reliable Higher-Level Phylogenies %A Gerard Talavera %A Lukhtanov, Vladimir %A Naomi E Pierce %A Vila, Roger %E Bond, Jason %X Abstract Taxa are frequently labeled incertae sedis when their placement is debated at ranks above the species level, such as their subgeneric, generic, or subtribal placement. This is a pervasive problem in groups with complex systematics due to difficulties in identifying suitable synapomorphies. In this study, we propose combining DNA barcodes with a multilocus backbone phylogeny in order to assign taxa to genus or other higher-level categories. This sampling strategy generates molecular matrices containing large amounts of missing data that are not distributed randomly: barcodes are sampled for all representatives, and additional markers are sampled only for a small percentage. We investigate the effects of the degree and randomness of missing data on phylogenetic accuracy using simulations for up to 100 markers in 1000-tips trees, as well as a real case: the subtribe Polyommatina (Lepidoptera: Lycaenidae), a large group including numerous species with unresolved taxonomy. Our simulation tests show that when a strategic and representative selection of species for higher-level categories has been made for multigene sequencing (approximately one per simulated genus), the addition of this multigene backbone DNA data for as few as 5–10% of the specimens in the total data set can produce high-quality phylogenies, comparable to those resulting from 100% multigene sampling. In contrast, trees based exclusively on barcodes performed poorly. This approach was applied to a 1365-specimen data set of Polyommatina (including ca. 80% of described species), with nearly 8% of representative species included in the multigene backbone and the remaining 92% included only by mitochondrial COI barcodes, a phylogeny was generated that highlighted potential misplacements, unrecognized major clades, and placement for incertae sedis taxa. We use this information to make systematic rearrangements within Polyommatina, and to describe two new genera. Finally, we propose a systematic workflow to assess higher-level taxonomy in hyperdiverse groups. This research identifies an additional, enhanced value of DNA barcodes for improvements in higher-level systematics using large data sets. [Birabiro; DNA barcoding; incertae sedis; Kipepeo; Lycaenidae; missing data; phylogenomic; phylogeny; Polyommatina; supermatrix; systematics; taxonomy] %B Systematic Biology %V 71 %P 382–395 %G eng %U https://academic.oup.com/sysbio/article/71/2/382/6281234 %N 2 %R 10.1093/sysbio/syab038 %0 Journal Article %J Environmental Microbiology %D 2022 %T Sarracenia pitcher plant‐associated microbial communities differ primarily by host species across a longitudinal gradient %A Heil, Jacob A. %A Wolock, Charles J. %A Naomi E. Pierce %A Pringle, Anne %A Leonora S. Bittleston %B Environmental Microbiology %V 24 %P 3500–3516 %G eng %U https://onlinelibrary.wiley.com/doi/10.1111/1462-2920.15993 %N 8 %R 10.1111/1462-2920.15993 %0 Book Section %B Caterpillars in the Middle %D 2022 %T The Natural History of Caterpillar-Ant Associations %A Naomi E. Pierce %A Dankowicz, Even %E Marquis, Robert J. %E Koptur, Suzanne %X Abstract The caterpillars of many Lepidoptera are neither attacked nor tended by ants but nevertheless appear to be obligately ant-associated and benefit from the enemy-free space created by ants. Obligate myrmecophiles that do not attract ants through stridulatory or chemical signaling are limited to habitats where ants are reliably present for other reasons, either among ant-attended hemipterans, on ant-plants, or around ant nests. Particularly in the tropics, obligate ant associates that passively coexist with ants are more diverse than previously recognized, including, for example, hundreds of African species in the lycaenid subfamily Poritiinae. Mutualists and parasites of ants have been reported in eleven families: Tineidae, Tortricidae, Cyclotornidae, Coleophoridae, Crambidae, Erebidae, Notodontidae, Hesperiidae, Pieridae, Lycaenidae, and Riodinidae. Altogether, myrmecophily has originated at least 30 times in Lepidoptera, and many groups may remain undiscovered. The butterfly families Lycaenidae and Riodinidae contain the vast majority of ant-associated species: larvae of at least 3841 (71%) of the \textasciitilde5390 described Lycaenidae and 308 (20%) of the \textasciitilde1562 described Riodinidae are known or inferred to be ant-associated, and both families possess specialized, convergently developed exocrine glands and stridulatory devices to communicate with ants. Many caterpillar-ant relationships previously characterized as mutualisms may actually be parasitic, as caterpillars can manipulate ants and ultimately exert a fitness cost. In the family Lycaenidae, highly specialized and obligate ant associations are found largely in the Old World tropics, Australia, and Southern Africa, where the stoichiometry of soil micronutrients, particularly sodium and phosphorus, climate, host plants, and geography may all selectively shape caterpillar-ant associations. %B Caterpillars in the Middle %I Springer International Publishing %C Cham %P 319–391 %@ 978-3-030-86687-7 978-3-030-86688-4 %G eng %U https://link.springer.com/10.1007/978-3-030-86688-4_11 %R 10.1007/978-3-030-86688-4_11 %0 Journal Article %J Nature Communications %D 2022 %T Measuring protected-area effectiveness using vertebrate distributions from leech iDNA %A Yinqiu Ji %A Baker, Christopher C. M. %A Viorel D. Popescu %A Jiaxin Wang %A Chunying Wu %A Zhengyang Wang %A Yuanheng Li %A Lin Wang %A Chaolang Hua %A Zhongxing Yang %A Chunyan Yang %A Charles C.Y. Xu %A Diana, Alex %A Qingzhong Wen %A Naomi E. Pierce %A Douglas W. Yu %X Abstract Protected areas are key to meeting biodiversity conservation goals, but direct measures of effectiveness have proven difficult to obtain. We address this challenge by using environmental DNA from leech-ingested bloodmeals to estimate spatially-resolved vertebrate occupancies across the 677 km 2 Ailaoshan reserve in Yunnan, China. From 30,468 leeches collected by 163 park rangers across 172 patrol areas, we identify 86 vertebrate species, including amphibians, mammals, birds and squamates. Multi-species occupancy modelling shows that species richness increases with elevation and distance to reserve edge. Most large mammals (e.g. sambar, black bear, serow, tufted deer) follow this pattern; the exceptions are the three domestic mammal species (cows, sheep, goats) and muntjak deer, which are more common at lower elevations. Vertebrate occupancies are a direct measure of conservation outcomes that can help guide protected-area management and improve the contributions that protected areas make towards global biodiversity goals. Here, we show the feasibility of using invertebrate-derived DNA to estimate spatially-resolved vertebrate occupancies across entire protected areas. %B Nature Communications %V 13 %P 1555 %G eng %U https://www.nature.com/articles/s41467-022-28778-8 %N 1 %R 10.1038/s41467-022-28778-8 %0 Journal Article %J Insect Sci %D 2022 %T Agent-based models reveal limits of mark-release-recapture estimates for the rare butterfly, Bhutanitis thaidina (Lepidoptera: Papilionidae) %A Zhengyang Wang %A Yuanheng Li %A Jain, Anuj %A Naomi E Pierce %K Animals %K butterflies %K Environment %K insecta %K Population Density %X Insect diversity and abundance are in drastic decline worldwide, but quantifying insect populations to better conserve them is a difficult task. Mark-release-recapture (MRR) is widely used as an ecological indicator for insect populations, but the accuracy of MRR estimates can vary with factors such as spatial scale, sampling effort and models of inference. We conducted a 3-year MRR study of B. thaidina in Yanzigou valley, Mt. Gongga but failed to obtain sufficient data for a robust population estimate. This prompted us to integrate B. thaidina life history information to parameterize agent-based models and evaluate the conditions under which successful MRR studies could be conducted. We evaluated: (1) the performance of MRR models under different landscape types, and (2) the influence of experimental design on the accuracy and variance of MRR-based estimates. Our simulations revealed systematic underestimates of true population parameters by MRR models when sampling effort was insufficient. In a total of 2772 simulations, subjective decisions in sampling protocol (e.g., frequency, number of sampling locations, use of spatially explicit models, type of estimands) accounted for nearly half of the variation in estimates. We conclude that MRR-based estimates could be improved with the addition of more field-specific parameters. %B Insect Sci %V 29 %P 550-566 %8 2022 Apr %G eng %N 2 %1 http://www.ncbi.nlm.nih.gov/pubmed/34263543?dopt=Abstract %R 10.1111/1744-7917.12949 %0 Journal Article %J Front Plant Sci %D 2022 %T Characterization and Comparison of Convergence Among Cephalotus follicularis Pitcher Plant-Associated Communities With Those of Nepenthes and Sarracenia Found Worldwide %A Bittleston, Leonora S %A Benson, Elizabeth L %A Bernardin, Jessica R %A Naomi E Pierce %X The Albany pitcher plant, Cephalotus follicularis, has evolved cup-shaped leaves and a carnivorous habit completely independently from other lineages of pitcher plants. It is the only species in the family Cephalotaceae and is restricted to a small region of Western Australia. Here, we used metabarcoding to characterize the bacterial and eukaryotic communities living in C. follicularis pitchers at two different sites. Bacterial and eukaryotic communities were correlated in both richness and composition; however, the factors associated with richness were not the same across bacteria and eukaryotes, with bacterial richness differing with fluid color, and eukaryotic richness differing with the concentration of DNA extracted from the fluid, a measure roughly related to biomass. For turnover in composition, the variation in both bacterial and eukaryotic communities primarily differed with fluid acidity, fluid color, and sampling site. We compared C. follicularis-associated community diversity with that of Australian Nepenthes mirabilis, as well as a global comparison of Southeast Asian Nepenthes and North American Sarracenia. Our results showed similarity in richness with communities from other pitcher plants, and specific bacterial taxa shared among all three independent lineages of pitcher plants. Overall, we saw convergence in richness and particular clades colonizing pitcher plants around the world, suggesting that these highly specialized habitats select for certain numbers and types of inhabitants. %B Front Plant Sci %V 13 %P 887635 %8 2022 %G eng %1 http://www.ncbi.nlm.nih.gov/pubmed/35734258?dopt=Abstract %R 10.3389/fpls.2022.887635 %0 Journal Article %J Nat Ecol Evol %D 2022 %T Edward O. Wilson (1929-2021) %A Moreau, Corrie S %A Naomi E Pierce %B Nat Ecol Evol %V 6 %P 240-241 %8 2022 03 %G eng %N 3 %1 http://www.ncbi.nlm.nih.gov/pubmed/35115671?dopt=Abstract %R 10.1038/s41559-022-01680-8 %0 Journal Article %J Ecol Evol %D 2022 %T Evidence of nutrient translocation in response to smoke exposure by the East African ant acacia, Vachellia drepanolobium %A Rabideau-Childers, Richard %A Angier, Katherine I W %A Dean, Brendan Z M %A Blumstein, Meghan %A Darling, Walker S %A Kennedy-Yoon, Annina %A Ziemke, Clayton H %A Perez-Martinez, Christian A %A Wu, Donghao %A Wenqing Ye %A Yekwayo, Inam %A Kimuyu, Duncan M %A Martins, Dino J %A Naomi E Pierce %X Fire is a major selective force on arid grassland communities, favoring traits such as the smoke-induced seed germination response seen in a wide variety of plant species. However, little is known about the relevance of smoke as a cue for plants beyond the seedling stage.We exposed a fire-adapted savanna tree, Vachellia (=Acacia) drepanolobium, to smoke and compared nutrient concentrations in leaf and root tissues to unexposed controls. Experiments were performed on three age cohorts: 2-year-old, 9-month-old, and 3-month-old plants.For the 2-year-old plants exposed to smoke, carbon and nitrogen concentrations were lower in the leaves and higher in the roots than controls. Less pronounced trends were found for boron and magnesium.In contrast, smoke-exposed 3-month-old plants had lower root nitrogen concentrations than controls. No significant differences were found in the 9-month-old plants, and no significant shifts in other nutrient concentrations were observed between plant tissues for any of the three age cohorts. Synthesis: Our findings are consistent with smoke-induced translocation of nutrients from leaves to roots in 2-year-old Vdrepanolobium. This could represent a novel form of fire adaptation, with variation over the course of plant development. The translocation differences between age cohorts highlight the need to investigate smoke response in older plants of other species. Accounting for this adaptation could better inform our understanding of savanna community structure and nutrient flows under fire regimes altered by anthropogenic land use and climate change. %B Ecol Evol %V 12 %P e8244 %8 2022 Jan %G eng %N 1 %1 http://www.ncbi.nlm.nih.gov/pubmed/35126997?dopt=Abstract %R 10.1002/ece3.8244 %0 Journal Article %J Mol Ecol %D 2022 %T Fine-scale genome-wide signature of Pleistocene glaciation in Thitarodes moths (Lepidoptera: Hepialidae), host of Ophiocordyceps fungus in the Hengduan Mountains %A Zhengyang Wang %A Naomi E Pierce %X The Hengduan Mountains region is a biodiversity hotspot known for its topologically complex, deep valleys and high mountains. While landscape and glacial refugia have been evoked to explain patterns of interspecies divergence, the accumulation of intra-species (i.e., population level) genetic divergence across the mountain-valley landscape in this region has received less attention. We used genome-wide restriction site-associated DNA sequencing (RADseq) to reveal signatures of Pleistocene glaciation in populations of Thitarodes shambalaensis (Lepidoptera: Hepialidae), the host moth of parasitic Ophiocordyceps sinensis (Hypocreales: Ophiocordycipitaceae) or "caterpillar fungus" endemic to the glacier of eastern Mt. Gongga. We used moraine history along the glacier valleys to model the distribution and environmental barriers to gene flow across populations of T. shambalaensis. We found that moth populations separated by less than 10 km exhibited valley-based population genetic clustering and isolation-by-distance (IBD), while gene flow among populations was best explained by models using information about their distributions at the local last glacial maximum (LGML , 58 kya), not their contemporary distribution. Maximum likelihood lineage history among populations, and among subpopulations as little as 500 m apart, recapitulated glaciation history across the landscape. We also found signals of isolated population expansion following the retreat of LGML glaciers. These results reveal the fine-scale, long-term historical influence of landscape and glaciation on the genetic structuring of populations of an endangered and economically important insect species. Similar mechanisms, given enough time and continued isolation, could explain the contribution of glacier refugia to the generation of species diversity among the Hengduan Mountains. %B Mol Ecol %8 2022 Apr 04 %G eng %1 http://www.ncbi.nlm.nih.gov/pubmed/35377501?dopt=Abstract %R 10.1111/mec.16457 %0 Journal Article %J Proc Biol Sci %D 2022 %T Profiling, monitoring and conserving caterpillar fungus in the Himalayan region using anchored hybrid enrichment markers %A Zhengyang Wang %A Da, Wa %A Negi, Chandra Singh %A Ghimire, Puspa Lal %A Wangdi, Karma %A Yadav, Pramod K %A Pubu, Zhuoma %A Lama, Laiku %A Yarpel, Kuenga %A Maunsell, Sarah C %A Yong Liu %A Kunte, Krushnamegh %A Bawa, Kamaljit S %A Darong Yang %A Naomi E Pierce %K Asia %K fungi %K Geography %X The collection of caterpillar fungus accounts for 50-70% of the household income of thousands of Himalayan communities and has an estimated market value of $5-11 billion across Asia. However, Himalayan collectors are at multiple economic disadvantages compared with collectors on the Tibetan Plateau because their product is not legally recognized. Using a customized hybrid-enrichment probe set and market-grade caterpillar fungus (with samples up to 30 years old) from 94 production zones across Asia, we uncovered clear geography-based signatures of historical dispersal and significant isolation-by-distance among caterpillar fungus hosts. This high-throughput approach can readily distinguish samples from major production zones with definitive geographical resolution, especially for samples from the Himalayan region that form monophyletic clades in our analysis. Based on these results, we propose a two-step procedure to help local communities authenticate their produce and improve this multi-national trade-route without creating opportunities for illegal exports and other forms of economic exploitation. We argue that policymakers and conservation practitioners must encourage the fair trade of caterpillar fungus in addition to sustainable harvesting to support a trans-boundary conservation effort that is much needed for this natural commodity in the Himalayan region. %B Proc Biol Sci %V 289 %P 20212650 %8 2022 04 27 %G eng %N 1973 %1 http://www.ncbi.nlm.nih.gov/pubmed/35473372?dopt=Abstract %R 10.1098/rspb.2021.2650 %0 Journal Article %J PNAS %D 2021 %T The evolution of red color vision is linked to coordinated rhodopsin tuning in lycaenid butterflies %A Marjorie A Lienard %A Gary D Bernard %A Andrew Allen %A Jean-Marc Lassance %A Siliang Song %A Richard Rabideau Childers %A Nanfang Yu %A Dajia Ye %A Adriana Stephenson %A Wendy A Valencia-Montoya %A Shayla Salzman %A Melissa R L Whitaker %A Michael Calonje %A Zhang, Feng %A Naomi E Pierce %B PNAS %V 118 %G eng %N 6 %0 Journal Article %J Frontiers in Plant Science %D 2021 %T Cycad-Weevil Pollination Symbiosis Is Characterized by Rapidly Evolving and Highly Specific Plant-Insect Chemical Communication %A Shayla Salzman %A Damon Crook %A Michael Calonje %A Stevenson, Dennis W. %A Naomi E. Pierce %A Robin Hopkins %X Coevolution between plants and insects is thought to be responsible for generating biodiversity. Extensive research has focused largely on antagonistic herbivorous relationships, but mutualistic pollination systems also likely contribute to diversification. Here we describe an example of chemically-mediated mutualistic species interactions affecting trait evolution and lineage diversification. We show that volatile compounds produced by closely related species of Zamia cycads are more strikingly different from each other than are other phenotypic characters, and that two distantly related pollinating weevil species have specialized responses only to volatiles from their specific host Zamia species. Plant transcriptomes show that approximately a fifth of genes related to volatile production are evolving under positive selection, but we find no differences in the relative proportion of genes under positive selection in different categories. The importance of phenotypic divergence coupled with chemical communication for the maintenance of this obligate mutualism highlights chemical signaling as a key mechanism of coevolution between cycads and their weevil pollinators. %B Frontiers in Plant Science %V 12 %P 639368 %G eng %U https://www.frontiersin.org/articles/10.3389/fpls.2021.639368/full %R 10.3389/fpls.2021.639368 %0 Journal Article %J Biology Letters %D 2021 %T Museum genomics reveals the Xerces blue butterfly (Glaucopsyche xerces) was a distinct species driven to extinction %A Grewe, Felix %A Kronforst, Marcus R. %A Naomi E. Pierce %A Moreau, Corrie S. %X The last Xerces blue butterfly was seen in the early 1940s, and its extinction is credited to human urban development. This butterfly has become a North American icon for insect conservation, but some have questioned whether it was truly a distinct species, or simply an isolated population of another living species. To address this question, we leveraged next-generation sequencing using a 93-year-old museum specimen. We applied a genome skimming strategy that aimed for the organellar genome and high-copy fractions of the nuclear genome by a shallow sequencing approach. From these data, we were able to recover over 200 million nucleotides, which assembled into several phylogenetically informative markers and the near-complete mitochondrial genome. From our phylogenetic analyses and haplotype network analysis we conclude that the Xerces blue butterfly was a distinct species driven to extinction. %B Biology Letters %V 17 %P 20210123 %G eng %U https://royalsocietypublishing.org/doi/10.1098/rsbl.2021.0123 %N 7 %R 10.1098/rsbl.2021.0123 %0 Journal Article %J Insect Conservation and Diversity %D 2021 %T Out of sight, out of mind: public and research interest in insects is negatively correlated with their conservation status %A Zhengyang Wang %A Zeng, Jimmy %A Meng, Weilin %A David J. Lohman %A Naomi E. Pierce %B Insect Conservation and Diversity %V 14 %P 700–708 %G eng %U https://onlinelibrary.wiley.com/doi/10.1111/icad.12499 %N 5 %R 10.1111/icad.12499 %0 Journal Article %J Proc Biol Sci %D 2021 %T Evolutionary trade-offs between male secondary sexual traits revealed by a phylogeny of the hyperdiverse tribe Eumaeini (Lepidoptera: Lycaenidae) %A Wendy A Valencia-Montoya %A Quental, Tiago B %A Tonini, João Filipe R %A Gerard Talavera %A Crall, James D %A Lamas, Gerardo %A Busby, Robert C %A Carvalho, Ana Paula S %A Morais, Ana B %A Oliveira Mega, Nicolás %A Romanowski, Helena Piccoli %A Liénard, Marjorie A %A Shayla Salzman %A Melissa R L Whitaker %A Akito Y. Kawahara %A Lohman, David J %A Robbins, Robert K %A Naomi E Pierce %K Animals %K Biological Evolution %K butterflies %K Male %K Phenotype %K pheromones %K Phylogeny %X Male butterflies in the hyperdiverse tribe Eumaeini possess an unusually complex and diverse repertoire of secondary sexual characteristics involved in pheromone production and dissemination. Maintaining multiple sexually selected traits is likely to be metabolically costly, potentially resulting in trade-offs in the evolution of male signals. However, a phylogenetic framework to test hypotheses regarding the evolution and maintenance of male sexual traits in Eumaeini has been lacking. Here, we infer a comprehensive, time-calibrated phylogeny from 379 loci for 187 species representing 91% of the 87 described genera. Eumaeini is a monophyletic group that originated in the late Oligocene and underwent rapid radiation in the Neotropics. We examined specimens of 818 of the 1096 described species (75%) and found that secondary sexual traits are present in males of 91% of the surveyed species. Scent pads and scent patches on the wings and brush organs associated with the genitalia were probably present in the common ancestor of Eumaeini and are widespread throughout the tribe. Brush organs and scent pads are negatively correlated across the phylogeny, exhibiting a trade-off in which lineages with brush organs are unlikely to regain scent pads and vice versa. In contrast, scent patches seem to facilitate the evolution of scent pads, although they are readily lost once scent pads have evolved. Our results illustrate the complex interplay between natural and sexual selection in the origin and maintenance of multiple male secondary sexual characteristics and highlight the potential role of sexual selection spurring diversification in this lineage. %B Proc Biol Sci %V 288 %P 20202512 %8 2021 05 12 %G eng %N 1950 %1 http://www.ncbi.nlm.nih.gov/pubmed/33975481?dopt=Abstract %R 10.1098/rspb.2020.2512 %0 Journal Article %J PNAS %D 2021 %T The evolution of red color vision is linked to coordinated rhodopsin tuning in lycaenid butterflies %A Marjorie A Lienard %A Gary D Bernard %A Andrew Allen %A Jean-Marc Lassance %A Siliang Song %A Richard Rabideau Childers %A Nanfang Yu %A Dajia Ye %A Adriana Stephenson %A Wendy A Valencia-Montoya %A Shayla Salzman %A Melissa R L Whitaker %A Michael Calonje %A Zhang, Feng %A Naomi E Pierce %B PNAS %V 118 %G eng %N 6 %0 Journal Article %J ZooKeys %D 2020 %T Ants of the Hengduan Mountains: a new altitudinal survey and updated checklist for Yunnan Province highlight an understudied insect biodiversity hotspot %A Cong Liu %A Georg Fischer %A Francisco Hita Garcia %A Seiki Yamane %A Qing Liu %A Yan Qiong Peng %A Evan P. Economo %A Benoit Guénard %A Naomi E. Pierce %B ZooKeys %V 978 %P 1-171 %G eng %0 Journal Article %J Applied and Environmental Microbiology %D 2020 %T In situ activation and heterologous production of a cryptic lantibiotic from an African plant ant-derived Saccharopolyspora species %A Eleni Vikeli %A David A. Widdick %A Sibyl F. Batey %A Daniel Heine %A Neil A. Holmes %A Mervyn J. Bibb %A Dino J. Martins %A Naomi E. Pierce %A Matthew I. Hutchings %A Barrie Wilkinson %E Charles M. Dozois %B Applied and Environmental Microbiology %V 86 %P e01876-19 %G eng %N 3 %0 Journal Article %J Fungal Ecology %D 2020 %T The entomophagous caterpillar fungus Ophiocordyceps sinensis is consumed by its lepidopteran host as a plant endophyte %A Zhengyang Wang %A Meng Li %A Wenbin Ju %A Wenqing Ye %A Longha iXue %A David E. Boufford %A Xinfen Gao %A Bisong Yue %A Yong Liu %A Naomi E. Pierce %B Fungal Ecology %V 47 %G eng %0 Journal Article %J Microbial Ecology %D 2020 %T Investigation of an Elevational Gradient Reveals Strong Differences Between Bacterial and Eukaryotic Communities Coinhabiting Nepenthes Phytotelmata %A Kadeem J. Gilbert %A Leonora S. Bittleston %A Mark Arcebal K. Naive %A Anthony E. Kiszewski %A Perry Archival C. Buenavente %A David J. Lohman %A Naomi E. Pierce %B Microbial Ecology %G eng %0 Journal Article %J Insectes Sociaux %D 2020 %T Mating system, population genetics, and phylogeography of the devil’s garden ant, Myrmelachista schumanni, in the Peruvian Amazon %A P. J. G. Malé %A Youngerman, E. %A Pierce, N. E. %A Frederickson, M. E. %B Insectes Sociaux %V 67 %P 113-125 %G eng %0 Journal Article %J Systematic Entomology %D 2020 %T Molecular phylogeny of the tribe Candalidini (Lepidoptera: Lycaenidae): systematics, diversification and evolutionary history %A Michael F. Braby %A Marianne Espeland %A Chris J. Müller %A Eastwood, Rod %A David J. Lohman %A Akito Y. Kawahara %A Sarah C. Maunsell %A Naomi E. Pierce %B Systematic Entomology %V 45 %P 703-722 %G eng %N 3 %0 Journal Article %J Nature Communications %D 2020 %T Physical and behavioral adaptations to prevent overheating of the living wings of butterflies %A Cheng-Chia Tsai %A Richard A. Childers %A Norman Nan Shi %A Crystal Ren %A Julianne N. Pelaez %A Gary D. Bernard %A Naomi E. Pierce %A Nanfang Yu %B Nature Communications %V 11 %G eng %0 Journal Article %J Molecular Phylogenetics and Evolution %D 2020 %T Recent diversification of Chrysoritis butterflies in the South African Cape (Lepidoptera: Lycaenidae) %A Gerard Talavera %A Zofia A. Kaliszeksa %A Alan Heath %A Naomi E. Pierce %B Molecular Phylogenetics and Evolution %V 128 %G eng %0 Journal Article %J Scientific Reports %D 2020 %T Tropical pitcher plants (Nepenthes) act as ecological filters by altering properties of their fluid microenvironments %A Kadeem J. Gilbert %A Leonora S. Bittleston %A Wenfei Tong %A Naomi E. Pierce %B Scientific Reports %V 10 %G eng %0 Journal Article %J Biology Letters %D 2020 %T Wind drives temporal variation in pollinator visitation in a fragmented tropical forest %A James D. Crall %A Julia Brokaw %A Susan F. Gagliardi %A Chase D. Mendenhall %A Naomi E. Pierce %A Stacey A. Combes %B Biology Letters %V 16 %G eng %N 4 %0 Journal Article %J ZooKeys %D 2020 %T Ants of the Hengduan Mountains: a new altitudinal survey and updated checklist for Yunnan Province highlight an understudied insect biodiversity hotspot %A Cong Liu %A Georg Fischer %A Francisco Hita Garcia %A Seiki Yamane %A Qing Liu %A Yan Qiong Peng %A Evan P. Economo %A Benoit Guénard %A Naomi E. Pierce %B ZooKeys %V 978 %P 1-171 %G eng %0 Journal Article %J Fungal Ecology %D 2020 %T The entomophagous caterpillar fungus Ophiocordyceps sinensis is consumed by its lepidopteran host as a plant endophyte %A Zhengyang Wang %A Meng Li %A Wenbin Ju %A Wenqing Ye %A Longha iXue %A David E. Boufford %A Xinfen Gao %A Bisong Yue %A Yong Liu %A Naomi E. Pierce %B Fungal Ecology %V 47 %G eng %0 Journal Article %J Systematic Entomology %D 2020 %T Molecular phylogeny of the tribe Candalidini (Lepidoptera: Lycaenidae): systematics, diversification and evolutionary history %A Michael F. Braby %A Marianne Espeland %A Chris J. Müller %A Eastwood, Rod %A David J. Lohman %A Akito Y. Kawahara %A Sarah C. Maunsell %A Naomi E. Pierce %B Systematic Entomology %V 45 %P 703-722 %G eng %N 3 %0 Journal Article %J Science Advances %D 2020 %T An ancient push-pull pollination mechanism in cycads %A Shayla Salzman %A Damon Crook %A James D. Crall %A Robin Hopkins %A Naomi E. Pierce %B Science Advances %V 6 %G eng %N 24 %0 Journal Article %J Journal of Economic Entomology %D 2020 %T Report on the Emergence Time of a Species of Thitarodes Ghost Moth (Lepidoptera: Hepialidae), Host of the Caterpillar Fungus Ophiocordyceps sinensis (Ascomycota: Ophiocordycipitaceae) in Uttarakhand, India %A Zhengyang Wang %A Darong Yang %A Rawal Janak %A Naomi E. Pierce %B Journal of Economic Entomology %V 113 %P 2031-2034 %G eng %N 4 %0 Journal Article %J Microbial Ecology %D 2020 %T Investigation of an Elevational Gradient Reveals Strong Differences Between Bacterial and Eukaryotic Communities Coinhabiting Nepenthes Phytotelmata %A Kadeem J. Gilbert %A Leonora S. Bittleston %A Mark Arcebal K. Naive %A Anthony E. Kiszewski %A Perry Archival C. Buenavente %A David J. Lohman %A Naomi E. Pierce %B Microbial Ecology %G eng %0 Journal Article %J Molecular Phylogenetics and Evolution %D 2020 %T Recent diversification of Chrysoritis butterflies in the South African Cape (Lepidoptera: Lycaenidae) %A Gerard Talavera %A Zofia A. Kaliszeksa %A Alan Heath %A Naomi E. Pierce %B Molecular Phylogenetics and Evolution %V 128 %G eng %0 Journal Article %J Scientific Reports %D 2020 %T Tropical pitcher plants (Nepenthes) act as ecological filters by altering properties of their fluid microenvironments %A Kadeem J. Gilbert %A Leonora S. Bittleston %A Wenfei Tong %A Naomi E. Pierce %B Scientific Reports %V 10 %G eng %0 Journal Article %J Insectes Sociaux %D 2020 %T Mating system, population genetics, and phylogeography of the devil’s garden ant, Myrmelachista schumanni, in the Peruvian Amazon %A P. J. G. Malé %A Youngerman, E. %A Pierce, N. E. %A Frederickson, M. E. %B Insectes Sociaux %V 67 %P 113-125 %G eng %0 Journal Article %J Nature Communications %D 2020 %T Physical and behavioral adaptations to prevent overheating of the living wings of butterflies %A Cheng-Chia Tsai %A Richard A. Childers %A Norman Nan Shi %A Crystal Ren %A Julianne N. Pelaez %A Gary D. Bernard %A Naomi E. Pierce %A Nanfang Yu %B Nature Communications %V 11 %G eng %0 Journal Article %J Applied and Environmental Microbiology %D 2020 %T In situ activation and heterologous production of a cryptic lantibiotic from an African plant ant-derived Saccharopolyspora species %A Eleni Vikeli %A David A. Widdick %A Sibyl F. Batey %A Daniel Heine %A Neil A. Holmes %A Mervyn J. Bibb %A Dino J. Martins %A Naomi E. Pierce %A Matthew I. Hutchings %A Barrie Wilkinson %E Charles M. Dozois %B Applied and Environmental Microbiology %V 86 %P e01876-19 %G eng %N 3 %0 Journal Article %J Biology Letters %D 2020 %T Wind drives temporal variation in pollinator visitation in a fragmented tropical forest %A James D. Crall %A Julia Brokaw %A Susan F. Gagliardi %A Chase D. Mendenhall %A Naomi E. Pierce %A Stacey A. Combes %B Biology Letters %V 16 %G eng %N 4 %0 Journal Article %J PLOS ONE %D 2019 %T Combining stable isotope analysis with DNA metabarcoding improves inferences of trophic ecology %A Melissa R. L. Whitaker %A Chistopher C. M. Baker %A Shayla M. Salzman %A Dino J. Martins %A Naomi E. Pierce %B PLOS ONE %G eng %0 Journal Article %J Zookeys %D 2019 %T Thitarodes shambalaensis sp. nov. (Lepidoptera, Hepialidae): a new host of the caterpillar fungus Ophiocordyceps sinensis supported by genome-wide SNP data %A Zhengyang Wang %A Hailing Zhuang %A Min Wang %A Naomi E. Pierce %B Zookeys %V 885 %P 85-113 %G eng %0 Journal Article %J Frontiers in Ecology and Evolution %D 2019 %T Population Genomics and Demographic Sampling of the Ant-Plant Vachellia drepanolobium and Its Symbiotic Ants From Sites Across Its Range in East Africa %A John H. Boyle %A Dino Martens %A Paul M. Musili %A Naomi E. Pierce %B Frontiers in Ecology and Evolution %V 7 %G eng %0 Journal Article %J Journal of Insect Conservation %D 2019 %T Radio telemetry helps record the dispersal patterns of birdwingbutterfies in mountainous habitats: Golden Birdwing (Troides aeacus)as an example %A Zhengyang Wang %A Yuan Huang %A Naomi E. Pierce %B Journal of Insect Conservation %V 23 %P 729-738 %G eng %0 Journal Article %J Frontiers in Ecology and Evolution %D 2019 %T Social Buffering of Pesticides in Bumblebees: Agent-Based Modeling of the Effects of Colony Size and Neonicotinoid Exposure on Behavior Within Nests %A James D. Crall %A Benjamin L. de Bivort %A Biswadip Dey %A Ashlee N. Ford %B Frontiers in Ecology and Evolution %V 7 %G eng %0 Journal Article %J PLOS Pathogens %D 2019 %T Symbiotic microbiota may reflect host adaptation by resident to invasive ant species %A Daifeng Cheng %A Siqi Chen %A Yuquan Huang %A Naomi E. Pierce %A Markus Riegler %A Fan Yang %A Ling Zeng %A Yongyue Lu %A Guangwen Liang %A Yijuan Xu %B PLOS Pathogens %V 15 %G eng %N 7 %0 Journal Article %J PLOS ONE %D 2019 %T Combining stable isotope analysis with DNA metabarcoding improves inferences of trophic ecology %A Melissa R. L. Whitaker %A Chistopher C. M. Baker %A Shayla M. Salzman %A Dino J. Martins %A Naomi E. Pierce %B PLOS ONE %G eng %0 Journal Article %J Zookeys %D 2019 %T Thitarodes shambalaensis sp. nov. (Lepidoptera, Hepialidae): a new host of the caterpillar fungus Ophiocordyceps sinensis supported by genome-wide SNP data %A Zhengyang Wang %A Hailing Zhuang %A Min Wang %A Naomi E. Pierce %B Zookeys %V 885 %P 85-113 %G eng %0 Journal Article %J PLOS Pathogens %D 2019 %T Symbiotic microbiota may reflect host adaptation by resident to invasive ant species %A Daifeng Cheng %A Siqi Chen %A Yuquan Huang %A Naomi E. Pierce %A Markus Riegler %A Fan Yang %A Ling Zeng %A Yongyue Lu %A Guangwen Liang %A Yijuan Xu %B PLOS Pathogens %V 15 %G eng %N 7 %0 Journal Article %J Frontiers in Ecology and Evolution %D 2019 %T Social Buffering of Pesticides in Bumblebees: Agent-Based Modeling of the Effects of Colony Size and Neonicotinoid Exposure on Behavior Within Nests %A James D. Crall %A Benjamin L. de Bivort %A Biswadip Dey %A Ashlee N. Ford %B Frontiers in Ecology and Evolution %V 7 %G eng %0 Journal Article %J Journal of Insect Conservation %D 2019 %T Radio telemetry helps record the dispersal patterns of birdwingbutterfies in mountainous habitats: Golden Birdwing (Troides aeacus)as an example %A Zhengyang Wang %A Yuan Huang %A Naomi E. Pierce %B Journal of Insect Conservation %V 23 %P 729-738 %G eng %0 Journal Article %J Frontiers in Ecology and Evolution %D 2019 %T Population Genomics and Demographic Sampling of the Ant-Plant Vachellia drepanolobium and Its Symbiotic Ants From Sites Across Its Range in East Africa %A John H. Boyle %A Dino Martens %A Paul M. Musili %A Naomi E. Pierce %B Frontiers in Ecology and Evolution %V 7 %G eng %0 Journal Article %J BMC Evolutionary Biology %D 2018 %T Anchored phylogenomics illuminates the skipper butterfly tree of life. %A Toussaint, E.F.A. %A Breinholt, J.W. %A Earl, C. %A Warren, A. D. %A Brower, J. V.Z. %A Yago, M. %A Dexter, K.M. %A Espeland, M. %A Pierce, N. E. %A Lohman, D. J. %A Kawahara, A. Y. %B BMC Evolutionary Biology %G eng %0 Journal Article %J Current Biology %D 2018 %T Comprehensive higher-level phylogeny of butterflies (Papilionoidea) inferred from genomic data. %A Espeland, M. %A Breinholt, J. %A Willmott, K. R. %A Warren, A. D. %A Vila, R. %A Toussaint, E.F.A. %A Maunsell, S.C. %A Kwaku, A. P. %A Talavera, G. %A Eastwood, R. %A Jarzyna, M.A. %A Ries, L. %A Guralinick, R. %A Lohman, D. J. %A Pierce, N. E. %A Kawahara, A. Y. %B Current Biology %V 28 %P 770-778 %G eng %U http://doi.org/10.1016/jcub.2018.01.061 %0 Journal Article %J Elife %D 2018 %T Convergence between the microcosms of Southeast Asian and North American pitcher plants. %A Bittleston, L. S. %A Wolock, C.J. %A Yahya, B.E. %A Chan, X.Y. %A Chan, K.G. %A Pierce, N. E. %A Pringle, A. %B Elife %G eng %0 Journal Article %J Biological Journal of the Linnaean Society %D 2018 %T Cycad-feeding insects share a core gut microbiome. %A Salzman, S. %A Whitaker, M. %A Pierce, N. E. %B Biological Journal of the Linnaean Society %V 123 %P 728-738 %G eng %U http://doi-org.ezp-prod1.hul.harvard.edu/10.1093/biolinnean/bly017 %0 Journal Article %J Proceedings of the Royal Society B %D 2018 %T Ecological specialization is associated with genetic structure. %A Schär, S. %A Eastwood, R. %A Arnaldi, K.G. %A Talavera, G. %A Kaliszewska, Z. A. %A Boyle, J. H. %A Espeland, M. %A Nash, D. R. %A Vila, R. %A Pierce, N. E. %B Proceedings of the Royal Society B %G eng %0 Journal Article %J Florida Entomologist %D 2018 %T A first record of Anatrachyntis badia (Hodges 1962)(Lepidoptera: Cosmopterigidae) on Zamia integrifolia (Zamiaceae) %A Hua, C. %A Salzman, S. %A Pierce, N. E. %B Florida Entomologist %G eng %0 Journal Article %J Nature Communications %D 2018 %T The genetic basis of a social polymorphism in halictid bees. %A Kocher, S. D. %A Mallarino, R. %A Rubin, E.R. %A Yu, D. W. %A Hoekstra, H. E. %A Pierce, N. E. %B Nature Communications %G eng %0 Journal Article %J Genome Biology and Evolution %D 2018 %T Genome evolution of Bartonellaceae symbionts of ants at the opposite ends of the trophic scale. %A Bisch, G. %A Minna-Maria Neuvonen, M.M %A Pierce, N. E. %A Russell. J.A. %A Koga, R. %A Sanders, J. G. %A Łukasik, P. %A Andersson, S.G.E. %B Genome Biology and Evolution %G eng %0 Journal Article %J Nature Communications %D 2018 %T Herbivorous turtle ants obtain essential nutrients from a conserved nitrogen-recycling gut microbiome. %A Hu, Y. %A Sanders, J. G. %A Łukasik, P. %A D’Amelio, K. %A Millar, J.S. %A Vann, D.R. %A Lan, Y. %A Newton, J.A. %A Schotanus, M. %A Kronauer, D. J. C. %A Pierce, N. E. %A Moreau, C.S %A Wertz, J.T. %A Engel, P. %A Russell, J. A. %B Nature Communications %V 9 %G eng %N 964 %0 Journal Article %J mBio %D 2018 %T An Introduced Crop Plant Is Driving Diversification of theVirulent Bacterial Pathogen Erwinia tracheiphila %A Shapiro, L. R. %A Paulson, J.N. %A Arnold, B.J. %A Scully, E.D. %A Zhaxybayeva, O. %A Pierce, N. E. %A Rocha, J. %A Klepac-Ceraj, V. %A Holton, K. %A Kolter, R. %B mBio %V 9 %G eng %N 5 %0 Journal Article %J Biological Journal of the Linnaean Society %D 2018 %T Keeping an eye on coloration: Ecological correlates of the evolution of pitcher traits in the genus Nepenthes(Caryophyllales) %A Gilbert, K.J. %A Nitta, J.H. %A Talavera, G. %A Pierce, N. E. %B Biological Journal of the Linnaean Society %V 123 %P 321-337 %G eng %0 Journal Article %J Science %D 2018 %T Neonicotinoid exposure disrupts bumblebee nest behavior, social networks, and thermoregulation %A Crall, J. D. %A Switzer, C.M. %A Oppenheimer, R.L. %A Ford Versypt, A.N. %A Dey, B. %A Brown, A. %A Eyster, M. %A Guerin, C. %A Pierce, N. E. %A Combes, S. A. %A de Bivort, B.L. %B Science %V 362 %P 683-686 %G eng %N 6415 %0 Journal Article %J Molecular Phylogenetics and Evolution %D 2018 %T Phylogenetics of moth-like butterflies (Papilionoidea: Hedylidae) based on a new 13-locus target capture probe set. %A Kawahara, A. Y. %A Breinholt, J.W. %A Espeland, M. %A Storer, C. %A Plotkin,D. %A Dexter, K.M. %A Toussaint, E.F.A. %A St Laurent, R.A. %A Brehm, G. %A Vargas, S. %A Forero, D. %A Pierce, N. E. %A Lohman, D. J. %B Molecular Phylogenetics and Evolution %G eng %0 Journal Article %J Molecular Ecology %D 2018 %T Rhizosphere-associated Pseudomonas induce systemic resistance to herbivores at the cost of susceptibility to bacterial pathogens. %A Haney, C.H. %A Wiesmann, C.L. %A Shapiro, L. R. %A O’Sullivan, L. R. %A Khorasani, S. %A Melnyk, R.A. %A Xiao, L. %A Han, J. %A Bush, J. %A Carrillo, J. %A Pierce, N. E. %A Ausubel, F. M. %B Molecular Ecology %V 27 %P 1833-1847 %G eng %U http://doi-org.ezp-prod1.hul.harvard.edu/10.1111/mec.14400 %0 Journal Article %J Royal Society Open Science %D 2018 %T Social behavior in bees influences the abundance of Sodalis (Enterobacteriaceae) symbionts. %A Rubin, B.E. %A Sanders, J. G. %A Turner, K.M. %A Pierce, N. E. %A Kocher, S. D. %B Royal Society Open Science %G eng %0 Journal Article %J Nature Communications %D 2018 %T Spatial fidelity of workers predicts collective response to disturbance in a social insect %A Crall, J. D. %A Kocher, S. %A Oppenheimer, R.L. %A Gravish, N. %A Mountcastle, A.M. %A Pierce, N. E. %A Combes, S. A. %B Nature Communications %V 9 %G eng %N 1201 %0 Journal Article %J mBio %D 2018 %T An Introduced Crop Plant Is Driving Diversification of theVirulent Bacterial Pathogen Erwinia tracheiphila %A Shapiro, L. R. %A Paulson, J.N. %A Arnold, B.J. %A Scully, E.D. %A Zhaxybayeva, O. %A Pierce, N. E. %A Rocha, J. %A Klepac-Ceraj, V. %A Holton, K. %A Kolter, R. %B mBio %V 9 %G eng %N 5 %0 Journal Article %J Science %D 2018 %T Neonicotinoid exposure disrupts bumblebee nest behavior, social networks, and thermoregulation %A Crall, J. D. %A Switzer, C.M. %A Oppenheimer, R.L. %A Ford Versypt, A.N. %A Dey, B. %A Brown, A. %A Eyster, M. %A Guerin, C. %A Pierce, N. E. %A Combes, S. A. %A de Bivort, B.L. %B Science %V 362 %P 683-686 %G eng %N 6415 %0 Journal Article %J Proceedings of the Royal Society B %D 2018 %T Ecological specialization is associated with genetic structure. %A Schär, S. %A Eastwood, R. %A Arnaldi, K.G. %A Talavera, G. %A Kaliszewska, Z. A. %A Boyle, J. H. %A Espeland, M. %A Nash, D. R. %A Vila, R. %A Pierce, N. E. %B Proceedings of the Royal Society B %G eng %0 Journal Article %J Royal Society Open Science %D 2018 %T Social behavior in bees influences the abundance of Sodalis (Enterobacteriaceae) symbionts. %A Rubin, B.E. %A Sanders, J. G. %A Turner, K.M. %A Pierce, N. E. %A Kocher, S. D. %B Royal Society Open Science %G eng %0 Journal Article %J Genome Biology and Evolution %D 2018 %T Genome evolution of Bartonellaceae symbionts of ants at the opposite ends of the trophic scale. %A Bisch, G. %A Minna-Maria Neuvonen, M.M %A Pierce, N. E. %A Russell. J.A. %A Koga, R. %A Sanders, J. G. %A Łukasik, P. %A Andersson, S.G.E. %B Genome Biology and Evolution %G eng %0 Journal Article %J Nature Communications %D 2018 %T The genetic basis of a social polymorphism in halictid bees. %A Kocher, S. D. %A Mallarino, R. %A Rubin, E.R. %A Yu, D. W. %A Hoekstra, H. E. %A Pierce, N. E. %B Nature Communications %G eng %0 Journal Article %J Elife %D 2018 %T Convergence between the microcosms of Southeast Asian and North American pitcher plants. %A Bittleston, L. S. %A Wolock, C.J. %A Yahya, B.E. %A Chan, X.Y. %A Chan, K.G. %A Pierce, N. E. %A Pringle, A. %B Elife %G eng %0 Journal Article %J BMC Evolutionary Biology %D 2018 %T Anchored phylogenomics illuminates the skipper butterfly tree of life. %A Toussaint, E.F.A. %A Breinholt, J.W. %A Earl, C. %A Warren, A. D. %A Brower, J. V.Z. %A Yago, M. %A Dexter, K.M. %A Espeland, M. %A Pierce, N. E. %A Lohman, D. J. %A Kawahara, A. Y. %B BMC Evolutionary Biology %G eng %0 Journal Article %J Molecular Phylogenetics and Evolution %D 2018 %T Phylogenetics of moth-like butterflies (Papilionoidea: Hedylidae) based on a new 13-locus target capture probe set. %A Kawahara, A. Y. %A Breinholt, J.W. %A Espeland, M. %A Storer, C. %A Plotkin,D. %A Dexter, K.M. %A Toussaint, E.F.A. %A St Laurent, R.A. %A Brehm, G. %A Vargas, S. %A Forero, D. %A Pierce, N. E. %A Lohman, D. J. %B Molecular Phylogenetics and Evolution %G eng %0 Journal Article %J Florida Entomologist %D 2018 %T A first record of Anatrachyntis badia (Hodges 1962)(Lepidoptera: Cosmopterigidae) on Zamia integrifolia (Zamiaceae) %A Hua, C. %A Salzman, S. %A Pierce, N. E. %B Florida Entomologist %G eng %0 Journal Article %J Nature Communications %D 2018 %T Spatial fidelity of workers predicts collective response to disturbance in a social insect %A Crall, J. D. %A Kocher, S. %A Oppenheimer, R.L. %A Gravish, N. %A Mountcastle, A.M. %A Pierce, N. E. %A Combes, S. A. %B Nature Communications %V 9 %G eng %N 1201 %0 Journal Article %J Nature Communications %D 2018 %T Herbivorous turtle ants obtain essential nutrients from a conserved nitrogen-recycling gut microbiome. %A Hu, Y. %A Sanders, J. G. %A Łukasik, P. %A D’Amelio, K. %A Millar, J.S. %A Vann, D.R. %A Lan, Y. %A Newton, J.A. %A Schotanus, M. %A Kronauer, D. J. C. %A Pierce, N. E. %A Moreau, C.S %A Wertz, J.T. %A Engel, P. %A Russell, J. A. %B Nature Communications %V 9 %G eng %N 964 %0 Journal Article %J Current Biology %D 2018 %T Comprehensive higher-level phylogeny of butterflies (Papilionoidea) inferred from genomic data. %A Espeland, M. %A Breinholt, J. %A Willmott, K. R. %A Warren, A. D. %A Vila, R. %A Toussaint, E.F.A. %A Maunsell, S.C. %A Kwaku, A. P. %A Talavera, G. %A Eastwood, R. %A Jarzyna, M.A. %A Ries, L. %A Guralinick, R. %A Lohman, D. J. %A Pierce, N. E. %A Kawahara, A. Y. %B Current Biology %V 28 %P 770-778 %G eng %U http://doi.org/10.1016/jcub.2018.01.061 %0 Journal Article %J Biological Journal of the Linnaean Society %D 2018 %T Keeping an eye on coloration: Ecological correlates of the evolution of pitcher traits in the genus Nepenthes(Caryophyllales) %A Gilbert, K.J. %A Nitta, J.H. %A Talavera, G. %A Pierce, N. E. %B Biological Journal of the Linnaean Society %V 123 %P 321-337 %G eng %0 Journal Article %J Molecular Ecology %D 2018 %T Rhizosphere-associated Pseudomonas induce systemic resistance to herbivores at the cost of susceptibility to bacterial pathogens. %A Haney, C.H. %A Wiesmann, C.L. %A Shapiro, L. R. %A O’Sullivan, L. R. %A Khorasani, S. %A Melnyk, R.A. %A Xiao, L. %A Han, J. %A Bush, J. %A Carrillo, J. %A Pierce, N. E. %A Ausubel, F. M. %B Molecular Ecology %V 27 %P 1833-1847 %G eng %U http://doi-org.ezp-prod1.hul.harvard.edu/10.1111/mec.14400 %0 Journal Article %J Biological Journal of the Linnaean Society %D 2018 %T Cycad-feeding insects share a core gut microbiome. %A Salzman, S. %A Whitaker, M. %A Pierce, N. E. %B Biological Journal of the Linnaean Society %V 123 %P 728-738 %G eng %U http://doi-org.ezp-prod1.hul.harvard.edu/10.1093/biolinnean/bly017 %0 Journal Article %J Proceedings of the Royal Society B %D 2017 %T Distinctive fungal communities in an obligate African ant plant mutualism. %A Baker, C. C. M. %A Martins, D. J. %A Pelaez, J.N. %A Billen, J.P.J. %A Pringle, A. %A Frederickson, M. E. %A Pierce, N. E. %B Proceedings of the Royal Society B %V 284 %G eng %U http://dx.doi.org/10.1098/rspb.2016.2501 %N 20162501 %0 Journal Article %J Integrative and Comparative Biology %D 2017 %T Dramatic differences in gut bacterial densities help to explain the relationship between diet and habitat in rainforest ants. %A Sanders, J. G. %A Lukasik, P. %A Frederickson, M. E. %A Pierce, N. E. %B Integrative and Comparative Biology %V 57 %P 705-722 %G eng %0 Journal Article %J Zootaxa %D 2017 %T LepNet: The Lepidoptera of North America Network. %A Seltman, K.C. %A Cobb, N.S. %A Gall, L.F. %A Bartlett, C.R. %A Basham, M.A. %A Betancourt, I. %A Bills, C. %A Brandt, B. %A Brown, R.L. %A Bundy, C. %A Caterino, M. S. %B Zootaxa %V 4247 %P 73-77 %G eng %U http://dx.doi.org/10.11646/zootaxa.4247.1.10 %0 Journal Article %J Ecography %D 2017 %T Molecular substitution rate increases with latitude in butterflies: Evidence for a trans‐glacial latitudinal layering of populations? %A S Schär %A R Vila %A A Petrović %A Ž Tomanović %A Pierce, N. E. %A DR Nash %B Ecography %G eng %0 Journal Article %J Ecology and Evolution %D 2017 %T Polygyny cannot explain the superior competitive ability of dominant ant associates in the African ant plant Acacia(Vachellia) drepanolobium. %A Boyle, J. H. %A Martins, D. J. %A Peleaz, J. %A Musili, P.M. %A Kibet, S. %A Ndung’u, K.N. %A Kenfack, D. %A Pierce, N. E. %B Ecology and Evolution %V 8 %P 1441-1450 %G eng %0 Journal Article %J Proceedings of the National Academy of Science USA %D 2017 %T Solitary bees reduce investment in communication compared with their social relatives. %A Wittwer, B. %A Hefetz, A. %A Simon, T. %A Murphy, L.E.K. %A Elgar, M. A. %A Pierce, N. E. %A Kocher, Sarah D. %B Proceedings of the National Academy of Science USA %V 114 %P 6569–6574 %G eng %U http://www.pnas.org/content/114/25/6569.full %0 Journal Article %J Ecology and Evolution %D 2017 %T Polygyny cannot explain the superior competitive ability of dominant ant associates in the African ant plant Acacia(Vachellia) drepanolobium. %A Boyle, J. H. %A Martins, D. J. %A Peleaz, J. %A Musili, P.M. %A Kibet, S. %A Ndung’u, K.N. %A Kenfack, D. %A Pierce, N. E. %B Ecology and Evolution %V 8 %P 1441-1450 %G eng %0 Journal Article %J Integrative and Comparative Biology %D 2017 %T Dramatic differences in gut bacterial densities help to explain the relationship between diet and habitat in rainforest ants. %A Sanders, J. G. %A Lukasik, P. %A Frederickson, M. E. %A Pierce, N. E. %B Integrative and Comparative Biology %V 57 %P 705-722 %G eng %0 Journal Article %J Proceedings of the National Academy of Science USA %D 2017 %T Solitary bees reduce investment in communication compared with their social relatives. %A Wittwer, B. %A Hefetz, A. %A Simon, T. %A Murphy, L.E.K. %A Elgar, M. A. %A Pierce, N. E. %A Kocher, Sarah D. %B Proceedings of the National Academy of Science USA %V 114 %P 6569–6574 %G eng %U http://www.pnas.org/content/114/25/6569.full %0 Journal Article %J Proceedings of the Royal Society B %D 2017 %T Distinctive fungal communities in an obligate African ant plant mutualism. %A Baker, C. C. M. %A Martins, D. J. %A Pelaez, J.N. %A Billen, J.P.J. %A Pringle, A. %A Frederickson, M. E. %A Pierce, N. E. %B Proceedings of the Royal Society B %V 284 %G eng %U http://dx.doi.org/10.1098/rspb.2016.2501 %N 20162501 %0 Journal Article %J Zootaxa %D 2017 %T LepNet: The Lepidoptera of North America Network. %A Seltman, K.C. %A Cobb, N.S. %A Gall, L.F. %A Bartlett, C.R. %A Basham, M.A. %A Betancourt, I. %A Bills, C. %A Brandt, B. %A Brown, R.L. %A Bundy, C. %A Caterino, M. S. %B Zootaxa %V 4247 %P 73-77 %G eng %U http://dx.doi.org/10.11646/zootaxa.4247.1.10 %0 Journal Article %J Ecography %D 2017 %T Molecular substitution rate increases with latitude in butterflies: Evidence for a trans‐glacial latitudinal layering of populations? %A S Schär %A R Vila %A A Petrović %A Ž Tomanović %A Pierce, N. E. %A DR Nash %B Ecography %G eng %0 Journal Article %J Frontiers in Ecology and Evolution %D 2016 %T Colony Diet Influences Ant Worker Foraging and Attendance of Myrmecophilous Lycaenid Caterpillars %A S Pohl %A ME Frederickson %A Elgar, MA %A Pierce, N. E. %B Frontiers in Ecology and Evolution %V 4 %G eng %U http://journal.frontiersin.org/article/10.3389/fevo.2016.00114/full %N 114 %0 Journal Article %J Trends in Ecology & Evolution %D 2016 %T Convergence in Multispecies Interactions %A Bittleston, L. S. %A Pierce, N. E. %A Ellison, A.M. %A Pringle, A. %B Trends in Ecology & Evolution %V 31 %P 269-280 %8 April 2016 %G eng %N 4 %0 Journal Article %J Philosophical Transactions of the Royal Society B %D 2016 %T Dissecting host-associated communities with DNA barcodes %A CCM Baker %A Bittleston, LS %A Sanders, J. G. %A Pierce, N. E. %B Philosophical Transactions of the Royal Society B %V 371 %G eng %N 1702 %0 Journal Article %J The Science of Nature %D 2016 %T Eavesdropping on cooperative communication within an ant-butterfly mutualism %A Elgar, M. A. %A Nash, D. R. %A Pierce, N. E. %X

DOI 10.1007/s00114-016-1409-5 

%B The Science of Nature %V 103 %P 84 %G eng %U http://link.springer.com/article/10.1007/s00114-016-1409-5 %0 Journal Article %J Molecular Ecology %D 2016 %T Gut microbiota of dung beetles correspond to dietary specializations of adults and larvae %A SP Shukla %A Sanders, J. G. %A MJ Byrne %A Pierce, N. E. %B Molecular Ecology %V 25 %G eng %N 24 %0 Journal Article %J Frontiers in Microbiology %D 2016 %T Microbial communities of lycaenid butterflies do not correlate with larval diet %A MRL Whitaker %A S Salzman %A Sanders, J. %A M Kaltenpoth %A Pierce, N. E. %B Frontiers in Microbiology %V 7 %G eng %U https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5129467/ %N 1920 %0 Journal Article %J Invertebrate Systematics %D 2016 %T Phylogeny and population genetic structure of the ant genus Acropyga (Hymenoptera : Formicidae) in Papua New Guinea %A Janda, M.F. %A Matos-Maravi, P. %A Borovanska, M. %A Zima Jr, J. %A Youngerman, E. %A Pierce, N. E. %B Invertebrate Systematics %V 30 %P 28-40 %8 16 March 2016 %G eng %0 Journal Article %J J Insect Physiol %D 2016 %T Pseudomonas syringae enhances herbivory by suppressing the reactive oxygen burst in Arabidopsis %A Groen, S. C. %A Humphrey, P. T. %A Chevasco, D. %A Ausubel, F. M. %A Pierce, N. E. %A Whiteman, N. K. %X

Plant-herbivore interactions have evolved in the presence of plant-colonizing microbes. These microbes can have important third-party effects on herbivore ecology, as exemplified by drosophilid flies that evolved from ancestors feeding on plant-associated microbes. Leaf-mining flies in the genus Scaptomyza, which is nested within the paraphyletic genus Drosophila, show strong associations with bacteria in the genus Pseudomonas, including Pseudomonas syringae. Adult females are capable of vectoring these bacteria between plants and larvae show a preference for feeding on P. syringae-infected leaves. Here we show that Scaptomyza flava larvae can also vector P. syringae to and from feeding sites, and that they not only feed more, but also develop faster on plants previously infected with P. syringae. Our genetic and physiological data show that P. syringae enhances S. flava feeding on infected plants at least in part by suppressing anti-herbivore defenses mediated by reactive oxygen species.

%B J Insect Physiol %V 84 %P 90-102 %8 21 Jul 2015 %@ 1879-1611 (Electronic)0022-1910 (Linking) %G eng %M 26205072 %! Journal of insect physiologyJournal of insect physiology %0 Journal Article %J Molecular Ecology %D 2016 %T Gut microbiota of dung beetles correspond to dietary specializations of adults and larvae %A SP Shukla %A Sanders, J. G. %A MJ Byrne %A Pierce, N. E. %B Molecular Ecology %V 25 %G eng %N 24 %0 Journal Article %J Philosophical Transactions of the Royal Society B %D 2016 %T Dissecting host-associated communities with DNA barcodes %A CCM Baker %A Bittleston, LS %A Sanders, J. G. %A Pierce, N. E. %B Philosophical Transactions of the Royal Society B %V 371 %G eng %N 1702 %0 Journal Article %J Frontiers in Ecology and Evolution %D 2016 %T Colony Diet Influences Ant Worker Foraging and Attendance of Myrmecophilous Lycaenid Caterpillars %A S Pohl %A ME Frederickson %A Elgar, MA %A Pierce, N. E. %B Frontiers in Ecology and Evolution %V 4 %G eng %U http://journal.frontiersin.org/article/10.3389/fevo.2016.00114/full %N 114 %0 Journal Article %J Frontiers in Microbiology %D 2016 %T Microbial communities of lycaenid butterflies do not correlate with larval diet %A MRL Whitaker %A S Salzman %A Sanders, J. %A M Kaltenpoth %A Pierce, N. E. %B Frontiers in Microbiology %V 7 %G eng %U https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5129467/ %N 1920 %0 Journal Article %J The Science of Nature %D 2016 %T Eavesdropping on cooperative communication within an ant-butterfly mutualism %A Elgar, M. A. %A Nash, D. R. %A Pierce, N. E. %X

DOI 10.1007/s00114-016-1409-5 

%B The Science of Nature %V 103 %P 84 %G eng %U http://link.springer.com/article/10.1007/s00114-016-1409-5 %0 Journal Article %J Trends in Ecology & Evolution %D 2016 %T Convergence in Multispecies Interactions %A Bittleston, L. S. %A Pierce, N. E. %A Ellison, A.M. %A Pringle, A. %B Trends in Ecology & Evolution %V 31 %P 269-280 %8 April 2016 %G eng %N 4 %0 Journal Article %J Invertebrate Systematics %D 2016 %T Phylogeny and population genetic structure of the ant genus Acropyga (Hymenoptera : Formicidae) in Papua New Guinea %A Janda, M.F. %A Matos-Maravi, P. %A Borovanska, M. %A Zima Jr, J. %A Youngerman, E. %A Pierce, N. E. %B Invertebrate Systematics %V 30 %P 28-40 %8 16 March 2016 %G eng %0 Journal Article %J Biological Journal of the Linnean Society %D 2016 %T The setae of parasitic Liphyra brassolis butterfly larvae form a flexible armour for resisting attack by their ant hosts (Lycaenidae: Lepidoptera) %A Dupont, S.T. %A Zemeitat, D.S. %A Lohman, D. J. %A Pierce, N. E. %B Biological Journal of the Linnean Society %V 117 %P 607-619 %8 12 October 2015 %G eng %0 Journal Article %J J Insect Physiol %D 2016 %T Pseudomonas syringae enhances herbivory by suppressing the reactive oxygen burst in Arabidopsis %A Groen, S. C. %A Humphrey, P. T. %A Chevasco, D. %A Ausubel, F. M. %A Pierce, N. E. %A Whiteman, N. K. %X

Plant-herbivore interactions have evolved in the presence of plant-colonizing microbes. These microbes can have important third-party effects on herbivore ecology, as exemplified by drosophilid flies that evolved from ancestors feeding on plant-associated microbes. Leaf-mining flies in the genus Scaptomyza, which is nested within the paraphyletic genus Drosophila, show strong associations with bacteria in the genus Pseudomonas, including Pseudomonas syringae. Adult females are capable of vectoring these bacteria between plants and larvae show a preference for feeding on P. syringae-infected leaves. Here we show that Scaptomyza flava larvae can also vector P. syringae to and from feeding sites, and that they not only feed more, but also develop faster on plants previously infected with P. syringae. Our genetic and physiological data show that P. syringae enhances S. flava feeding on infected plants at least in part by suppressing anti-herbivore defenses mediated by reactive oxygen species.

%B J Insect Physiol %V 84 %P 90-102 %8 21 Jul 2015 %@ 1879-1611 (Electronic)0022-1910 (Linking) %G eng %M 26205072 %! Journal of insect physiologyJournal of insect physiology %0 Journal Article %J Molecular Phylogenetics and Evolution %D 2015 %T Ancient Neotropical origin and recent recolonisation: Phylogeny, biogeography and diversification of the Riodinidae (Lepidoptera: Papilionoidea) %A Espeland, M. %A Hall, J. P. W. %A DeVries, P. J. %A Lees, D. C. %A Cornwall, M. %A Hsu, Y. F. %A Wu, L. W. %A Campbell, D. L. %A Talavera, G. %A Vila, R. %A Salzman, S. %A Ruehr, S. %A Lohman, D. J. %A Pierce, N. E. %K biogeography %K Dispersal %K diversification %K dominican amber %K Evolution %K higher-level phylogeny %K hostplant %K larval %K life-history %K maximum %K metalmark butterflies %K mimetic ithomiine butterflies %K models %K nymphidiini %K revision %K styx %X

We present the first dated higher-level phylogenetic and biogeographic analysis of the butterfly family Riodinidae. This family is distributed worldwide, but more than 90% of the c. 1500 species are found in the Neotropics, while the c. 120 Old World species are concentrated in the Southeast Asian tropics, with minor Afrotropical and Australasian tropical radiations, and few temperate species. Morphologically based higher classification is partly unresolved, with genera incompletely assigned to tribes. Using 3666 bp from one mitochondrial and four nuclear markers for each of 23 outgroups and 178 riodinid taxa representing all subfamilies, tribes and subtribes, and 98 out of 145 described genera of riodinids, we estimate that Riodinidae split from Lycaenidae about 96 Mya in the mid-Cretaceous and started to diversify about 81 Mya. The Riodinidae are monophyletic and originated in the Neotropics, most likely in lowland proto-Amazonia. Neither the subfamily Euselasiinae nor the Nemeobiinae are monophyletic as currently constituted. The enigmatic, monotypic Neotropical genera Styx and Corrachia (most recently treated in Euselasiinae: Corrachiini) are highly supported as derived taxa in the Old World Nemeobiinae, with dispersal most likely occurring across the Beringia land bridge during the Oligocene. Styx and Corrachia, together with all other nemeobiines, are the only exclusively Primulaceae-feeding riodinids. The steadily increasing proliferation of the Neotropical Riodininae subfamily contrasts with the decrease in diversification in the Old World, and may provide insights into factors influencing the diversification rate of this relatively ancient clade of Neotropical insects. (C) 2015 Elsevier Inc. All rights reserved.

%B Molecular Phylogenetics and Evolution %V 93 %P 296-306 %8 Dec %@ 1055-7903 %G English %M WOS:000362613200027 %! Mol Phylogenet EvolMol Phylogenet Evol %0 Journal Article %J Genome Announcements %D 2015 %T Draft genome sequence ofErwinia tracheiphila, an economically important bacterial pathogen of cucurbits. %A Shapiro, LR %A ED Scully %A D. Roberts %A TJ Straub %A SM Geib %A J Park %A A Stephenson %A ES Rojas %A Liu, Q. %A G Beattie %A M. Gleason %A CM De Moraes %A MC Mescher %A SG Fleischer %A Kolter, R. %A Pierce, N. E. %A O Zhaxybayeva %X

Erwinia tracheiphila is one of the most economically important pathogens of cucumbers, melons, squashes, pumpkins, and gourds in the northeastern and midwestern United States, yet its molecular pathology remains uninvestigated. Here, we report the first draft genome sequence of an E. tracheiphila strain isolated from an infected wild gourd (Cucurbita pepo subsp. texana) plant. The genome assembly consists of 7 contigs and includes a putative plasmid and at least 20 phage and prophage elements.

%B Genome Announcements %V 3 %P e00482-15 %G eng %N 3 %0 Journal Article %J Austral Ecology %D 2015 %T Metabarcoding as a tool for investigating arthropod diversity inNepenthespitcher plants. %A Bittleston, LS %A CCM Baker %A LB Strominger %A Pringle, A. %A Pierce, N. E. %X

The biodiversity of tropical forests consists primarily of small organisms that are difficult to detect and characterize. Next-generation sequencing (NGS) methods can facilitate analyses of these arthropod and microbial communities, leading to a better understanding of existing diversity and factors influencing community assembly. The pitchers of carnivorous pitcher plants often house surprisingly discrete communities and provide ideal systems for analysis using an NGS approach. The plants digest insects in order to access essential nutrients while growing in poor soils; however, the pitchers are also home to communities of living organisms, called inquilines. Certain arthropods appear to have coevolved with their pitcher plant hosts and are not found in other environments. We used Illumina amplicon sequencing of 18S rDNA to characterize the eukaryotes in three species of Nepenthes (Nepenthaceae) pitcher plants – N. gracilisN. rafflesiana and N. ampullaria – in each of three different parks in Singapore. The data reveal an unexpected diversity of eukaryotes, significant differences in community diversity among host species, variation in host specificity of inquilines and the presence of gregarine parasites. Counts of whole inquiline arthropods from the first collection year were roughly correlated with scaled 18S sequence abundances, indicating that amplicon sequencing is an effective means of gauging community structure. We barcoded a subset of the dipteran larvae using COI primers, and the resulting phylogenetic tree is mostly congruent with that found using the 18S locus, with the exception of one of five morphospecies. For many 18S and COI sequences, the best BLASTn matches showed low sequence identity, illustrating the need for better databases of Southeast Asian dipterans. Finally, networks of core arthropods and their host species were used to investigate degree of host specificity across multiple hosts, and this revealed significant specialization of certain arthropod fauna.

%B Austral Ecology %G eng %0 Journal Article %J Systematic Entomology %D 2015 %T Phylogeny of the Aphnaeinae: myrmecophilous African butterflies with carnivorous and herbivorous life histories %A Boyle, J. H. %A Kaliszewska, Z. A. %A Espeland, M. %A Suderman, T. R. %A Fleming, J. %A Heath, A. %A Pierce, N. E. %K Evolution %K inference %K lepidoptera %K lycaenidae %K maximum-likelihood %K mitochondrial %K mixed models %K molecular-data %K moths %K sequences %X

The Aphnaeinae (Lepidoptera: Lycaenidae) are a largely African subfamily of 278 described species that exhibit extraordinary life-history variation. The larvae of these butterflies typically form mutualistic associations with ants, and feed on a wide variety of plants, including 23 families in 19 orders. However, at least one species in each of 9 of the 17 genera is aphytophagous, parasitically feeding on the eggs, brood or regurgitations of ants. This diversity in diet and type of symbiotic association makes the phylogenetic relations of the Aphnaeinae of particular interest. A phylogenetic hypothesis for the Aphnaeinae was inferred from 4.4kb covering the mitochondrial marker COI and five nuclear markers (wg, H3, CAD, GAPDH and EF1) for each of 79 ingroup taxa representing 15 of the 17 currently recognized genera, as well as three outgroup taxa. Maximum Parsimony, Maximum Likelihood and Bayesian Inference analyses all support Heath's systematic revision of the clade based on morphological characters. Ancestral range inference suggests an African origin for the subfamily with a single dispersal into Asia. The common ancestor of the aphnaeines likely associated with myrmicine ants in the genus Crematogaster and plants of the order Fabales.

%B Systematic Entomology %V 40 %P 169-182 %8 Jan %@ 0307-6970 %G English %M WOS:000347453400010 %! Syst EntomolSyst Entomol %0 Journal Article %J Molecular Phylogenetics and Evolution %D 2015 %T Ancient Neotropical origin and recent recolonisation: Phylogeny, biogeography and diversification of the Riodinidae (Lepidoptera: Papilionoidea) %A Espeland, M. %A Hall, J. P. W. %A DeVries, P. J. %A Lees, D. C. %A Cornwall, M. %A Hsu, Y. F. %A Wu, L. W. %A Campbell, D. L. %A Talavera, G. %A Vila, R. %A Salzman, S. %A Ruehr, S. %A Lohman, D. J. %A Pierce, N. E. %K biogeography %K Dispersal %K diversification %K dominican amber %K Evolution %K higher-level phylogeny %K hostplant %K larval %K life-history %K maximum %K metalmark butterflies %K mimetic ithomiine butterflies %K models %K nymphidiini %K revision %K styx %X

We present the first dated higher-level phylogenetic and biogeographic analysis of the butterfly family Riodinidae. This family is distributed worldwide, but more than 90% of the c. 1500 species are found in the Neotropics, while the c. 120 Old World species are concentrated in the Southeast Asian tropics, with minor Afrotropical and Australasian tropical radiations, and few temperate species. Morphologically based higher classification is partly unresolved, with genera incompletely assigned to tribes. Using 3666 bp from one mitochondrial and four nuclear markers for each of 23 outgroups and 178 riodinid taxa representing all subfamilies, tribes and subtribes, and 98 out of 145 described genera of riodinids, we estimate that Riodinidae split from Lycaenidae about 96 Mya in the mid-Cretaceous and started to diversify about 81 Mya. The Riodinidae are monophyletic and originated in the Neotropics, most likely in lowland proto-Amazonia. Neither the subfamily Euselasiinae nor the Nemeobiinae are monophyletic as currently constituted. The enigmatic, monotypic Neotropical genera Styx and Corrachia (most recently treated in Euselasiinae: Corrachiini) are highly supported as derived taxa in the Old World Nemeobiinae, with dispersal most likely occurring across the Beringia land bridge during the Oligocene. Styx and Corrachia, together with all other nemeobiines, are the only exclusively Primulaceae-feeding riodinids. The steadily increasing proliferation of the Neotropical Riodininae subfamily contrasts with the decrease in diversification in the Old World, and may provide insights into factors influencing the diversification rate of this relatively ancient clade of Neotropical insects. (C) 2015 Elsevier Inc. All rights reserved.

%B Molecular Phylogenetics and Evolution %V 93 %P 296-306 %8 Dec %@ 1055-7903 %G English %M WOS:000362613200027 %! Mol Phylogenet EvolMol Phylogenet Evol %0 Journal Article %J Genome Announcements %D 2015 %T Draft genome sequence ofErwinia tracheiphila, an economically important bacterial pathogen of cucurbits. %A Shapiro, LR %A ED Scully %A D. Roberts %A TJ Straub %A SM Geib %A J Park %A A Stephenson %A ES Rojas %A Liu, Q. %A G Beattie %A M. Gleason %A CM De Moraes %A MC Mescher %A SG Fleischer %A Kolter, R. %A Pierce, N. E. %A O Zhaxybayeva %X

Erwinia tracheiphila is one of the most economically important pathogens of cucumbers, melons, squashes, pumpkins, and gourds in the northeastern and midwestern United States, yet its molecular pathology remains uninvestigated. Here, we report the first draft genome sequence of an E. tracheiphila strain isolated from an infected wild gourd (Cucurbita pepo subsp. texana) plant. The genome assembly consists of 7 contigs and includes a putative plasmid and at least 20 phage and prophage elements.

%B Genome Announcements %V 3 %P e00482-15 %G eng %N 3 %0 Journal Article %J Current Biology %D 2015 %T Lycaenid Caterpillar Secretions Manipulate Attendant Ant Behavior %A Hojo, M. K. %A Pierce, N. E. %A Tsuji, K. %K butterflies %K cocaine %K dopamine %K Drosophila %K Ecology %K lepidoptera %K mutualism %K nectar %K Parasite %X

Mutualistic interactions typically involve the exchange of different commodities between species [1]. Nutritious secretions are produced by a number of insects and plants in exchange for services such as defense [2, 3]. These rewards are valuable metabolically and can be used to reinforce the behavior of symbiotic partners that can learn and remember them effectively [4, 5]. We show here novel effects of insect exocrine secretions produced by caterpillars in modulating the behavior of attendant ants in the food-for-defense interaction between lycaenid butterflies and ants [6]. Reward secretions from the dorsal nectary organ (DNO) of Narathura japonica caterpillars function to reduce the locomotory activities of their attendant ants, Pristomyrmex punctatus workers. Moreover, workers that feed from caterpillar secretions are significantly more likely to show aggressive responses to eversion of the tentacle organs of the caterpillars. Analysis of the neurogenic amines in the brains of workers that consumed caterpillar secretions showed a significant decrease in levels of dopamine compared with controls. Experimental treatments in which reserpine, a known inhibitor of dopamine in Drosophila, was fed to workers similarly reduced their locomotory activity. We conclude that DNO secretions of lycaenid caterpillars can manipulate attendant ant behavior by altering dopaminergic regulation and increasing partner fidelity. Unless manipulated ants also receive a net nutritional benefit from DNO secretions, this suggests that similar reward-for-defense interactions that have been traditionally considered to be mutualisms may in fact be parasitic in nature.

%B Current Biology %V 25 %P 2260-2264 %8 Aug 31 %@ 0960-9822 %G English %M WOS:000360711700023 %! Curr BiolCurr Biol %0 Journal Article %J Austral Ecology %D 2015 %T Metabarcoding as a tool for investigating arthropod diversity inNepenthespitcher plants. %A Bittleston, LS %A CCM Baker %A LB Strominger %A Pringle, A. %A Pierce, N. E. %X

The biodiversity of tropical forests consists primarily of small organisms that are difficult to detect and characterize. Next-generation sequencing (NGS) methods can facilitate analyses of these arthropod and microbial communities, leading to a better understanding of existing diversity and factors influencing community assembly. The pitchers of carnivorous pitcher plants often house surprisingly discrete communities and provide ideal systems for analysis using an NGS approach. The plants digest insects in order to access essential nutrients while growing in poor soils; however, the pitchers are also home to communities of living organisms, called inquilines. Certain arthropods appear to have coevolved with their pitcher plant hosts and are not found in other environments. We used Illumina amplicon sequencing of 18S rDNA to characterize the eukaryotes in three species of Nepenthes (Nepenthaceae) pitcher plants – N. gracilisN. rafflesiana and N. ampullaria – in each of three different parks in Singapore. The data reveal an unexpected diversity of eukaryotes, significant differences in community diversity among host species, variation in host specificity of inquilines and the presence of gregarine parasites. Counts of whole inquiline arthropods from the first collection year were roughly correlated with scaled 18S sequence abundances, indicating that amplicon sequencing is an effective means of gauging community structure. We barcoded a subset of the dipteran larvae using COI primers, and the resulting phylogenetic tree is mostly congruent with that found using the 18S locus, with the exception of one of five morphospecies. For many 18S and COI sequences, the best BLASTn matches showed low sequence identity, illustrating the need for better databases of Southeast Asian dipterans. Finally, networks of core arthropods and their host species were used to investigate degree of host specificity across multiple hosts, and this revealed significant specialization of certain arthropod fauna.

%B Austral Ecology %G eng %0 Journal Article %J Systematic Entomology %D 2015 %T Phylogeny of the Aphnaeinae: myrmecophilous African butterflies with carnivorous and herbivorous life histories %A Boyle, J. H. %A Kaliszewska, Z. A. %A Espeland, M. %A Suderman, T. R. %A Fleming, J. %A Heath, A. %A Pierce, N. E. %K Evolution %K inference %K lepidoptera %K lycaenidae %K maximum-likelihood %K mitochondrial %K mixed models %K molecular-data %K moths %K sequences %X

The Aphnaeinae (Lepidoptera: Lycaenidae) are a largely African subfamily of 278 described species that exhibit extraordinary life-history variation. The larvae of these butterflies typically form mutualistic associations with ants, and feed on a wide variety of plants, including 23 families in 19 orders. However, at least one species in each of 9 of the 17 genera is aphytophagous, parasitically feeding on the eggs, brood or regurgitations of ants. This diversity in diet and type of symbiotic association makes the phylogenetic relations of the Aphnaeinae of particular interest. A phylogenetic hypothesis for the Aphnaeinae was inferred from 4.4kb covering the mitochondrial marker COI and five nuclear markers (wg, H3, CAD, GAPDH and EF1) for each of 79 ingroup taxa representing 15 of the 17 currently recognized genera, as well as three outgroup taxa. Maximum Parsimony, Maximum Likelihood and Bayesian Inference analyses all support Heath's systematic revision of the clade based on morphological characters. Ancestral range inference suggests an African origin for the subfamily with a single dispersal into Asia. The common ancestor of the aphnaeines likely associated with myrmicine ants in the genus Crematogaster and plants of the order Fabales.

%B Systematic Entomology %V 40 %P 169-182 %8 Jan %@ 0307-6970 %G English %M WOS:000347453400010 %! Syst EntomolSyst Entomol %0 Journal Article %J Evolution %D 2015 %T When caterpillars attack: Biogeography and life history evolution of the Miletinae (Lepidoptera: Lycaenidae) %A Kaliszewska, Z. A. %A Lohman, D. J. %A Sommer, K. %A Adelson, G. %A Rand, D. B. %A Mathew, J. %A Talavera, G. %A Pierce, N. E. %K Africa %K ant association %K ants %K aphytophagy %K butterflies %K coevolution %K Dispersal %K diversification %K feniseca-tarquinius %K geographic range %K molecular evolution %K myrmecophagy %K myrmecophily %K natural-history %K phylogenetic inference %K social parasitism %X

Of the four most diverse insect orders, Lepidoptera contains remarkably few predatory and parasitic species. Although species with these habits have evolved multiple times in moths and butterflies, they have rarely been associated with diversification. The wholly aphytophagous subfamily Miletinae (Lycaenidae) is an exception, consisting of nearly 190 species distributed primarily throughout the Old World tropics and subtropics. Most miletines eat Hemiptera, although some consume ant brood or are fed by ant trophallaxis. A well-resolved phylogeny inferred using 4915 bp from seven markers sampled from representatives of all genera and nearly one-third the described species was used to examine the biogeography and evolution of biotic associations in this group. Biogeographic analyses indicate that Miletinae likely diverged from an African ancestor near the start of the Eocene, and four lineages dispersed between Africa and Asia. Phylogenetic constraint in prey selection is apparent at two levels: related miletine species are more likely to feed on related Hemiptera, and related miletines are more likely to associate with related ants, either directly by eating the ants, or indirectly by eating hemipteran prey that are attended by those ants. These results suggest that adaptations for host ant location by ovipositing female miletines may have been retained from phytophagous ancestors that associated with ants mutualistically.

%B Evolution %V 69 %P 571-588 %8 Mar %@ 0014-3820 %G English %M WOS:000351446000001 %! EvolutionEvolution %0 Journal Article %J Conservation Genetics Resources %D 2014 %T Development and characterization of twenty-two polymorphic microsatellite markers for the leafcutter ant, Acromyrmex lundii, utilizing Illumina sequencing %A Rabeling, C. %A Bollazzi, M. %A Bacci, M. %A Beasley, R. R. %A Lance, S. L. %A Jones, K. L. %A Pierce, N. E. %K acromyrmex %K attini %K fungus-growing ants %K Illumina %K leafcutter ants %K microsatellites %K pal_finder %K pcr primers %K polyandry %K polygyny %K social parasitism %K ssr %X

We isolated and characterized a total of 22 microsatellite loci for the leafcutter ant, Acromyrmex lundii. The loci were screened for 24 individuals from southern Brazil and Uruguay. The number of alleles per locus ranged from 5 to 20, the observed heterozygosity ranged from 0.417 to 0.917, and the probability of identity values ranged from 0.011 to 0.38. These genetic markers will be useful for understanding the population and conservation biology of the leafcutter ant A. lundii and closely related species, and will provide novel insights into the evolutionary biology of social parasitism and leafcutter ant mating systems.

%B Conservation Genetics Resources %V 6 %P 319-322 %8 Jun %@ 1877-7252 %G English %M WOS:000335959600019 %! Conserv Genet ResourConserv Genet Resour %0 Journal Article %J Conservation Genetics Resources %D 2014 %T Development of twenty-one polymorphic microsatellite markers for the fungus-growing ant, Mycocepurus goeldii (Formicidae: Attini), using Illumina paired-end genomic sequencing %A Rabeling, C. %A Love, C. N. %A Lance, S. L. %A Jones, K. L. %A Pierce, N. E. %A Bacci, M. %K hymenoptera-formicidae %K inquilinism %K pal_finder %K polyandry %K polygyny %K social parasitism %K ssr %X

Obligate social parasites, or inquilines, exploit the colonies of free-living social species and evolved at least 80 times in ants alone. Most species of the highly specialized inquiline social parasites are rare, only known from one or very few, geographically isolated populations, and the sexual offspring of most inquiline species mates inside the maternal colony. Therefore, inquiline populations are believed to be small and genetically homogeneous due to inbreeding. To comparatively study the genetic diversity of the socially parasitic fungus-growing ant, Mycocepurus castrator, and its only known host species, Mycocepurus goeldii, and to infer the parasite's conservation status, we developed 21 microsatellite markers for the host species, M. goeldii, and evaluated whether these markers cross-amplify in the social parasite, M. castrator. We isolated and characterized a total of 21 microsatellite loci for M. goeldii. The loci were screened for 24 individuals from geographically distant and genetically divergent populations in Brazil. The number of alleles per locus ranged from 18 to 4, the observed heterozygosity ranged from 0.25 to 0.636, and the probability of identity values ranged from 0.011 to 0.146. Preliminary analyses show that these markers cross amplify in the closely related social parasite species M. castrator. These newly developed loci provide tools for studying the genetic diversity and the evolution of social parasitism in the Mycocepurus host-parasite system.

%B Conservation Genetics Resources %V 6 %P 739-741 %8 Sep %@ 1877-7252 %G English %M WOS:000340404900066 %! Conserv Genet ResourConserv Genet Resour %0 Journal Article %J Zoologica Scripta %D 2014 %T Revised systematics and higher classification of pierid butterflies (Lepidoptera: Pieridae) based on molecular data %A Wahlberg, N. %A Rota, J. %A Braby, M. F. %A Pierce, N. E. %A Wheat, C. W. %K biogeography %K delias hubner lepidoptera %K diversification %K Evolution %K genera lepidoptera %K immature stages %K larval food plants %K neotropical mistletoe butterflies %K origin %K phylogenetic inference %X

The butterfly family Pieridae comprises approximately 1000 described species placed in 85 genera, but the higher classification has not yet been settled. We used molecular data from eight gene regions (one mitochondrial and seven nuclear protein-coding genes) comprising a total of similar to 6700bp from 96 taxa to infer a well-supported phylogenetic hypothesis for the family. Based on this hypothesis, we revise the higher classification for all pierid genera. We resurrect the tribe Teracolini stat. rev. in the subfamily Pierinae to include the genera Teracolus, Pinacopteryx, Gideona, Ixias, Eronia, Colotis and most likely Calopieris. We transfer Hebomoia to the tribe Anthocharidini and assign the previously unplaced genera Belenois and Dixeia to the subtribe Aporiina. Three lineages near the base of Pierinae (Leptosia, Elodina and Nepheronia + Pareronia) remain unplaced. For each of these, we describe and delineate new tribes: Elodinini Braby tribus nova, Leptosiaini Braby tribus nova and Nepheroniini Braby tribus nova. The proposed higher classification is based on well-supported monophyletic groups and is likely to remain stable even with the addition of more data.

%B Zoologica Scripta %V 43 %P 641-650 %8 Nov %@ 0300-3256 %G English %M WOS:000343866100007 %! Zool ScrZool Scr %0 Journal Article %J Current Biology %D 2014 %T A Social Parasite Evolved Reproductive Isolation from Its Fungus-Growing Ant Host in Sympatry %A Rabeling, C. %A Schultz, T. R. %A Pierce, N. E. %A Bacci, M. %K alternative adaptations %K Evolution %K hymenoptera-formicidae %K inquiline %K mycocepurus-smithii %K phylogenetic analysis %K sequences %K Speciation %K species trees %K x-linked genes %X

Inquiline social parasitic ant species exploit colonies of other ant species mainly by producing sexual offspring that are raised by the host. Ant social parasites and their hosts are often close relatives (Emery's rule), and two main hypotheses compete to explain the parasites' evolutionary origins: (1) the interspecific hypothesis proposes an allopatric speciation scenario for the parasite, whereas (2) the intraspecific hypothesis postulates that the parasite evolves directly from its host in sympatry [1-10]. Evidence in support of the intraspecific hypothesis has been accumulating for ants [3, 5, 7, 9-12], but sympatric speciation remains controversial as a general speciation mechanism for inquiline parasites. Here we use molecular phylogenetics to assess whether the socially parasitic fungus-growing ant Mycocepurus castrator speciated from its host Mycocepurus goeldii in sympatry. Based on differing patterns of relationship in mitochondrial and individual nuclear genes, we conclude that host and parasite occupy a temporal window in which lineage sorting has taken place in the mitochondrial genes but not yet in the nuclear alleles. We infer that the host originated first and that the parasite originated subsequently from a subset of the host species' populations, providing empirical support for the hypothesis that inquiline parasites can evolve reproductive isolation while living sympatrically with their hosts.

%B Current Biology %V 24 %P 2047-2052 %8 Sep 8 %@ 0960-9822 %G English %M WOS:000341541300031 %! Curr BiolCurr Biol %0 Journal Article %J Molecular Ecology %D 2014 %T Stability and phylogenetic correlation in gut microbiota: lessons from ants and apes %A Sanders, J. G. %A Powell, S. %A Kronauer, D. J. %A Vasconcelos, H. L. %A Frederickson, M. E. %A Pierce, N. E. %K *Microbiota %K *Phylogeny %K Animals %K Ants/genetics/*microbiology %K Biodiversity %K Biological Evolution %K Cluster Analysis %K Digestive System/*microbiology %K DNA, Bacterial/genetics %K Hominidae/microbiology %K RNA, Ribosomal, 16S/genetics %K Sequence Analysis, DNA %X

Correlation between gut microbiota and host phylogeny could reflect codiversification over shared evolutionary history or a selective environment that is more similar in related hosts. These alternatives imply substantial differences in the relationship between host and symbiont, but can they be distinguished based on patterns in the community data themselves? We explored patterns of phylogenetic correlation in the distribution of gut bacteria among species of turtle ants (genus Cephalotes), which host a dense gut microbial community. We used 16S rRNA pyrosequencing from 25 Cephalotes species to show that their gut community is remarkably stable, from the colony to the genus level. Despite this overall similarity, the existing differences among species' microbiota significantly correlated with host phylogeny. We introduced a novel analytical technique to test whether these phylogenetic correlations are derived from recent bacterial evolution, as would be expected in the case of codiversification, or from broader shifts more likely to reflect environmental filters imposed by factors such as diet or habitat. We also tested this technique on a published data set of ape microbiota, confirming earlier results while revealing previously undescribed patterns of phylogenetic correlation. Our results indicated a high degree of partner fidelity in the Cephalotes microbiota, suggesting that vertical transmission of the entire community could play an important role in the evolution and maintenance of the association. As additional comparative microbiota data become available, the techniques presented here can be used to explore trends in the evolution of host-associated microbial communities.

%B Molecular Ecology %V 23 %P 1268-83 %8 Mar %@ 1365-294X (Electronic)0962-1083 (Linking) %G eng %M 24304129 %! Mol EcolMol Ecol %0 Journal Article %J Proceedings of the Royal Society B-Biological Sciences %D 2014 %T Transitions in social complexity along elevational gradients reveal a combined impact of season length and development time on social evolution %A Kocher, S. D. %A Pellissier, L. %A Veller, C. %A Purcell, J. %A Nowak, M. A. %A Chapuisat, M. %A Pierce, N. E. %K Altitude %K bumble bees %K development time %K eusociality %K halictidae %K hymenoptera %K hypothesis %K Insects %K mountain passes %K polistes %K Population %K social behaviour %K solitary %K species distributions %X

Eusociality is taxonomically rare, yet associated with great ecological success. Surprisingly, studies of environmental conditions favouring eusociality are often contradictory. Harsh conditions associated with increasing altitude and latitude seem to favour increased sociality in bumblebees and ants, but the reverse pattern is found in halictid bees and polistine wasps. Here, we compare the life histories and distributions of populations of 176 species of Hymenoptera from the Swiss Alps. We show that differences in altitudinal distributions and development times among social forms can explain these contrasting patterns: highly social taxa develop more quickly than intermediate social taxa, and are thus able to complete the reproductive cycle in shorter seasons at higher elevations. This dual impact of altitude and development time on sociality illustrates that ecological constraints can elicit dynamic shifts in behaviour, and helps explain the complex distribution of sociality across ecological gradients.

%B Proceedings of the Royal Society B-Biological Sciences %V 281 %8 Jul 22 %@ 0962-8452 %G English %M WOS:000338335800019 %! P Roy Soc B-Biol SciP Roy Soc B-Biol Sci %0 Journal Article %J Conservation Genetics Resources %D 2014 %T Development and characterization of twenty-two polymorphic microsatellite markers for the leafcutter ant, Acromyrmex lundii, utilizing Illumina sequencing %A Rabeling, C. %A Bollazzi, M. %A Bacci, M. %A Beasley, R. R. %A Lance, S. L. %A Jones, K. L. %A Pierce, N. E. %K acromyrmex %K attini %K fungus-growing ants %K Illumina %K leafcutter ants %K microsatellites %K pal_finder %K pcr primers %K polyandry %K polygyny %K social parasitism %K ssr %X

We isolated and characterized a total of 22 microsatellite loci for the leafcutter ant, Acromyrmex lundii. The loci were screened for 24 individuals from southern Brazil and Uruguay. The number of alleles per locus ranged from 5 to 20, the observed heterozygosity ranged from 0.417 to 0.917, and the probability of identity values ranged from 0.011 to 0.38. These genetic markers will be useful for understanding the population and conservation biology of the leafcutter ant A. lundii and closely related species, and will provide novel insights into the evolutionary biology of social parasitism and leafcutter ant mating systems.

%B Conservation Genetics Resources %V 6 %P 319-322 %8 Jun %@ 1877-7252 %G English %M WOS:000335959600019 %! Conserv Genet ResourConserv Genet Resour %0 Journal Article %J Conservation Genetics Resources %D 2014 %T Development of twenty-one polymorphic microsatellite markers for the fungus-growing ant, Mycocepurus goeldii (Formicidae: Attini), using Illumina paired-end genomic sequencing %A Rabeling, C. %A Love, C. N. %A Lance, S. L. %A Jones, K. L. %A Pierce, N. E. %A Bacci, M. %K hymenoptera-formicidae %K inquilinism %K pal_finder %K polyandry %K polygyny %K social parasitism %K ssr %X

Obligate social parasites, or inquilines, exploit the colonies of free-living social species and evolved at least 80 times in ants alone. Most species of the highly specialized inquiline social parasites are rare, only known from one or very few, geographically isolated populations, and the sexual offspring of most inquiline species mates inside the maternal colony. Therefore, inquiline populations are believed to be small and genetically homogeneous due to inbreeding. To comparatively study the genetic diversity of the socially parasitic fungus-growing ant, Mycocepurus castrator, and its only known host species, Mycocepurus goeldii, and to infer the parasite's conservation status, we developed 21 microsatellite markers for the host species, M. goeldii, and evaluated whether these markers cross-amplify in the social parasite, M. castrator. We isolated and characterized a total of 21 microsatellite loci for M. goeldii. The loci were screened for 24 individuals from geographically distant and genetically divergent populations in Brazil. The number of alleles per locus ranged from 18 to 4, the observed heterozygosity ranged from 0.25 to 0.636, and the probability of identity values ranged from 0.011 to 0.146. Preliminary analyses show that these markers cross amplify in the closely related social parasite species M. castrator. These newly developed loci provide tools for studying the genetic diversity and the evolution of social parasitism in the Mycocepurus host-parasite system.

%B Conservation Genetics Resources %V 6 %P 739-741 %8 Sep %@ 1877-7252 %G English %M WOS:000340404900066 %! Conserv Genet ResourConserv Genet Resour %0 Journal Article %J Journal of Evolutionary Biology %D 2014 %T Renewed diversification is associated with new ecological opportunity in the Neotropical turtle ants %A Price, S. L. %A Powell, S. %A Kronauer, D. J. C. %A Tran, L. A. P. %A Pierce, N. E. %A Wayne, R. K. %K adaptive radiation %K biogeography %K brazilian cerrado %K cephalotes %K chacoan %K community phylogenetics %K continental radiation %K dominican amber %K extinction rates %K molecular phylogenies %K morphological evolution %K Phylogeny %K plant diversity %K radiation %K range evolution %K Speciation %K species-diversity %X

Ecological opportunity, defined as access to new resources free from competitors, is thought to be a catalyst for the process of adaptive radiation. Much of what we know about ecological opportunity, and the larger process of adaptive radiation, is derived from vertebrate diversification on islands. Here, we examine lineage diversification in the turtle ants (Cephalotes), a species-rich group of ants that has diversified throughout the Neotropics. We show that crown group turtle ants originated during the Eocene (around 46 mya), coincident with global warming and the origin of many other clades. We also show a marked lineage-wide slowdown in diversification rates in the Miocene. Contrasting this overall pattern, a species group associated with the young and seasonally harsh Chacoan biogeographic region underwent a recent burst of diversification. Subsequent analyses also indicated that there is significant phylogenetic clustering within the Chacoan region and that speciation rates are highest there. Together, these findings suggest that recent ecological opportunity, from successful colonization of novel habitat, may have facilitated renewed turtle ant diversification. Our findings highlight a central role of ecological opportunity within a successful continental radiation.

%B Journal of Evolutionary Biology %V 27 %P 242-258 %8 Feb %@ 1010-061X %G English %M WOS:000330016400002 %! J Evolution BiolJ Evolution Biol %0 Journal Article %J Zoologica Scripta %D 2014 %T Revised systematics and higher classification of pierid butterflies (Lepidoptera: Pieridae) based on molecular data %A Wahlberg, N. %A Rota, J. %A Braby, M. F. %A Pierce, N. E. %A Wheat, C. W. %K biogeography %K delias hubner lepidoptera %K diversification %K Evolution %K genera lepidoptera %K immature stages %K larval food plants %K neotropical mistletoe butterflies %K origin %K phylogenetic inference %X

The butterfly family Pieridae comprises approximately 1000 described species placed in 85 genera, but the higher classification has not yet been settled. We used molecular data from eight gene regions (one mitochondrial and seven nuclear protein-coding genes) comprising a total of similar to 6700bp from 96 taxa to infer a well-supported phylogenetic hypothesis for the family. Based on this hypothesis, we revise the higher classification for all pierid genera. We resurrect the tribe Teracolini stat. rev. in the subfamily Pierinae to include the genera Teracolus, Pinacopteryx, Gideona, Ixias, Eronia, Colotis and most likely Calopieris. We transfer Hebomoia to the tribe Anthocharidini and assign the previously unplaced genera Belenois and Dixeia to the subtribe Aporiina. Three lineages near the base of Pierinae (Leptosia, Elodina and Nepheronia + Pareronia) remain unplaced. For each of these, we describe and delineate new tribes: Elodinini Braby tribus nova, Leptosiaini Braby tribus nova and Nepheroniini Braby tribus nova. The proposed higher classification is based on well-supported monophyletic groups and is likely to remain stable even with the addition of more data.

%B Zoologica Scripta %V 43 %P 641-650 %8 Nov %@ 0300-3256 %G English %M WOS:000343866100007 %! Zool ScrZool Scr %0 Journal Article %J Current Biology %D 2014 %T A Social Parasite Evolved Reproductive Isolation from Its Fungus-Growing Ant Host in Sympatry %A Rabeling, C. %A Schultz, T. R. %A Pierce, N. E. %A Bacci, M. %K alternative adaptations %K Evolution %K hymenoptera-formicidae %K inquiline %K mycocepurus-smithii %K phylogenetic analysis %K sequences %K Speciation %K species trees %K x-linked genes %X

Inquiline social parasitic ant species exploit colonies of other ant species mainly by producing sexual offspring that are raised by the host. Ant social parasites and their hosts are often close relatives (Emery's rule), and two main hypotheses compete to explain the parasites' evolutionary origins: (1) the interspecific hypothesis proposes an allopatric speciation scenario for the parasite, whereas (2) the intraspecific hypothesis postulates that the parasite evolves directly from its host in sympatry [1-10]. Evidence in support of the intraspecific hypothesis has been accumulating for ants [3, 5, 7, 9-12], but sympatric speciation remains controversial as a general speciation mechanism for inquiline parasites. Here we use molecular phylogenetics to assess whether the socially parasitic fungus-growing ant Mycocepurus castrator speciated from its host Mycocepurus goeldii in sympatry. Based on differing patterns of relationship in mitochondrial and individual nuclear genes, we conclude that host and parasite occupy a temporal window in which lineage sorting has taken place in the mitochondrial genes but not yet in the nuclear alleles. We infer that the host originated first and that the parasite originated subsequently from a subset of the host species' populations, providing empirical support for the hypothesis that inquiline parasites can evolve reproductive isolation while living sympatrically with their hosts.

%B Current Biology %V 24 %P 2047-2052 %8 Sep 8 %@ 0960-9822 %G English %M WOS:000341541300031 %! Curr BiolCurr Biol %0 Journal Article %J Molecular Ecology %D 2014 %T Stability and phylogenetic correlation in gut microbiota: lessons from ants and apes %A Sanders, J. G. %A Powell, S. %A Kronauer, D. J. %A Vasconcelos, H. L. %A Frederickson, M. E. %A Pierce, N. E. %K *Microbiota %K *Phylogeny %K Animals %K Ants/genetics/*microbiology %K Biodiversity %K Biological Evolution %K Cluster Analysis %K Digestive System/*microbiology %K DNA, Bacterial/genetics %K Hominidae/microbiology %K RNA, Ribosomal, 16S/genetics %K Sequence Analysis, DNA %X

Correlation between gut microbiota and host phylogeny could reflect codiversification over shared evolutionary history or a selective environment that is more similar in related hosts. These alternatives imply substantial differences in the relationship between host and symbiont, but can they be distinguished based on patterns in the community data themselves? We explored patterns of phylogenetic correlation in the distribution of gut bacteria among species of turtle ants (genus Cephalotes), which host a dense gut microbial community. We used 16S rRNA pyrosequencing from 25 Cephalotes species to show that their gut community is remarkably stable, from the colony to the genus level. Despite this overall similarity, the existing differences among species' microbiota significantly correlated with host phylogeny. We introduced a novel analytical technique to test whether these phylogenetic correlations are derived from recent bacterial evolution, as would be expected in the case of codiversification, or from broader shifts more likely to reflect environmental filters imposed by factors such as diet or habitat. We also tested this technique on a published data set of ape microbiota, confirming earlier results while revealing previously undescribed patterns of phylogenetic correlation. Our results indicated a high degree of partner fidelity in the Cephalotes microbiota, suggesting that vertical transmission of the entire community could play an important role in the evolution and maintenance of the association. As additional comparative microbiota data become available, the techniques presented here can be used to explore trends in the evolution of host-associated microbial communities.

%B Molecular Ecology %V 23 %P 1268-83 %8 Mar %@ 1365-294X (Electronic)0962-1083 (Linking) %G eng %M 24304129 %! Mol EcolMol Ecol %0 Journal Article %J Proceedings of the Royal Society B-Biological Sciences %D 2014 %T Transitions in social complexity along elevational gradients reveal a combined impact of season length and development time on social evolution %A Kocher, S. D. %A Pellissier, L. %A Veller, C. %A Purcell, J. %A Nowak, M. A. %A Chapuisat, M. %A Pierce, N. E. %K Altitude %K bumble bees %K development time %K eusociality %K halictidae %K hymenoptera %K hypothesis %K Insects %K mountain passes %K polistes %K Population %K social behaviour %K solitary %K species distributions %X

Eusociality is taxonomically rare, yet associated with great ecological success. Surprisingly, studies of environmental conditions favouring eusociality are often contradictory. Harsh conditions associated with increasing altitude and latitude seem to favour increased sociality in bumblebees and ants, but the reverse pattern is found in halictid bees and polistine wasps. Here, we compare the life histories and distributions of populations of 176 species of Hymenoptera from the Swiss Alps. We show that differences in altitudinal distributions and development times among social forms can explain these contrasting patterns: highly social taxa develop more quickly than intermediate social taxa, and are thus able to complete the reproductive cycle in shorter seasons at higher elevations. This dual impact of altitude and development time on sociality illustrates that ecological constraints can elicit dynamic shifts in behaviour, and helps explain the complex distribution of sociality across ecological gradients.

%B Proceedings of the Royal Society B-Biological Sciences %V 281 %8 Jul 22 %@ 0962-8452 %G English %M WOS:000338335800019 %! P Roy Soc B-Biol SciP Roy Soc B-Biol Sci %0 Journal Article %J Proceedings of the National Academy of Sciences of the United States of America %D 2013 %T Animals in a bacterial world, a new imperative for the life sciences %A McFall-Ngai, M. %A Hadfield, M. G. %A Bosch, T. C. G. %A Carey, H. V. %A Domazet-Loso, T. %A Douglas, A. E. %A Dubilier, N. %A Eberl, G. %A Fukami, T. %A Gilbert, S. F. %A Hentschel, U. %A King, N. %A Kjelleberg, S. %A Knoll, A. H. %A Kremer, N. %A Mazmanian, S. K. %A Metcalf, J. L. %A Nealson, K. %A Pierce, N. E. %A Rawls, J. F. %A Reid, A. %A Ruby, E. G. %A Rumpho, M. %A Sanders, J. G. %A Tautz, D. %A Wernegreen, J. J. %K bacteria-driven development %K bacterial roles animal origins %K colonization %K euprymna-scolopes %K gut microbiota %K host %K innate immune homeostasis %K intestinal microbiota %K larval settlement %K marine-bacteria %K microbiome and host physiology %K nested ecosystems %K ocean acidification %K reciprocal effects animal-bacterial genomics %K vibrio-fischeri %X

In the last two decades, the widespread application of genetic and genomic approaches has revealed a bacterial world astonishing in its ubiquity and diversity. This review examines how a growing knowledge of the vast range of animal-bacterial interactions, whether in shared ecosystems or intimate symbioses, is fundamentally altering our understanding of animal biology. Specifically, we highlight recent technological and intellectual advances that have changed our thinking about five questions: how have bacteria facilitated the origin and evolution of animals; how do animals and bacteria affect each other's genomes; how does normal animal development depend on bacterial partners; how is homeostasis maintained between animals and their symbionts; and how can ecological approaches deepen our understanding of the multiple levels of animal-bacterial interaction. As answers to these fundamental questions emerge, all biologists will be challenged to broaden their appreciation of these interactions and to include investigations of the relationships between and among bacteria and their animal partners as we seek a better understanding of the natural world.

%B Proceedings of the National Academy of Sciences of the United States of America %V 110 %P 3229-3236 %8 Feb 26 %@ 0027-8424 %G English %M WOS:000315841900016 %! P Natl Acad Sci USAP Natl Acad Sci USA %0 Journal Article %J Biological Journal of the Linnean Society %D 2013 %T Association between the African lycaenid, Anthene usamba, and an obligate acacia ant, Crematogaster mimosae %A Martins, D. J. %A Collins, S. C. %A Congdon, C. %A Pierce, N. E. %K acacia drepanolobium %K attendant ants %K Ecology %K jalmenus-evagoras %K kenya %K larvae %K lepidoptera %K mutualism %K myrmicinae %K oecophylla-smaragdina %K oviposition %K parasitism %K plant-selection %K polyommatinae %K swollen thorn acacias %X

The African lycaenid butterfly, Anthene usamba, is an obligate myrmecophile of the acacia ant, Crematogaster mimosae. Female butterflies use the presence of C.mimosae as an oviposition cue. The eggs are laid on the foliage and young branches of the host plant, Acacia drepanolobium. Larvae shelter in the swollen thorns (domatia) of the host tree, where they live in close association with the acacia ants, and each larva occupies a domatium singly. Anthene usamba are tended by ants that feed from the dorsal nectary organ at regular intervals. Larvae also possess tentacle organs flanking the dorsal nectary organ and appear to signal to ants by everting these structures. Larvae were observed to spend most of their time within the domatia. Stable isotope analysis of matched host plantantbutterfly samples revealed that Anthene usamba are 15N enriched relative to the ants with which they associate. These data, based on the increase in 15N through trophic levels, indicate that the caterpillars of these butterflies are aphytophagous and either exploit the ant brood of C.mimosae within the domatia, or are fed mouth to mouth by adult workers via trophallaxis. This is the first documented case of aphytophagy in African Anthene. Pupation occurs inside the domatium and the imago emerges and departs via the hole chewed by the larva. The adult females remain closely associated with their natal patch of trees, whereas males disperse more widely across the acacia savannah. Females prefer to oviposit on trees with the specific host ant, C.mimosae, an aggressive obligate mutualist, and avoid neighbouring trees with other ant species. Adult butterflies are active during most months of the year, and there are at least two to three generations each year. Observations made over a 5-year period indicate that a number of different lycaenid species utilize ant-acacias in East Africa, and these observations are summarized, together with comparisons from the literature.(c) 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109, 302312.

%B Biological Journal of the Linnean Society %V 109 %P 302-312 %8 Jun %@ 0024-4066 %G English %M WOS:000318809500006 %! Biol J Linn SocBiol J Linn Soc %0 Journal Article %J Insect Conservation and Diversity %D 2013 %T Cross-continental comparisons of butterfly assemblages in tropical rainforests: implications for biological monitoring %A Basset, Y. %A Eastwood, R. %A Sam, L. %A Lohman, D. J. %A Novotny, V. %A Treuer, T. %A Miller, S. E. %A Weiblen, G. D. %A Pierce, N. E. %A Bunyavejchewin, S. %A Sakchoowong, W. %A Kongnoo, P. %A Osorio-Arenas, M. A. %K abundance %K barro colorado island %K biological monitoring %K center for tropical forest science %K Climate %K community %K Conservation %K disturbance %K dry forest %K Ecology %K insect diversity %K lepidoptera %K Panama %K papua new guinea %K pollard walks %K species-diversity %K Temperature %K thailand %K tropical rainforest %X

. 1.Standardised transect counts of butterflies in old-growth rainforests in different biogeographical regions are lacking. Such data are needed to mitigate the influence of methodological and environmental factors within and between sites and, ultimately, to discriminate between long-term trends and short-term stochastic changes in abundance and community composition. 2.We compared butterfly assemblages using standardised Pollard Walks in the understory of closed-canopy lowland tropical rainforests across three biogeographical regions: Barro Colorado Island (BCI), Panama; Khao Chong (KHC), Thailand; and Wanang (WAN), Papua New Guinea. 3.The length and duration of transects, their spatial autocorrelation, and number of surveys per year represented important methodological factors that strongly influenced estimates of butterfly abundance. Of these, the effect of spatial autocorrelation was most difficult to mitigate across study sites. 4.Butterfly abundance and faunal composition were best explained by air temperature, elevation, rainfall, wind velocity, and human disturbance at BCI and KHC. In the absence of weather data at WAN, duration of transects and number of forest gaps accounted for most of the explained variance, which was rather low in all cases (<33%). 5.Adequate monitoring of the abundance of common butterflies was achieved at the 50ha BCI plot, with three observers walking each of 10 transects of 500m for 30min each, during each of four surveys per year. These data may be standardised further after removing outliers of temperature and rainfall. Practical procedures are suggested to implement global monitoring of rainforest butterflies with Pollard Walks.

%B Insect Conservation and Diversity %V 6 %P 223-233 %8 May %@ 1752-458X %G English %M WOS:000319406400003 %! Insect Conserv DiverInsect Conserv Diver %0 Journal Article %J Genome Biology %D 2013 %T The draft genome of a socially polymorphic halictid bee, Lasioglossum albipes %A Kocher, S. D. %A Li, C. %A Yang, W. %A Tan, H. %A Yi, S. V. %A Yang, X. Y. %A Hoekstra, H. E. %A Zhang, G. J. %A Pierce, N. E. %A Yu, D. W. %K adaptive evolution %K ants camponotus-floridanus %K DNA Methylation %K eusocial nests %K family %K gene %K harpegnathos-saltator %K honey-bees %K maximum-likelihood %K phylogenetic inference %X

Background: Taxa that harbor natural phenotypic variation are ideal for ecological genomic approaches aimed at understanding how the interplay between genetic and environmental factors can lead to the evolution of complex traits. Lasioglossum albipes is a polymorphic halictid bee that expresses variation in social behavior among populations, and common-garden experiments have suggested that this variation is likely to have a genetic component.Results: We present the L. albipes genome assembly to characterize the genetic and ecological factors associated with the evolution of social behavior. The de novo assembly is comparable to other published social insect genomes, with an N50 scaffold length of 602 kb. Gene families unique to L. albipes are associated with integrin-mediated signaling and DNA-binding domains, and several appear to be expanded in this species, including the glutathione-s-transferases and the inositol monophosphatases. L. albipes has an intact DNA methylation system, and in silico analyses suggest that methylation occurs primarily in exons. Comparisons to other insect genomes indicate that genes associated with metabolism and nucleotide binding undergo accelerated evolution in the halictid lineage. Whole-genome resequencing data from one solitary and one social L. albipes female identify six genes that appear to be rapidly diverging between social forms, including a putative odorant receptor and a cuticular protein.Conclusions: L. albipes represents a novel genetic model system for understanding the evolution of social behavior. It represents the first published genome sequence of a primitively social insect, thereby facilitating comparative genomic studies across the Hymenoptera as a whole.

%B Genome Biology %V 14 %@ 1465-6906 %G English %M WOS:000331376200010 %! Genome BiolGenome Biol %0 Journal Article %J Molecular Phylogenetics and Evolution %D 2013 %T In the shadow of phylogenetic uncertainty: The recent diversification of Lysandra butterflies through chromosomal change %A Talavera, G. %A Lukhtanov, V. A. %A Rieppel, L. %A Pierce, N. E. %A Vila, R. %K agrodiaetus butterflies %K chromosomal evolution %K gene trees %K hybridization %K incomplete lineage sorting %K karyotype evolution %K lepidoptera %K lepidoptera lycaenidae %K maximum-likelihood %K mitochondrial-DNA introgression %K molecular-data %K Speciation %K species trees %X

The phylogeny of the butterfly genus Lysandra (Lycaenidae, Polyommatinae) has been intractable using both molecular and morphological characters, which could be a result of speciation due to karyotype instability. Here we reconstruct the phylogeny of the group using multi-locus coalescent-based methods on seven independent genetic markers. While the genus is ca. 4.9 Mya old, the diversification of the extant lineages was extremely recent (ca. 1.5 Mya) and involved multiple chromosomal rearrangements. We find that relationships are uncertain due to both incomplete lineage sorting and hybridization. Minimizing the impact of reticulation in inferring the species tree by testing for mitochondria] introgression events yields a partially resolved tree with three main supported clades: L. punctifera + L. bellargus, the corydonius taxa, and L coridon + the Iberian taxa, plus three independent lineages without apparently close relatives (L. ossmar, L syriaca and L. dezina). Based on these results and new karyotypic data, we propose a rearrangement recognizing ten species within the genus. Finally, we hypothesize that chromosomal instability may have played a crucial role in the Lysandra recent diversification. New chromosome rearrangements might be fixed in populations after severe bottlenecks, which at the same time might promote rapid sorting of neutral molecular markers. We argue that population bottlenecks might be a prerequisite for chromosomal speciation in this group. (C) 2013 Elsevier Inc. All rights reserved.

%B Molecular Phylogenetics and Evolution %V 69 %P 469-478 %8 Dec %@ 1055-7903 %G English %M WOS:000326417600006 %! Mol Phylogenet EvolMol Phylogenet Evol %0 Journal Article %J Behavioral Ecology %D 2013 %T Non-nest mate discrimination and clonal colony structure in the parthenogenetic ant Cerapachys biroi %A Kronauer, D. J. C. %A Tsuji, K. %A Pierce, N. E. %A Keller, L. %K aggression %K asexuality %K chimera %K cooperation %K fire ant %K forel formicidae %K formicidae %K genetic conflicts %K hymenoptera-formicidae %K kin recognition %K platythyrea-punctata %K pristomyrmex-punctatus %K sexual reproduction %K social insects %K thelytocous-parthenogenesis %K thelytoky %X

Understanding the interplay between cooperation and conflict in social groups is a major goal of biology. One important factor is genetic relatedness, and animal societies are usually composed of related but genetically different individuals, setting the stage for conflicts over reproductive allocation. Recently, however, it has been found that several ant species reproduce predominantly asexually. Although this can potentially give rise to clonal societies, in the few well-studied cases, colonies are often chimeric assemblies of different genotypes, due to worker drifting or colony fusion. In the ant Cerapachys biroi, queens are absent and all individuals reproduce via thelytokous parthenogenesis, making this species an ideal study system of asexual reproduction and its consequences for social dynamics. Here, we show that colonies in our study population on Okinawa, Japan, recognize and effectively discriminate against foreign workers, especially those from unrelated asexual lineages. In accord with this finding, colonies never contained more than a single asexual lineage and average pairwise genetic relatedness within colonies was extremely high (r = 0.99). This implies that the scope for social conflict in C. biroi is limited, with unusually high potential for cooperation and altruism.

%B Behavioral Ecology %V 24 %P 617-622 %8 May-Jun %@ 1045-2249 %G English %M WOS:000317320100010 %! Behav EcolBehav Ecol %0 Journal Article %J Plant Cell %D 2013 %T Pathogen-Triggered Ethylene Signaling Mediates Systemic-Induced Susceptibility to Herbivory in Arabidopsis %A Groen, S. C. %A Whiteman, N. K. %A Bahrami, A. K. %A Wilczek, A. M. %A Cui, J. P. %A Russell, J. A. %A Cibrian-Jaramillo, A. %A Butler, I. A. %A Rana, J. D. %A Huang, G. H. %A Bush, J. %A Ausubel, F. M. %A Pierce, N. E. %K acquired-resistance %K defense responses %K dependent cell-death %K disease resistance %K jasmonic acid %K nicotiana-attenuata %K plant defenses %K pseudomonas-syringae %K salicylic-acid %K syringae pv. tomato %X

Multicellular eukaryotic organisms are attacked by numerous parasites from diverse phyla, often simultaneously or sequentially. An outstanding question in these interactions is how hosts integrate signals induced by the attack of different parasites. We used a model system comprised of the plant host Arabidopsis thaliana, the hemibiotrophic bacterial phytopathogen Pseudomonas syringae, and herbivorous larvae of the moth Trichoplusia ni (cabbage looper) to characterize mechanisms involved in systemic-induced susceptibility (SIS) to T. ni herbivory caused by prior infection by virulent P. syringae. We uncovered a complex multilayered induction mechanism for SIS to herbivory. In this mechanism, antiherbivore defenses that depend on signaling via (1) the jasmonic acid-isoleucine conjugate (JA-Ile) and (2) other octadecanoids are suppressed by microbe-associated molecular pattern-triggered salicylic acid (SA) signaling and infection-triggered ethylene signaling, respectively. SIS to herbivory is, in turn, counteracted by a combination of the bacterial JA-Ile mimic coronatine and type III virulence-associated effectors. Our results show that SIS to herbivory involves more than antagonistic signaling between SA and JA-Ile and provide insight into the unexpectedly complex mechanisms behind a seemingly simple trade-off in plant defense against multiple enemies.

%B Plant Cell %V 25 %P 4755-4766 %8 Nov %@ 1040-4651 %G English %M WOS:000329174400034 %! Plant CellPlant Cell %0 Journal Article %J Proceedings of the National Academy of Sciences of the United States of America %D 2013 %T Animals in a bacterial world, a new imperative for the life sciences %A McFall-Ngai, M. %A Hadfield, M. G. %A Bosch, T. C. G. %A Carey, H. V. %A Domazet-Loso, T. %A Douglas, A. E. %A Dubilier, N. %A Eberl, G. %A Fukami, T. %A Gilbert, S. F. %A Hentschel, U. %A King, N. %A Kjelleberg, S. %A Knoll, A. H. %A Kremer, N. %A Mazmanian, S. K. %A Metcalf, J. L. %A Nealson, K. %A Pierce, N. E. %A Rawls, J. F. %A Reid, A. %A Ruby, E. G. %A Rumpho, M. %A Sanders, J. G. %A Tautz, D. %A Wernegreen, J. J. %K bacteria-driven development %K bacterial roles animal origins %K colonization %K euprymna-scolopes %K gut microbiota %K host %K innate immune homeostasis %K intestinal microbiota %K larval settlement %K marine-bacteria %K microbiome and host physiology %K nested ecosystems %K ocean acidification %K reciprocal effects animal-bacterial genomics %K vibrio-fischeri %X

In the last two decades, the widespread application of genetic and genomic approaches has revealed a bacterial world astonishing in its ubiquity and diversity. This review examines how a growing knowledge of the vast range of animal-bacterial interactions, whether in shared ecosystems or intimate symbioses, is fundamentally altering our understanding of animal biology. Specifically, we highlight recent technological and intellectual advances that have changed our thinking about five questions: how have bacteria facilitated the origin and evolution of animals; how do animals and bacteria affect each other's genomes; how does normal animal development depend on bacterial partners; how is homeostasis maintained between animals and their symbionts; and how can ecological approaches deepen our understanding of the multiple levels of animal-bacterial interaction. As answers to these fundamental questions emerge, all biologists will be challenged to broaden their appreciation of these interactions and to include investigations of the relationships between and among bacteria and their animal partners as we seek a better understanding of the natural world.

%B Proceedings of the National Academy of Sciences of the United States of America %V 110 %P 3229-3236 %8 Feb 26 %@ 0027-8424 %G English %M WOS:000315841900016 %! P Natl Acad Sci USAP Natl Acad Sci USA %0 Journal Article %J Biological Journal of the Linnean Society %D 2013 %T Association between the African lycaenid, Anthene usamba, and an obligate acacia ant, Crematogaster mimosae %A Martins, D. J. %A Collins, S. C. %A Congdon, C. %A Pierce, N. E. %K acacia drepanolobium %K attendant ants %K Ecology %K jalmenus-evagoras %K kenya %K larvae %K lepidoptera %K mutualism %K myrmicinae %K oecophylla-smaragdina %K oviposition %K parasitism %K plant-selection %K polyommatinae %K swollen thorn acacias %X

The African lycaenid butterfly, Anthene usamba, is an obligate myrmecophile of the acacia ant, Crematogaster mimosae. Female butterflies use the presence of C.mimosae as an oviposition cue. The eggs are laid on the foliage and young branches of the host plant, Acacia drepanolobium. Larvae shelter in the swollen thorns (domatia) of the host tree, where they live in close association with the acacia ants, and each larva occupies a domatium singly. Anthene usamba are tended by ants that feed from the dorsal nectary organ at regular intervals. Larvae also possess tentacle organs flanking the dorsal nectary organ and appear to signal to ants by everting these structures. Larvae were observed to spend most of their time within the domatia. Stable isotope analysis of matched host plantantbutterfly samples revealed that Anthene usamba are 15N enriched relative to the ants with which they associate. These data, based on the increase in 15N through trophic levels, indicate that the caterpillars of these butterflies are aphytophagous and either exploit the ant brood of C.mimosae within the domatia, or are fed mouth to mouth by adult workers via trophallaxis. This is the first documented case of aphytophagy in African Anthene. Pupation occurs inside the domatium and the imago emerges and departs via the hole chewed by the larva. The adult females remain closely associated with their natal patch of trees, whereas males disperse more widely across the acacia savannah. Females prefer to oviposit on trees with the specific host ant, C.mimosae, an aggressive obligate mutualist, and avoid neighbouring trees with other ant species. Adult butterflies are active during most months of the year, and there are at least two to three generations each year. Observations made over a 5-year period indicate that a number of different lycaenid species utilize ant-acacias in East Africa, and these observations are summarized, together with comparisons from the literature.(c) 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109, 302312.

%B Biological Journal of the Linnean Society %V 109 %P 302-312 %8 Jun %@ 0024-4066 %G English %M WOS:000318809500006 %! Biol J Linn SocBiol J Linn Soc %0 Journal Article %J Insect Conservation and Diversity %D 2013 %T Cross-continental comparisons of butterfly assemblages in tropical rainforests: implications for biological monitoring %A Basset, Y. %A Eastwood, R. %A Sam, L. %A Lohman, D. J. %A Novotny, V. %A Treuer, T. %A Miller, S. E. %A Weiblen, G. D. %A Pierce, N. E. %A Bunyavejchewin, S. %A Sakchoowong, W. %A Kongnoo, P. %A Osorio-Arenas, M. A. %K abundance %K barro colorado island %K biological monitoring %K center for tropical forest science %K Climate %K community %K Conservation %K disturbance %K dry forest %K Ecology %K insect diversity %K lepidoptera %K Panama %K papua new guinea %K pollard walks %K species-diversity %K Temperature %K thailand %K tropical rainforest %X

. 1.Standardised transect counts of butterflies in old-growth rainforests in different biogeographical regions are lacking. Such data are needed to mitigate the influence of methodological and environmental factors within and between sites and, ultimately, to discriminate between long-term trends and short-term stochastic changes in abundance and community composition. 2.We compared butterfly assemblages using standardised Pollard Walks in the understory of closed-canopy lowland tropical rainforests across three biogeographical regions: Barro Colorado Island (BCI), Panama; Khao Chong (KHC), Thailand; and Wanang (WAN), Papua New Guinea. 3.The length and duration of transects, their spatial autocorrelation, and number of surveys per year represented important methodological factors that strongly influenced estimates of butterfly abundance. Of these, the effect of spatial autocorrelation was most difficult to mitigate across study sites. 4.Butterfly abundance and faunal composition were best explained by air temperature, elevation, rainfall, wind velocity, and human disturbance at BCI and KHC. In the absence of weather data at WAN, duration of transects and number of forest gaps accounted for most of the explained variance, which was rather low in all cases (<33%). 5.Adequate monitoring of the abundance of common butterflies was achieved at the 50ha BCI plot, with three observers walking each of 10 transects of 500m for 30min each, during each of four surveys per year. These data may be standardised further after removing outliers of temperature and rainfall. Practical procedures are suggested to implement global monitoring of rainforest butterflies with Pollard Walks.

%B Insect Conservation and Diversity %V 6 %P 223-233 %8 May %@ 1752-458X %G English %M WOS:000319406400003 %! Insect Conserv DiverInsect Conserv Diver %0 Journal Article %J Genome Biology %D 2013 %T The draft genome of a socially polymorphic halictid bee, Lasioglossum albipes %A Kocher, S. D. %A Li, C. %A Yang, W. %A Tan, H. %A Yi, S. V. %A Yang, X. Y. %A Hoekstra, H. E. %A Zhang, G. J. %A Pierce, N. E. %A Yu, D. W. %K adaptive evolution %K ants camponotus-floridanus %K DNA Methylation %K eusocial nests %K family %K gene %K harpegnathos-saltator %K honey-bees %K maximum-likelihood %K phylogenetic inference %X

Background: Taxa that harbor natural phenotypic variation are ideal for ecological genomic approaches aimed at understanding how the interplay between genetic and environmental factors can lead to the evolution of complex traits. Lasioglossum albipes is a polymorphic halictid bee that expresses variation in social behavior among populations, and common-garden experiments have suggested that this variation is likely to have a genetic component.Results: We present the L. albipes genome assembly to characterize the genetic and ecological factors associated with the evolution of social behavior. The de novo assembly is comparable to other published social insect genomes, with an N50 scaffold length of 602 kb. Gene families unique to L. albipes are associated with integrin-mediated signaling and DNA-binding domains, and several appear to be expanded in this species, including the glutathione-s-transferases and the inositol monophosphatases. L. albipes has an intact DNA methylation system, and in silico analyses suggest that methylation occurs primarily in exons. Comparisons to other insect genomes indicate that genes associated with metabolism and nucleotide binding undergo accelerated evolution in the halictid lineage. Whole-genome resequencing data from one solitary and one social L. albipes female identify six genes that appear to be rapidly diverging between social forms, including a putative odorant receptor and a cuticular protein.Conclusions: L. albipes represents a novel genetic model system for understanding the evolution of social behavior. It represents the first published genome sequence of a primitively social insect, thereby facilitating comparative genomic studies across the Hymenoptera as a whole.

%B Genome Biology %V 14 %@ 1465-6906 %G English %M WOS:000331376200010 %! Genome BiolGenome Biol %0 Journal Article %J Molecular Phylogenetics and Evolution %D 2013 %T In the shadow of phylogenetic uncertainty: The recent diversification of Lysandra butterflies through chromosomal change %A Talavera, G. %A Lukhtanov, V. A. %A Rieppel, L. %A Pierce, N. E. %A Vila, R. %K agrodiaetus butterflies %K chromosomal evolution %K gene trees %K hybridization %K incomplete lineage sorting %K karyotype evolution %K lepidoptera %K lepidoptera lycaenidae %K maximum-likelihood %K mitochondrial-DNA introgression %K molecular-data %K Speciation %K species trees %X

The phylogeny of the butterfly genus Lysandra (Lycaenidae, Polyommatinae) has been intractable using both molecular and morphological characters, which could be a result of speciation due to karyotype instability. Here we reconstruct the phylogeny of the group using multi-locus coalescent-based methods on seven independent genetic markers. While the genus is ca. 4.9 Mya old, the diversification of the extant lineages was extremely recent (ca. 1.5 Mya) and involved multiple chromosomal rearrangements. We find that relationships are uncertain due to both incomplete lineage sorting and hybridization. Minimizing the impact of reticulation in inferring the species tree by testing for mitochondria] introgression events yields a partially resolved tree with three main supported clades: L. punctifera + L. bellargus, the corydonius taxa, and L coridon + the Iberian taxa, plus three independent lineages without apparently close relatives (L. ossmar, L syriaca and L. dezina). Based on these results and new karyotypic data, we propose a rearrangement recognizing ten species within the genus. Finally, we hypothesize that chromosomal instability may have played a crucial role in the Lysandra recent diversification. New chromosome rearrangements might be fixed in populations after severe bottlenecks, which at the same time might promote rapid sorting of neutral molecular markers. We argue that population bottlenecks might be a prerequisite for chromosomal speciation in this group. (C) 2013 Elsevier Inc. All rights reserved.

%B Molecular Phylogenetics and Evolution %V 69 %P 469-478 %8 Dec %@ 1055-7903 %G English %M WOS:000326417600006 %! Mol Phylogenet EvolMol Phylogenet Evol %0 Journal Article %J Behavioral Ecology %D 2013 %T Non-nest mate discrimination and clonal colony structure in the parthenogenetic ant Cerapachys biroi %A Kronauer, D. J. C. %A Tsuji, K. %A Pierce, N. E. %A Keller, L. %K aggression %K asexuality %K chimera %K cooperation %K fire ant %K forel formicidae %K formicidae %K genetic conflicts %K hymenoptera-formicidae %K kin recognition %K platythyrea-punctata %K pristomyrmex-punctatus %K sexual reproduction %K social insects %K thelytocous-parthenogenesis %K thelytoky %X

Understanding the interplay between cooperation and conflict in social groups is a major goal of biology. One important factor is genetic relatedness, and animal societies are usually composed of related but genetically different individuals, setting the stage for conflicts over reproductive allocation. Recently, however, it has been found that several ant species reproduce predominantly asexually. Although this can potentially give rise to clonal societies, in the few well-studied cases, colonies are often chimeric assemblies of different genotypes, due to worker drifting or colony fusion. In the ant Cerapachys biroi, queens are absent and all individuals reproduce via thelytokous parthenogenesis, making this species an ideal study system of asexual reproduction and its consequences for social dynamics. Here, we show that colonies in our study population on Okinawa, Japan, recognize and effectively discriminate against foreign workers, especially those from unrelated asexual lineages. In accord with this finding, colonies never contained more than a single asexual lineage and average pairwise genetic relatedness within colonies was extremely high (r = 0.99). This implies that the scope for social conflict in C. biroi is limited, with unusually high potential for cooperation and altruism.

%B Behavioral Ecology %V 24 %P 617-622 %8 May-Jun %@ 1045-2249 %G English %M WOS:000317320100010 %! Behav EcolBehav Ecol %0 Journal Article %J Plant Cell %D 2013 %T Pathogen-Triggered Ethylene Signaling Mediates Systemic-Induced Susceptibility to Herbivory in Arabidopsis %A Groen, S. C. %A Whiteman, N. K. %A Bahrami, A. K. %A Wilczek, A. M. %A Cui, J. P. %A Russell, J. A. %A Cibrian-Jaramillo, A. %A Butler, I. A. %A Rana, J. D. %A Huang, G. H. %A Bush, J. %A Ausubel, F. M. %A Pierce, N. E. %K acquired-resistance %K defense responses %K dependent cell-death %K disease resistance %K jasmonic acid %K nicotiana-attenuata %K plant defenses %K pseudomonas-syringae %K salicylic-acid %K syringae pv. tomato %X

Multicellular eukaryotic organisms are attacked by numerous parasites from diverse phyla, often simultaneously or sequentially. An outstanding question in these interactions is how hosts integrate signals induced by the attack of different parasites. We used a model system comprised of the plant host Arabidopsis thaliana, the hemibiotrophic bacterial phytopathogen Pseudomonas syringae, and herbivorous larvae of the moth Trichoplusia ni (cabbage looper) to characterize mechanisms involved in systemic-induced susceptibility (SIS) to T. ni herbivory caused by prior infection by virulent P. syringae. We uncovered a complex multilayered induction mechanism for SIS to herbivory. In this mechanism, antiherbivore defenses that depend on signaling via (1) the jasmonic acid-isoleucine conjugate (JA-Ile) and (2) other octadecanoids are suppressed by microbe-associated molecular pattern-triggered salicylic acid (SA) signaling and infection-triggered ethylene signaling, respectively. SIS to herbivory is, in turn, counteracted by a combination of the bacterial JA-Ile mimic coronatine and type III virulence-associated effectors. Our results show that SIS to herbivory involves more than antagonistic signaling between SA and JA-Ile and provide insight into the unexpectedly complex mechanisms behind a seemingly simple trade-off in plant defense against multiple enemies.

%B Plant Cell %V 25 %P 4755-4766 %8 Nov %@ 1040-4651 %G English %M WOS:000329174400034 %! Plant CellPlant Cell %0 Journal Article %J Molecular Ecology %D 2012 %T Asexual reproduction in introduced and native populations of the ant Cerapachys biroi %A Kronauer, D. J. C. %A Pierce, N. E. %A Keller, L. %K anthropogenic introduction %K argentine ant %K asexuality %K automixis %K clonal reproduction %K Evolution %K fire ant %K formicidae %K fungus-growing ant %K hymenoptera-formicidae %K invasion %K maximum-likelihood %K parthenogenesis %K platythyrea-punctata %K thelytocous-parthenogenesis %K thelytoky %X

Asexual reproduction is particularly common among introduced species, probably because it helps to overcome the negative effects associated with low population densities during colonization. The ant Cerapachys biroi has been introduced to tropical and subtropical islands around the world since the beginning of the last century. In this species, workers can reproduce via thelytokous parthenogenesis. Here, we use genetic markers to reconstruct the history of anthropogenic introductions of C.biroi, and to address the prevalence of female parthenogenesis in introduced and native populations. We show that at least four genetically distinct lineages have been introduced from continental Asia and have led to the species' circumtropical establishment. Our analyses demonstrate that asexual reproduction dominates in the introduced range and is also common in the native range. Given that C.biroi is the only dorylomorph ant that has successfully become established outside of its native range, this unusual mode of reproduction probably facilitated the species' worldwide spread. On the other hand, the rare occurrence of haploid males and at least one clear case of sexual recombination in the introduced range show that C.biroi has not lost the potential for sex. Finally, we show that thelytoky in C.biroi probably has a genetic rather than an infectious origin, and that automixis with central fusion is the most likely underlying cytological mechanism. This is in accordance with what is known for other thelytokous eusocial Hymenoptera.

%B Molecular Ecology %V 21 %P 5221-5235 %8 Nov %@ 0962-1083 %G English %M WOS:000310065600008 %! Mol EcolMol Ecol %0 Journal Article %J Insect Conservation and Diversity %D 2012 %T Cross-continental comparisons of butterfly assemblages in tropical rainforests: implications for biological monitoring %A Basset, Y %A Eastwood, R. %A L Sam %A DJ Lohman %A V Novotny %A T Treuer %A SE Miller %A GD Weiblen %A Pierce, N. E. %A Bunyavejchewin, S. %A W Sakchoowong %A P Kongnoo %A MA Osorio-Arenas %X

. 1. Standardised transect counts of butterflies in old-growth rainforests in different biogeographical regions are lacking. Such data are needed to mitigate the influence of methodological and environmental factors within and between sites and, ultimately, to discriminate between long-term trends and short-term stochastic changes in abundance and community composition. 2. We compared butterfly assemblages using standardised Pollard Walks in the understory of closed-canopy lowland tropical rainforests across three biogeographical regions: Barro Colorado Island (BCI), Panama; Khao Chong (KHC), Thailand; and Wanang (WAN), Papua New Guinea. 3. The length and duration of transects, their spatial autocorrelation, and number of surveys per year represented important methodological factors that strongly influenced estimates of butterfly abundance. Of these, the effect of spatial autocorrelation was most difficult to mitigate across study sites. 4. Butterfly abundance and faunal composition were best explained by air temperature, elevation, rainfall, wind velocity, and human disturbance at BCI and KHC. In the absence of weather data at WAN, duration of transects and number of forest gaps accounted for most of the explained variance, which was rather low in all cases (<33%). 5. Adequate monitoring of the abundance of common butterflies was achieved at the 50 ha BCI plot, with three observers walking each of 10 transects of 500 m for 30 min each, during each of four surveys per year. These data may be standardised further after removing outliers of temperature and rainfall. Practical procedures are suggested to implement global monitoring of rainforest butterflies with Pollard Walks.

%B Insect Conservation and Diversity %G eng %0 Journal Article %J American Naturalist %D 2012 %T The Direct and Ecological Costs of an Ant-Plant Symbiosis %A Frederickson, M. E. %A Ravenscraft, A. %A Miller, G. A. %A Hernandez, L. M. A. %A Booth, G. %A Pierce, N. E. %K acacia %K ant-plant interactions %K castration %K cheating %K conflict %K cordia-nodosa %K costs of resistance %K Evolution %K evolution of mutualism %K extrafloral nectar production %K food bodies %K herbivory %K indirect plant defense %K mutualism %K selection %X

How strong is selection for cheating in mutualisms? The answer depends on the type and magnitude of the costs of the mutualism. Here we investigated the direct and ecological costs of plant defense by ants in the association between Cordia nodosa, a myr-mecophytic plant, and Allomerus octoarticulatus, a phytoecious ant. Cordia nodosa trees produce food and housing to reward ants that protect them against herbivores. For nearly 1 year, we manipulated the presence of A. octoarticulatus ants and most insect herbivores on C. nodosa in a full-factorial experiment. Ants increased plant growth when herbivores were present but decreased plant growth when herbivores were absent, indicating that hosting ants can be costly to plants. However, we did not detect a cost to ant colonies of defending host plants against herbivores. Although this asymmetry in costs suggests that the plants may be under stronger selection than the ants to cheat by withholding investment in their partner, the costs to C. nodosa are probably at least partly ecological, arising because ants tend scale insects on their host plants. We argue that ecological costs should favor resistance or traits other than cheating and thus that neither partner may face much temptation to cheat.

%B American Naturalist %V 179 %P 768-778 %8 Jun %@ 0003-0147 %G English %M WOS:000304354600010 %! Am NatAm Nat %0 Journal Article %J Cladistics %D 2012 %T Establishing criteria for higher-level classification using molecular data: the systematics of Polyommatusblue butterflies (Lepidoptera, Lycaenidae). %A Talavera, G %A Lukhtanov, V. A. %A Pierce, N. E. %A Villa, R. %X

Most taxonomists agree on the need to adapt current classifications to recognize monophyletic units. However, delineations between higher taxonomic units can be based on the relative ages of different lineages and ⁄or the level of morphological differentiation. In this paper, we address these issues in considering the species-rich Polyommatus section, a group of butterflies whose taxonomy has been highly controversial. We propose a taxonomy-friendly, flexible temporal scheme for higher-level classification. Using molecular data from nine markers (6666 bp) for 104 representatives of the Polyommatus section, representing all but two of the 81 described genera ⁄subgenera and five outgroups, we obtained a complete and well resolved phylogeny for this clade. We use this to revise the systematics of the Polyommatus blues, and to define criteria that best accommodate the described genera within a phylogenetic framework. First, we normalize the concept of section (Polyommatus) and propose the use of subtribe (Polyommatina) instead. To preserve taxonomic stability and traditionally recognized taxa, we designate an age interval (4–5 Myr) instead of a fixed minimum age to define genera. The application of these criteria results in the retention of 31 genera of the 81 formally described generic names, and necessitates the description of one new genus (Rueckbeilia gen. nov.). We note that while classifications should be based on phylogenetic data, applying a rigid universal scheme is rarely feasible. Ideally, taxon age limits should be applied according to the particularities and pre-existing taxonomy of each group. We demonstrate that the concept of a morphological gap may be misleading at the genus level and can produce polyphyletic genera, and we propose that recognition of the existence of cryptic genera may be useful in taxonomy.

%B Cladistics %P 1-27 %G eng %0 Journal Article %J Genome Biology and Evolution %D 2012 %T Genes Involved in the Evolution of Herbivory by a Leaf-Mining, Drosophilid Fly %A Whiteman, N. K. %A Gloss, A. D. %A Sackton, T. B. %A Groen, S. C. %A Humphrey, P. T. %A Lapoint, R. T. %A Sonderby, I. E. %A Halkier, B. A. %A Kocks, C. %A Ausubel, F. M. %A Pierce, N. E. %K arabidopsis %K cytochrome-p450 %K DNA-sequences %K Drosophila %K expression analysis %K genome %K glucosinolates %K herbivory %K host specialization %K induced resistance %K melanogaster %K molecular evolution %K phylogenetic analysis %K stress %K Transcriptome %X

Herbivorous insects are among the most successful radiations of life. However, we know little about the processes underpinning the evolution of herbivory. We examined the evolution of herbivory in the fly, Scaptomyza flava, whose larvae are leaf miners on species of Brassicaceae, including the widely studied reference plant, Arabidopsis thaliana (Arabidopsis). Scaptomyza flava is phylogenetically nested within the paraphyletic genus Drosophila, and the whole genome sequences available for 12 species of Drosophila facilitated phylogenetic analysis and assembly of a transcriptome for S. flava. A time-calibrated phylogeny indicated that leaf mining in Scaptomyza evolved between 6 and 16 million years ago. Feeding assays showed that biosynthesis of glucosinolates, the major class of antiherbivore chemical defense compounds in mustard leaves, was upregulated by S. flava larval feeding. The presence of glucosinolates in wild-type (WT) Arabidopsis plants reduced S. flava larval weight gain and increased egg-adult development time relative to flies reared in glucosinolate knockout (GKO) plants. An analysis of gene expression differences in 5-day-old larvae reared on WT versus GKO plants showed a total of 341 transcripts that were differentially regulated by glucosinolate uptake in larval S. flava. Of these, approximately a third corresponded to homologs of Drosophila melanogaster genes associated with starvation, dietary toxin-, heat-, oxidation-, and aging-related stress. The upregulated transcripts exhibited elevated rates of protein evolution compared with unregulated transcripts. The remaining differentially regulated transcripts also contained a higher proportion of novel genes than the unregulated transcripts. Thus, the transition to herbivory in Scaptomyza appears to be coupled with the evolution of novel genes and the co-option of conserved stress-related genes.

%B Genome Biology and Evolution %V 4 %P 900-916 %@ 1759-6653 %G English %M WOS:000313212900002 %! Genome Biol EvolGenome Biol Evol %0 Journal Article %J Plos One %D 2012 %T A Veritable Menagerie of Heritable Bacteria from Ants, Butterflies, and Beyond: Broad Molecular Surveys and a Systematic Review %A Russell, J. A. %A Funaro, C. F. %A Giraldo, Y. M. %A Goldman-Huertas, B. %A Suh, D. %A Kronauer, D. J. C. %A Moreau, C. S. %A Pierce, N. E. %K cytoplasmic incompatibility %K fungus-growing ants %K horizontal transmission %K hymenoptera-formicidae %K introduced populations %K phylogenetic analysis %K sex-ratio %K solenopsis-invicta %K symbiont infection %K wolbachia infections %X

Maternally transmitted bacteria have been important players in the evolution of insects and other arthropods, affecting their nutrition, defense, development, and reproduction. Wolbachia are the best studied among these and typically the most prevalent. While several other bacteria have independently evolved a heritable lifestyle, less is known about their host ranges. Moreover, most groups of insects have not had their heritable microflora systematically surveyed across a broad range of their taxonomic diversity. To help remedy these shortcomings we used diagnostic PCR to screen for five groups of heritable symbionts-Arsenophonus spp., Cardinium hertigii, Hamiltonella defensa, Spiroplasma spp., and Wolbachia spp.-across the ants and lepidopterans (focusing, in the latter case, on two butterfly families-the Lycaenidae and Nymphalidae). We did not detect Cardinium or Hamiltonella in any host. Wolbachia were the most widespread, while Spiroplasma (ants and lepidopterans) and Arsenophonus (ants only) were present at low levels. Co-infections with different Wolbachia strains appeared especially common in ants and less so in lepidopterans. While no additional facultative heritable symbionts were found among ants using universal bacterial primers, microbes related to heritable enteric bacteria were detected in several hosts. In summary, our findings show that Wolbachia are the dominant heritable symbionts of ants and at least some lepidopterans. However, a systematic review of symbiont frequencies across host taxa revealed that this is not always the case across other arthropods. Furthermore, comparisons of symbiont frequencies revealed that the prevalence of Wolbachia and other heritable symbionts varies substantially across lower-level arthropod taxa. We discuss the correlates, potential causes, and implications of these patterns, providing hypotheses on host attributes that may shape the distributions of these influential bacteria.

%B Plos One %V 7 %8 Dec 20 %@ 1932-6203 %G English %M WOS:000312794500019 %! Plos OnePlos One %0 Journal Article %J Molecular Ecology %D 2012 %T Asexual reproduction in introduced and native populations of the ant Cerapachys biroi %A Kronauer, D. J. C. %A Pierce, N. E. %A Keller, L. %K anthropogenic introduction %K argentine ant %K asexuality %K automixis %K clonal reproduction %K Evolution %K fire ant %K formicidae %K fungus-growing ant %K hymenoptera-formicidae %K invasion %K maximum-likelihood %K parthenogenesis %K platythyrea-punctata %K thelytocous-parthenogenesis %K thelytoky %X

Asexual reproduction is particularly common among introduced species, probably because it helps to overcome the negative effects associated with low population densities during colonization. The ant Cerapachys biroi has been introduced to tropical and subtropical islands around the world since the beginning of the last century. In this species, workers can reproduce via thelytokous parthenogenesis. Here, we use genetic markers to reconstruct the history of anthropogenic introductions of C.biroi, and to address the prevalence of female parthenogenesis in introduced and native populations. We show that at least four genetically distinct lineages have been introduced from continental Asia and have led to the species' circumtropical establishment. Our analyses demonstrate that asexual reproduction dominates in the introduced range and is also common in the native range. Given that C.biroi is the only dorylomorph ant that has successfully become established outside of its native range, this unusual mode of reproduction probably facilitated the species' worldwide spread. On the other hand, the rare occurrence of haploid males and at least one clear case of sexual recombination in the introduced range show that C.biroi has not lost the potential for sex. Finally, we show that thelytoky in C.biroi probably has a genetic rather than an infectious origin, and that automixis with central fusion is the most likely underlying cytological mechanism. This is in accordance with what is known for other thelytokous eusocial Hymenoptera.

%B Molecular Ecology %V 21 %P 5221-5235 %8 Nov %@ 0962-1083 %G English %M WOS:000310065600008 %! Mol EcolMol Ecol %0 Journal Article %J Insect Conservation and Diversity %D 2012 %T Cross-continental comparisons of butterfly assemblages in tropical rainforests: implications for biological monitoring %A Basset, Y %A Eastwood, R. %A L Sam %A DJ Lohman %A V Novotny %A T Treuer %A SE Miller %A GD Weiblen %A Pierce, N. E. %A Bunyavejchewin, S. %A W Sakchoowong %A P Kongnoo %A MA Osorio-Arenas %X

. 1. Standardised transect counts of butterflies in old-growth rainforests in different biogeographical regions are lacking. Such data are needed to mitigate the influence of methodological and environmental factors within and between sites and, ultimately, to discriminate between long-term trends and short-term stochastic changes in abundance and community composition. 2. We compared butterfly assemblages using standardised Pollard Walks in the understory of closed-canopy lowland tropical rainforests across three biogeographical regions: Barro Colorado Island (BCI), Panama; Khao Chong (KHC), Thailand; and Wanang (WAN), Papua New Guinea. 3. The length and duration of transects, their spatial autocorrelation, and number of surveys per year represented important methodological factors that strongly influenced estimates of butterfly abundance. Of these, the effect of spatial autocorrelation was most difficult to mitigate across study sites. 4. Butterfly abundance and faunal composition were best explained by air temperature, elevation, rainfall, wind velocity, and human disturbance at BCI and KHC. In the absence of weather data at WAN, duration of transects and number of forest gaps accounted for most of the explained variance, which was rather low in all cases (<33%). 5. Adequate monitoring of the abundance of common butterflies was achieved at the 50 ha BCI plot, with three observers walking each of 10 transects of 500 m for 30 min each, during each of four surveys per year. These data may be standardised further after removing outliers of temperature and rainfall. Practical procedures are suggested to implement global monitoring of rainforest butterflies with Pollard Walks.

%B Insect Conservation and Diversity %G eng %0 Journal Article %J American Naturalist %D 2012 %T The Direct and Ecological Costs of an Ant-Plant Symbiosis %A Frederickson, M. E. %A Ravenscraft, A. %A Miller, G. A. %A Hernandez, L. M. A. %A Booth, G. %A Pierce, N. E. %K acacia %K ant-plant interactions %K castration %K cheating %K conflict %K cordia-nodosa %K costs of resistance %K Evolution %K evolution of mutualism %K extrafloral nectar production %K food bodies %K herbivory %K indirect plant defense %K mutualism %K selection %X

How strong is selection for cheating in mutualisms? The answer depends on the type and magnitude of the costs of the mutualism. Here we investigated the direct and ecological costs of plant defense by ants in the association between Cordia nodosa, a myr-mecophytic plant, and Allomerus octoarticulatus, a phytoecious ant. Cordia nodosa trees produce food and housing to reward ants that protect them against herbivores. For nearly 1 year, we manipulated the presence of A. octoarticulatus ants and most insect herbivores on C. nodosa in a full-factorial experiment. Ants increased plant growth when herbivores were present but decreased plant growth when herbivores were absent, indicating that hosting ants can be costly to plants. However, we did not detect a cost to ant colonies of defending host plants against herbivores. Although this asymmetry in costs suggests that the plants may be under stronger selection than the ants to cheat by withholding investment in their partner, the costs to C. nodosa are probably at least partly ecological, arising because ants tend scale insects on their host plants. We argue that ecological costs should favor resistance or traits other than cheating and thus that neither partner may face much temptation to cheat.

%B American Naturalist %V 179 %P 768-778 %8 Jun %@ 0003-0147 %G English %M WOS:000304354600010 %! Am NatAm Nat %0 Journal Article %J Cladistics %D 2012 %T Establishing criteria for higher-level classification using molecular data: the systematics of Polyommatusblue butterflies (Lepidoptera, Lycaenidae). %A Talavera, G %A Lukhtanov, V. A. %A Pierce, N. E. %A Villa, R. %X

Most taxonomists agree on the need to adapt current classifications to recognize monophyletic units. However, delineations between higher taxonomic units can be based on the relative ages of different lineages and ⁄or the level of morphological differentiation. In this paper, we address these issues in considering the species-rich Polyommatus section, a group of butterflies whose taxonomy has been highly controversial. We propose a taxonomy-friendly, flexible temporal scheme for higher-level classification. Using molecular data from nine markers (6666 bp) for 104 representatives of the Polyommatus section, representing all but two of the 81 described genera ⁄subgenera and five outgroups, we obtained a complete and well resolved phylogeny for this clade. We use this to revise the systematics of the Polyommatus blues, and to define criteria that best accommodate the described genera within a phylogenetic framework. First, we normalize the concept of section (Polyommatus) and propose the use of subtribe (Polyommatina) instead. To preserve taxonomic stability and traditionally recognized taxa, we designate an age interval (4–5 Myr) instead of a fixed minimum age to define genera. The application of these criteria results in the retention of 31 genera of the 81 formally described generic names, and necessitates the description of one new genus (Rueckbeilia gen. nov.). We note that while classifications should be based on phylogenetic data, applying a rigid universal scheme is rarely feasible. Ideally, taxon age limits should be applied according to the particularities and pre-existing taxonomy of each group. We demonstrate that the concept of a morphological gap may be misleading at the genus level and can produce polyphyletic genera, and we propose that recognition of the existence of cryptic genera may be useful in taxonomy.

%B Cladistics %P 1-27 %G eng %0 Journal Article %J Genome Biology and Evolution %D 2012 %T Genes Involved in the Evolution of Herbivory by a Leaf-Mining, Drosophilid Fly %A Whiteman, N. K. %A Gloss, A. D. %A Sackton, T. B. %A Groen, S. C. %A Humphrey, P. T. %A Lapoint, R. T. %A Sonderby, I. E. %A Halkier, B. A. %A Kocks, C. %A Ausubel, F. M. %A Pierce, N. E. %K arabidopsis %K cytochrome-p450 %K DNA-sequences %K Drosophila %K expression analysis %K genome %K glucosinolates %K herbivory %K host specialization %K induced resistance %K melanogaster %K molecular evolution %K phylogenetic analysis %K stress %K Transcriptome %X

Herbivorous insects are among the most successful radiations of life. However, we know little about the processes underpinning the evolution of herbivory. We examined the evolution of herbivory in the fly, Scaptomyza flava, whose larvae are leaf miners on species of Brassicaceae, including the widely studied reference plant, Arabidopsis thaliana (Arabidopsis). Scaptomyza flava is phylogenetically nested within the paraphyletic genus Drosophila, and the whole genome sequences available for 12 species of Drosophila facilitated phylogenetic analysis and assembly of a transcriptome for S. flava. A time-calibrated phylogeny indicated that leaf mining in Scaptomyza evolved between 6 and 16 million years ago. Feeding assays showed that biosynthesis of glucosinolates, the major class of antiherbivore chemical defense compounds in mustard leaves, was upregulated by S. flava larval feeding. The presence of glucosinolates in wild-type (WT) Arabidopsis plants reduced S. flava larval weight gain and increased egg-adult development time relative to flies reared in glucosinolate knockout (GKO) plants. An analysis of gene expression differences in 5-day-old larvae reared on WT versus GKO plants showed a total of 341 transcripts that were differentially regulated by glucosinolate uptake in larval S. flava. Of these, approximately a third corresponded to homologs of Drosophila melanogaster genes associated with starvation, dietary toxin-, heat-, oxidation-, and aging-related stress. The upregulated transcripts exhibited elevated rates of protein evolution compared with unregulated transcripts. The remaining differentially regulated transcripts also contained a higher proportion of novel genes than the unregulated transcripts. Thus, the transition to herbivory in Scaptomyza appears to be coupled with the evolution of novel genes and the co-option of conserved stress-related genes.

%B Genome Biology and Evolution %V 4 %P 900-916 %@ 1759-6653 %G English %M WOS:000313212900002 %! Genome Biol EvolGenome Biol Evol %0 Journal Article %J Plos One %D 2012 %T A Veritable Menagerie of Heritable Bacteria from Ants, Butterflies, and Beyond: Broad Molecular Surveys and a Systematic Review %A Russell, J. A. %A Funaro, C. F. %A Giraldo, Y. M. %A Goldman-Huertas, B. %A Suh, D. %A Kronauer, D. J. C. %A Moreau, C. S. %A Pierce, N. E. %K cytoplasmic incompatibility %K fungus-growing ants %K horizontal transmission %K hymenoptera-formicidae %K introduced populations %K phylogenetic analysis %K sex-ratio %K solenopsis-invicta %K symbiont infection %K wolbachia infections %X

Maternally transmitted bacteria have been important players in the evolution of insects and other arthropods, affecting their nutrition, defense, development, and reproduction. Wolbachia are the best studied among these and typically the most prevalent. While several other bacteria have independently evolved a heritable lifestyle, less is known about their host ranges. Moreover, most groups of insects have not had their heritable microflora systematically surveyed across a broad range of their taxonomic diversity. To help remedy these shortcomings we used diagnostic PCR to screen for five groups of heritable symbionts-Arsenophonus spp., Cardinium hertigii, Hamiltonella defensa, Spiroplasma spp., and Wolbachia spp.-across the ants and lepidopterans (focusing, in the latter case, on two butterfly families-the Lycaenidae and Nymphalidae). We did not detect Cardinium or Hamiltonella in any host. Wolbachia were the most widespread, while Spiroplasma (ants and lepidopterans) and Arsenophonus (ants only) were present at low levels. Co-infections with different Wolbachia strains appeared especially common in ants and less so in lepidopterans. While no additional facultative heritable symbionts were found among ants using universal bacterial primers, microbes related to heritable enteric bacteria were detected in several hosts. In summary, our findings show that Wolbachia are the dominant heritable symbionts of ants and at least some lepidopterans. However, a systematic review of symbiont frequencies across host taxa revealed that this is not always the case across other arthropods. Furthermore, comparisons of symbiont frequencies revealed that the prevalence of Wolbachia and other heritable symbionts varies substantially across lower-level arthropod taxa. We discuss the correlates, potential causes, and implications of these patterns, providing hypotheses on host attributes that may shape the distributions of these influential bacteria.

%B Plos One %V 7 %8 Dec 20 %@ 1932-6203 %G English %M WOS:000312794500019 %! Plos OnePlos One %0 Journal Article %J Applied and Environmental Microbiology %D 2011 %T Army Ants Harbor a Host-Specific Clade of Entomoplasmatales Bacteria %A Funaro, C. F. %A Kronauer, D. J. C. %A Moreau, C. S. %A Goldman-Huertas, B. %A Pierce, N. E. %A Russell, J. A. %K Community structure %K diversity %K Drosophila %K fungus-growing ants %K hymenoptera-formicidae %K molecular-identification %K parasite relationships %K sp. nov %K spiroplasma %K symbionts %X

In this article, we describe the distributions of Entomoplasmatales bacteria across the ants, identifying a novel lineage of gut bacteria that is unique to the army ants. While our findings indicate that the Entomoplasmatales are not essential for growth or development, molecular analyses suggest that this relationship is host specific and potentially ancient. The documented trends add to a growing body of literature that hints at a diversity of undiscovered associations between ants and bacterial symbionts.

%B Applied and Environmental Microbiology %V 77 %P 346-350 %8 Jan %@ 0099-2240 %G English %M WOS:000285550300041 %! Appl Environ MicrobAppl Environ Microb %0 Journal Article %J Science %D 2011 %T Asynchronous Diversification in a Specialized Plant-Pollinator Mutualism %A Ramirez, S. R. %A Eltz, T. %A Fujiwara, M. K. %A Gerlach, G. %A Goldman-Huertas, B. %A Tsutsui, N. D. %A Pierce, N. E. %K epidendroideae %K euglossine bees %K Evolution %K Insects %K models %K networks %K orchid bees %X

Most flowering plants establish mutualistic associations with insect pollinators to facilitate sexual reproduction. However, the evolutionary processes that gave rise to these associations remain poorly understood. We reconstructed the times of divergence, diversification patterns, and interaction networks of a diverse group of specialized orchids and their bee pollinators. In contrast to a scenario of coevolution by race formation, we show that fragrance-producing orchids originated at least three times independently after their fragrance-collecting bee mutualists. Whereas orchid diversification has apparently tracked the diversification of orchids' bee pollinators, bees appear to have depended on the diverse chemical environment of neotropical forests. We corroborated this apparent asymmetrical dependency by simulating co-extinction cascades in real interaction networks that lacked reciprocal specialization. These results suggest that the diversification of insect-pollinated angiosperms may have been facilitated by the exploitation of preexisting sensory biases of insect pollinators.

%B Science %V 333 %P 1742-1746 %8 Sep 23 %@ 0036-8075 %G English %M WOS:000295121500041 %! ScienceScience %0 Journal Article %J Ecology Letters %D 2011 %T Economic game theory for mutualism and cooperation %A Archetti, M. %A Scheuring, I. %A Hoffman, M. %A Frederickson, M. E. %A Pierce, N. E. %A Yu, D. W. %K coevolution %K collective action %K common-pool resource %K contract theory %K cooperation %K Game Theory %K host sanctions %K legume-rhizobium mutualism %K mutualism %K n-person prisoner's dilemma %K partner choice %K prisoners-dilemma %K public goods %K public-goods games %K reproductive success %K social dilemmas %K symbiosis %K volunteer's dilemma %K volunteers dilemma %X

We review recent work at the interface of economic game theory and evolutionary biology that provides new insights into the evolution of partner choice, host sanctions, partner fidelity feedback and public goods. (1) The theory of games with asymmetrical information shows that the right incentives allow hosts to screen-out parasites and screen-in mutualists, explaining successful partner choice in the absence of signalling. Applications range from ant-plants to microbiomes. (2) Contract theory distinguishes two longstanding but weakly differentiated explanations of host response to defectors: host sanctions and partner fidelity feedback. Host traits that selectively punish misbehaving symbionts are parsimoniously interpreted as pre-adaptations. Yucca-moth and legume-rhizobia mutualisms are argued to be examples of partner fidelity feedback. (3) The theory of public goods shows that cooperation in multi-player interactions can evolve in the absence of assortment, in one-shot social dilemmas among non-kin. Applications include alarm calls in vertebrates and exoenzymes in microbes.

%B Ecology Letters %V 14 %P 1300-1312 %8 Dec %@ 1461-023X %G English %M WOS:000297147500013 %! Ecol LettEcol Lett %0 Journal Article %J American Naturalist %D 2011 %T Let the Right One In: A Microeconomic Approach to Partner Choice in Mutualisms %A Archetti, M. %A Ubeda, F. %A Fudenberg, D. %A Green, J. %A Pierce, N. E. %A Yu, D. W. %K ant-plant mutualism %K cooperation %K euprymna-scolopes %K fischeri %K host %K light organ %K microbiome %K myrmecophytes %K parasitism %K species coexistence %K specificity %K squid-vibrio symbiosis %K symbiosis %K vibrio fischeri %K Virulence %K volunteers dilemma %X

One of the main problems impeding the evolution of cooperation is partner choice. When information is asymmetric (the quality of a potential partner is known only to himself), it may seem that partner choice is not possible without signaling. Many mutualisms, however, exist without signaling, and the mechanisms by which hosts might select the right partners are unclear. Here we propose a general mechanism of partner choice, "screening," that is similar to the economic theory of mechanism design. Imposing the appropriate costs and rewards may induce the informed individuals to screen themselves according to their types and therefore allow a noninformed individual to establish associations with the correct partners in the absence of signaling. Several types of biological symbioses are good candidates for screening, including bobtail squid, ant-plants, gut microbiomes, and many animal and plant species that produce reactive oxygen species. We describe a series of diagnostic tests for screening. Screening games can apply to the cases where by-products, partner fidelity feedback, or host sanctions do not apply, therefore explaining the evolution of mutualism in systems where it is impossible for potential symbionts to signal their cooperativeness beforehand and where the host does not punish symbiont misbehavior.

%B American Naturalist %V 177 %P 75-85 %8 Jan %@ 0003-0147 %G English %M WOS:000285136700010 %! Am NatAm Nat %0 Journal Article %J Molecular Ecology %D 2011 %T Mining the plant-herbivore interface with a leafmining Drosophila of Arabidopsis %A Whiteman, N. K. %A Groen, S. C. %A Chevasco, D. %A Bear, A. %A Beckwith, N. %A Gregory, T. R. %A Denoux, C. %A Mammarella, N. %A Ausubel, F. M. %A Pierce, N. E. %K arabidopsis %K cardamine-cordifolia gray,a. %K defense responses %K Drosophila %K Gene Expression %K hawaiian drosophilidae %K herbivory %K host plants %K insect interactions %K jasmonate %K larval performance %K model system %K molecular-patterns %K native crucifer %K parkinsons-disease %K pseudomonas-syringae %X

Experimental infections of Arabidopsis thaliana (Arabidopsis) with genomically characterized plant pathogens such as Pseudomonas syringae have facilitated the dissection of canonical eukaryotic defence pathways and parasite virulence factors. Plants are also attacked by herbivorous insects, and the development of an ecologically relevant genetic model herbivore that feeds on Arabidopsis will enable the parallel dissection of host defence and reciprocal resistance pathways such as those involved in xenobiotic metabolism. An ideal candidate is Scaptomyza flava, a drosophilid fly whose leafmining larvae are true herbivores that can be found in nature feeding on Arabidopsis and other crucifers. Here, we describe the life cycle of S. flava on Arabidopsis and use multiple approaches to characterize the response of Arabidopsis to S. flava attack. Oviposition choice tests and growth performance assays on different Arabidopsis ecotypes, defence-related mutants, and hormone and chitin-treated plants revealed significant differences in host preference and variation in larval performance across Arabidopsis accessions. The jasmonate and glucosinolate pathways in Arabidopsis are important in mediating quantitative resistance against S. flava, and priming with jasmonate or chitin resulted in increased resistance. Expression of xenobiotic detoxification genes was reduced in S. flava larvae reared on Arabidopsis jasmonate signalling mutants and increased in plants pretreated with chitin. These results and future research directions are discussed in the context of developing a genetic model system to analyse insect-plant interactions.

%B Molecular Ecology %V 20 %P 995-1014 %8 Mar %@ 0962-1083 %G English %M WOS:000287397600013 %! Mol EcolMol Ecol %0 Journal Article %J Current Biology %D 2011 %T Myrmecophiles %A Kronauer, D. J. C. %A Pierce, N. E. %B Current Biology %V 21 %P R208-R209 %8 Mar 22 %@ 0960-9822 %G English %M WOS:000288826100004 %! Curr BiolCurr Biol %0 Journal Article %J Conservation Genetics Resources %D 2011 %T Nine novel microsatellite markers for the army ant Simopelta pergandei (subfamily Ponerinae) %A Kronauer, D. J. C. %A Boomsma, J. J. %A Pierce, N. E. %K eciton-burchellii %K Evolution %K formicidae %K genetic marker %K microsatellite %K Population genetics %K sociobiology %X

Simopelta (subfamily Ponerinae) army ants are specialized predators of other ants in New World tropical forests. Although they show a striking convergence in overall life-history with the well known army ants of the subfamilies Aenictinae, Dorylinae, and Ecitoninae, the genus has been little studied. We developed and characterized nine novel microsatellite loci for S. pergandei with 2-8 observed alleles (mean: 5.2) and expected heterozygosities between 0.16 and 0.87 (mean: 0.68). Three of these loci reliably cross-amplified in a second species, S. pentadentata, with 4-8 alleles (mean: 8.0) and expected heterozygosities between 0.32 and 0.85 (mean: 0.65). These genetic markers will be useful in studying the sociobiology and molecular ecology of Simopelta army ants and in elucidating convergent evolutionary trajectories that have culminated in the army ant lifestyle.

%B Conservation Genetics Resources %V 3 %P 61-63 %8 Jan %@ 1877-7252 %G English %M WOS:000290438500016 %! Conserv Genet ResourConserv Genet Resour %0 Journal Article %J Proceedings of the Royal Society B-Biological Sciences %D 2011 %T Phylogeny and palaeoecology of Polyommatus blue butterflies show Beringia was a climate-regulated gateway to the New World %A Vila, R. %A Bell, C. D. %A Macniven, R. %A Goldman-Huertas, B. %A Ree, R. H. %A Marshall, C. R. %A Balint, Z. %A Johnson, K. %A Benyamini, D. %A Pierce, N. E. %K beringia %K biogeography %K climate change %K conservatism %K Dispersal %K diversification %K Evolution %K historical biogeography %K lepidoptera lycaenidae %K lycaenidae %K nabokov %K northern-hemisphere %K patterns %K Phylogeny %K species distribution %K vicariance analysis %X

Transcontinental dispersals by organisms usually represent improbable events that constitute a major challenge for biogeographers. By integrating molecular phylogeny, historical biogeography and palaeoecology, we test a bold hypothesis proposed by Vladimir Nabokov regarding the origin of Neotropical Polyommatus blue butterflies, and show that Beringia has served as a biological corridor for the dispersal of these insects from Asia into the New World. We present a novel method to estimate ancestral temperature tolerances using distribution range limits of extant organisms, and find that climatic conditions in Beringia acted as a decisive filter in determining which taxa crossed into the New World during five separate invasions over the past 11 Myr. Our results reveal a marked effect of the Miocene-Pleistocene global cooling, and demonstrate that palaeoclimatic conditions left a strong signal on the ecology of present-day taxa in the New World. The phylogenetic conservatism in thermal tolerances that we have identified may permit the reconstruction of the palaeoecology of ancestral organisms, especially mobile taxa that can easily escape from hostile environments rather than adapt to them.

%B Proceedings of the Royal Society B-Biological Sciences %V 278 %P 2737-2744 %8 Sep 22 %@ 0962-8452 %G English %M WOS:000293733600006 %! P Roy Soc B-Biol SciP Roy Soc B-Biol Sci %0 Journal Article %J Proceedings of the National Academy of Sciences of the United States of America %D 2011 %T Reply to Kiers et al.: Economic and biological clarity in the theory of mutualism %A Weyl, E. G. %A Frederickson, M. E. %A Yu, D. W. %A Pierce, N. E. %K host sanctions %B Proceedings of the National Academy of Sciences of the United States of America %V 108 %P E8-E8 %8 Jan 18 %@ 0027-8424 %G English %M WOS:000286310300002 %! P Natl Acad Sci USAP Natl Acad Sci USA %0 Journal Article %J Molecular Ecology %D 2011 %T Strict monandry in the ponerine army ant genus Simopelta suggests that colony size and complexity drive mating system evolution in social insects %A Kronauer, D. J. C. %A O'Donnell, S. %A Boomsma, J. J. %A Pierce, N. E. %K allele homo-heterozygosity %K bees apis-mellifera %K colony fission %K eusocial hymenoptera %K eusociality %K formicidae %K honey production %K kin selection %K male parentage %K mating frequency %K microsatellites %K paternity frequency %K sex determination %K stingless bees %K worker reproduction %X

Altruism in social insects has evolved between closely related full-siblings. It is therefore of considerable interest why some groups have secondarily evolved low within-colony relatedness, which in turn affects the relatedness incentives of within-colony cooperation and conflict. The highest queen mating frequencies, and therefore among the lowest degrees of colony relatedness, occur in Apis honeybees and army ants of the subfamilies Aenictinae, Ecitoninae, and Dorylinae, suggesting that common life history features such as reproduction by colony fission and male biased numerical sex-ratios have convergently shaped these mating systems. Here we show that ponerine army ants of the genus Simopelta, which are distantly related but similar in general biology to other army ants, have strictly monandrous queens. Preliminary data suggest that workers reproduce in queenright colonies, which is in sharp contrast to other army ants. We hypothesize that differences in mature colony size and social complexity may explain these striking discrepancies.

%B Molecular Ecology %V 20 %P 420-428 %8 Jan %@ 0962-1083 %G English %M WOS:000285970200019 %! Mol EcolMol Ecol %0 Journal Article %J Applied and Environmental Microbiology %D 2011 %T Army Ants Harbor a Host-Specific Clade of Entomoplasmatales Bacteria %A Funaro, C. F. %A Kronauer, D. J. C. %A Moreau, C. S. %A Goldman-Huertas, B. %A Pierce, N. E. %A Russell, J. A. %K Community structure %K diversity %K Drosophila %K fungus-growing ants %K hymenoptera-formicidae %K molecular-identification %K parasite relationships %K sp. nov %K spiroplasma %K symbionts %X

In this article, we describe the distributions of Entomoplasmatales bacteria across the ants, identifying a novel lineage of gut bacteria that is unique to the army ants. While our findings indicate that the Entomoplasmatales are not essential for growth or development, molecular analyses suggest that this relationship is host specific and potentially ancient. The documented trends add to a growing body of literature that hints at a diversity of undiscovered associations between ants and bacterial symbionts.

%B Applied and Environmental Microbiology %V 77 %P 346-350 %8 Jan %@ 0099-2240 %G English %M WOS:000285550300041 %! Appl Environ MicrobAppl Environ Microb %0 Journal Article %J Science %D 2011 %T Asynchronous Diversification in a Specialized Plant-Pollinator Mutualism %A Ramirez, S. R. %A Eltz, T. %A Fujiwara, M. K. %A Gerlach, G. %A Goldman-Huertas, B. %A Tsutsui, N. D. %A Pierce, N. E. %K epidendroideae %K euglossine bees %K Evolution %K Insects %K models %K networks %K orchid bees %X

Most flowering plants establish mutualistic associations with insect pollinators to facilitate sexual reproduction. However, the evolutionary processes that gave rise to these associations remain poorly understood. We reconstructed the times of divergence, diversification patterns, and interaction networks of a diverse group of specialized orchids and their bee pollinators. In contrast to a scenario of coevolution by race formation, we show that fragrance-producing orchids originated at least three times independently after their fragrance-collecting bee mutualists. Whereas orchid diversification has apparently tracked the diversification of orchids' bee pollinators, bees appear to have depended on the diverse chemical environment of neotropical forests. We corroborated this apparent asymmetrical dependency by simulating co-extinction cascades in real interaction networks that lacked reciprocal specialization. These results suggest that the diversification of insect-pollinated angiosperms may have been facilitated by the exploitation of preexisting sensory biases of insect pollinators.

%B Science %V 333 %P 1742-1746 %8 Sep 23 %@ 0036-8075 %G English %M WOS:000295121500041 %! ScienceScience %0 Journal Article %J Ecology Letters %D 2011 %T Economic game theory for mutualism and cooperation %A Archetti, M. %A Scheuring, I. %A Hoffman, M. %A Frederickson, M. E. %A Pierce, N. E. %A Yu, D. W. %K coevolution %K collective action %K common-pool resource %K contract theory %K cooperation %K Game Theory %K host sanctions %K legume-rhizobium mutualism %K mutualism %K n-person prisoner's dilemma %K partner choice %K prisoners-dilemma %K public goods %K public-goods games %K reproductive success %K social dilemmas %K symbiosis %K volunteer's dilemma %K volunteers dilemma %X

We review recent work at the interface of economic game theory and evolutionary biology that provides new insights into the evolution of partner choice, host sanctions, partner fidelity feedback and public goods. (1) The theory of games with asymmetrical information shows that the right incentives allow hosts to screen-out parasites and screen-in mutualists, explaining successful partner choice in the absence of signalling. Applications range from ant-plants to microbiomes. (2) Contract theory distinguishes two longstanding but weakly differentiated explanations of host response to defectors: host sanctions and partner fidelity feedback. Host traits that selectively punish misbehaving symbionts are parsimoniously interpreted as pre-adaptations. Yucca-moth and legume-rhizobia mutualisms are argued to be examples of partner fidelity feedback. (3) The theory of public goods shows that cooperation in multi-player interactions can evolve in the absence of assortment, in one-shot social dilemmas among non-kin. Applications include alarm calls in vertebrates and exoenzymes in microbes.

%B Ecology Letters %V 14 %P 1300-1312 %8 Dec %@ 1461-023X %G English %M WOS:000297147500013 %! Ecol LettEcol Lett %0 Book Section %B Losos J (ed) In the Light of Evolution: Essays From the Laboratory and Field %D 2011 %T The herbivore's dilemma: Never enough nitrogen %A Pierce, N. E. %A Berry, A. J. %B Losos J (ed) In the Light of Evolution: Essays From the Laboratory and Field %7 1st %I Roberts & Company %P 121-134 %G eng %0 Journal Article %J American Naturalist %D 2011 %T Let the Right One In: A Microeconomic Approach to Partner Choice in Mutualisms %A Archetti, M. %A Ubeda, F. %A Fudenberg, D. %A Green, J. %A Pierce, N. E. %A Yu, D. W. %K ant-plant mutualism %K cooperation %K euprymna-scolopes %K fischeri %K host %K light organ %K microbiome %K myrmecophytes %K parasitism %K species coexistence %K specificity %K squid-vibrio symbiosis %K symbiosis %K vibrio fischeri %K Virulence %K volunteers dilemma %X

One of the main problems impeding the evolution of cooperation is partner choice. When information is asymmetric (the quality of a potential partner is known only to himself), it may seem that partner choice is not possible without signaling. Many mutualisms, however, exist without signaling, and the mechanisms by which hosts might select the right partners are unclear. Here we propose a general mechanism of partner choice, "screening," that is similar to the economic theory of mechanism design. Imposing the appropriate costs and rewards may induce the informed individuals to screen themselves according to their types and therefore allow a noninformed individual to establish associations with the correct partners in the absence of signaling. Several types of biological symbioses are good candidates for screening, including bobtail squid, ant-plants, gut microbiomes, and many animal and plant species that produce reactive oxygen species. We describe a series of diagnostic tests for screening. Screening games can apply to the cases where by-products, partner fidelity feedback, or host sanctions do not apply, therefore explaining the evolution of mutualism in systems where it is impossible for potential symbionts to signal their cooperativeness beforehand and where the host does not punish symbiont misbehavior.

%B American Naturalist %V 177 %P 75-85 %8 Jan %@ 0003-0147 %G English %M WOS:000285136700010 %! Am NatAm Nat %0 Journal Article %J Molecular Ecology %D 2011 %T Mining the plant-herbivore interface with a leafmining Drosophila of Arabidopsis %A Whiteman, N. K. %A Groen, S. C. %A Chevasco, D. %A Bear, A. %A Beckwith, N. %A Gregory, T. R. %A Denoux, C. %A Mammarella, N. %A Ausubel, F. M. %A Pierce, N. E. %K arabidopsis %K cardamine-cordifolia gray,a. %K defense responses %K Drosophila %K Gene Expression %K hawaiian drosophilidae %K herbivory %K host plants %K insect interactions %K jasmonate %K larval performance %K model system %K molecular-patterns %K native crucifer %K parkinsons-disease %K pseudomonas-syringae %X

Experimental infections of Arabidopsis thaliana (Arabidopsis) with genomically characterized plant pathogens such as Pseudomonas syringae have facilitated the dissection of canonical eukaryotic defence pathways and parasite virulence factors. Plants are also attacked by herbivorous insects, and the development of an ecologically relevant genetic model herbivore that feeds on Arabidopsis will enable the parallel dissection of host defence and reciprocal resistance pathways such as those involved in xenobiotic metabolism. An ideal candidate is Scaptomyza flava, a drosophilid fly whose leafmining larvae are true herbivores that can be found in nature feeding on Arabidopsis and other crucifers. Here, we describe the life cycle of S. flava on Arabidopsis and use multiple approaches to characterize the response of Arabidopsis to S. flava attack. Oviposition choice tests and growth performance assays on different Arabidopsis ecotypes, defence-related mutants, and hormone and chitin-treated plants revealed significant differences in host preference and variation in larval performance across Arabidopsis accessions. The jasmonate and glucosinolate pathways in Arabidopsis are important in mediating quantitative resistance against S. flava, and priming with jasmonate or chitin resulted in increased resistance. Expression of xenobiotic detoxification genes was reduced in S. flava larvae reared on Arabidopsis jasmonate signalling mutants and increased in plants pretreated with chitin. These results and future research directions are discussed in the context of developing a genetic model system to analyse insect-plant interactions.

%B Molecular Ecology %V 20 %P 995-1014 %8 Mar %@ 0962-1083 %G English %M WOS:000287397600013 %! Mol EcolMol Ecol %0 Journal Article %J Current Biology %D 2011 %T Myrmecophiles %A Kronauer, D. J. C. %A Pierce, N. E. %B Current Biology %V 21 %P R208-R209 %8 Mar 22 %@ 0960-9822 %G English %M WOS:000288826100004 %! Curr BiolCurr Biol %0 Journal Article %J Conservation Genetics Resources %D 2011 %T Nine novel microsatellite markers for the army ant Simopelta pergandei (subfamily Ponerinae) %A Kronauer, D. J. C. %A Boomsma, J. J. %A Pierce, N. E. %K eciton-burchellii %K Evolution %K formicidae %K genetic marker %K microsatellite %K Population genetics %K sociobiology %X

Simopelta (subfamily Ponerinae) army ants are specialized predators of other ants in New World tropical forests. Although they show a striking convergence in overall life-history with the well known army ants of the subfamilies Aenictinae, Dorylinae, and Ecitoninae, the genus has been little studied. We developed and characterized nine novel microsatellite loci for S. pergandei with 2-8 observed alleles (mean: 5.2) and expected heterozygosities between 0.16 and 0.87 (mean: 0.68). Three of these loci reliably cross-amplified in a second species, S. pentadentata, with 4-8 alleles (mean: 8.0) and expected heterozygosities between 0.32 and 0.85 (mean: 0.65). These genetic markers will be useful in studying the sociobiology and molecular ecology of Simopelta army ants and in elucidating convergent evolutionary trajectories that have culminated in the army ant lifestyle.

%B Conservation Genetics Resources %V 3 %P 61-63 %8 Jan %@ 1877-7252 %G English %M WOS:000290438500016 %! Conserv Genet ResourConserv Genet Resour %0 Journal Article %J Molecular Phylogenetics and Evolution %D 2011 %T A phylogenetic revision of the Glaucopsyche section (Lepidoptera: Lycaenidae), with special focus on the Phengaris-Maculinea clade %A Ugelvig, L. V. %A Vila, R. %A Pierce, N. E. %A Nash, D. R. %K ants %K bayesian inference %K butterflies %K characters %K coevolution %K Europe %K Evolution %K fauna %K inference %K lycaenid butterflies %K maximum likelihood %K parasitic caterpillars %K Phylogeny %K selection %X

Despite much research on the socially parasitic large blue butterflies (genus Maculinea) in the past 40 years, their relationship to their closest relatives, Phengaris, is controversial and the relationships among the remaining genera in the Glaucopsyche section are largely unresolved. The evolutionary history of this butterfly section is particularly important to understand the evolution of life history diversity connected to food-plant and host-ant associations in the larval stage. In the present study, we use a combination of four nuclear and two mitochondrial genes to reconstruct the phylogeny of the Glaucopsyche section, and in particular, to study the relationships among and within the Phengaris-Maculinea species.We find a clear pattern between the clades recovered in the Glaucopsyche section phylogeny and their food-plant associations, with only the Phengaris-Maculinea clade utilising more than one plant family. Maculinea is, for the first time, recovered with strong support as a monophyletic group nested within Phengaris, with the closest relative being the rare genus Caerulea. The genus Glaucopsyche is polyphyletic, including the genera Sinia and lolana. Interestingly, we find evidence for additional potential cryptic species within the highly endangered Maculinea, which has long been suspected from morphological, ecological and molecular studies. (C) 2011 Elsevier Inc. All rights reserved.

%B Molecular Phylogenetics and Evolution %V 61 %P 237-243 %8 Oct %@ 1055-7903 %G English %M WOS:000293810000020 %! Mol Phylogenet EvolMol Phylogenet Evol %0 Journal Article %J Proceedings of the Royal Society B-Biological Sciences %D 2011 %T Phylogeny and palaeoecology of Polyommatus blue butterflies show Beringia was a climate-regulated gateway to the New World %A Vila, R. %A Bell, C. D. %A Macniven, R. %A Goldman-Huertas, B. %A Ree, R. H. %A Marshall, C. R. %A Balint, Z. %A Johnson, K. %A Benyamini, D. %A Pierce, N. E. %K beringia %K biogeography %K climate change %K conservatism %K Dispersal %K diversification %K Evolution %K historical biogeography %K lepidoptera lycaenidae %K lycaenidae %K nabokov %K northern-hemisphere %K patterns %K Phylogeny %K species distribution %K vicariance analysis %X

Transcontinental dispersals by organisms usually represent improbable events that constitute a major challenge for biogeographers. By integrating molecular phylogeny, historical biogeography and palaeoecology, we test a bold hypothesis proposed by Vladimir Nabokov regarding the origin of Neotropical Polyommatus blue butterflies, and show that Beringia has served as a biological corridor for the dispersal of these insects from Asia into the New World. We present a novel method to estimate ancestral temperature tolerances using distribution range limits of extant organisms, and find that climatic conditions in Beringia acted as a decisive filter in determining which taxa crossed into the New World during five separate invasions over the past 11 Myr. Our results reveal a marked effect of the Miocene-Pleistocene global cooling, and demonstrate that palaeoclimatic conditions left a strong signal on the ecology of present-day taxa in the New World. The phylogenetic conservatism in thermal tolerances that we have identified may permit the reconstruction of the palaeoecology of ancestral organisms, especially mobile taxa that can easily escape from hostile environments rather than adapt to them.

%B Proceedings of the Royal Society B-Biological Sciences %V 278 %P 2737-2744 %8 Sep 22 %@ 0962-8452 %G English %M WOS:000293733600006 %! P Roy Soc B-Biol SciP Roy Soc B-Biol Sci %0 Journal Article %J Proceedings of the National Academy of Sciences of the United States of America %D 2011 %T Reply to Kiers et al.: Economic and biological clarity in the theory of mutualism %A Weyl, E. G. %A Frederickson, M. E. %A Yu, D. W. %A Pierce, N. E. %K host sanctions %B Proceedings of the National Academy of Sciences of the United States of America %V 108 %P E8-E8 %8 Jan 18 %@ 0027-8424 %G English %M WOS:000286310300002 %! P Natl Acad Sci USAP Natl Acad Sci USA %0 Journal Article %J Molecular Ecology %D 2011 %T Strict monandry in the ponerine army ant genus Simopelta suggests that colony size and complexity drive mating system evolution in social insects %A Kronauer, D. J. C. %A O'Donnell, S. %A Boomsma, J. J. %A Pierce, N. E. %K allele homo-heterozygosity %K bees apis-mellifera %K colony fission %K eusocial hymenoptera %K eusociality %K formicidae %K honey production %K kin selection %K male parentage %K mating frequency %K microsatellites %K paternity frequency %K sex determination %K stingless bees %K worker reproduction %X

Altruism in social insects has evolved between closely related full-siblings. It is therefore of considerable interest why some groups have secondarily evolved low within-colony relatedness, which in turn affects the relatedness incentives of within-colony cooperation and conflict. The highest queen mating frequencies, and therefore among the lowest degrees of colony relatedness, occur in Apis honeybees and army ants of the subfamilies Aenictinae, Ecitoninae, and Dorylinae, suggesting that common life history features such as reproduction by colony fission and male biased numerical sex-ratios have convergently shaped these mating systems. Here we show that ponerine army ants of the genus Simopelta, which are distantly related but similar in general biology to other army ants, have strictly monandrous queens. Preliminary data suggest that workers reproduce in queenright colonies, which is in sharp contrast to other army ants. We hypothesize that differences in mature colony size and social complexity may explain these striking discrepancies.

%B Molecular Ecology %V 20 %P 420-428 %8 Jan %@ 0962-1083 %G English %M WOS:000285970200019 %! Mol EcolMol Ecol %0 Journal Article %J Proceedings of the National Academy of Sciences %D 2010 %T Economic contract theory tests models of mutualism. %A Weyl, EG %A ME Frederickson %A Yu, D. W. %A Pierce, N. E. %X

Although mutualisms are common in all ecological communities and have played key roles in the diversification of life, our current understanding of the evolution of cooperation applies mostly to social behavior within a species. A central question is whether mutualisms persist because hosts have evolved costly punishment of cheaters. Here, we use the economic theory of employment contracts to formulate and distinguish between two mechanisms that have been proposed to prevent cheating in host–symbiont mutualisms, partner fidelity feedback (PFF) and host sanctions (HS). Under PFF, positive feedback between host fitness and symbiont fitness is sufficient to prevent cheating; in contrast, HS posits the necessity of costly punishment to maintain mutualism. A coevolutionary model of mutualism finds that HS are unlikely to evolve de novo, and published data on legume–rhizobia and yucca–moth mutualisms are consistent with PFF and not with HS. Thus, in systems considered to be textbook cases of HS, we find poor support for the theory that hosts have evolved to punish cheating symbionts; instead, we show that even horizontally transmitted mutualisms can be stabilized via PFF. PFF theory may place previously underappreciated constraints on the evolution of mutualism and explain why punishment is far from ubiquitous in nature.

%B Proceedings of the National Academy of Sciences %V 107 %P 15712-15716 %G eng %N 36 %0 Journal Article %J Biological Journal of the Linnean Society %D 2010 %T How common are dot-like distributions? Taxonomical oversplitting in western European Agrodiaetus (Lepidoptera: Lycaenidae) revealed by chromosomal and molecular markers %A Vila, R. %A Lukhtanov, V. A. %A Talavera, G. %A Gil, F. %A Pierce, N. E. %K blue butterflies %K Conservation %K distribution %K Ecology %K Evolution %K gene-sequences %K maximum-likelihood %K mitochondrial-DNA %K Phylogeny %K phylogeny estimation %K polyommatus %K radiation %K range %K reinforcement %K species %K Systematics %X

Approximately 50 taxa of butterflies in Western Europe have been described as new species or elevated to the level of species during the last 40 years. Many, especially those belonging to the genus Agrodiaetus, have unusually localized, 'dot-like' distributional ranges. In the present study, we use a combination of chromosomal and molecular markers to re-evaluate the species status of Agrodiaetus distributed west of the 17th meridian. The results obtained do not support the current designations of Agrodiaetus galloi, Agrodiaetus exuberans, and Agrodiaetus agenjoi as endemic species with highly restricted distribution ranges, but indicate that these taxa are more likely to be local populations of a widely distributed species, Agrodiaetus ripartii. Agrodiaetus violetae is shown to be a polytypic species consisting of at least two subspecies, including Agrodiaetus violetae subbaeticus comb. nov. and Agrodiaetus violetae violetae. Agrodiaetus violetae is genetically (but not chromosomally) distinct from Agrodiaetus fabressei and has a wider distribution in southern Spain than previously believed. Agrodiaetus humedasae from northern Italy is supported as a highly localized species that is distinct from its nearest relatives. We propose a revision of the species lists for Agrodiaetus taking these new data into account. The results reported in the present study are relevant to animal conservation efforts in Europe because of their implications for IUCN Red List priorities. (C) 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 130-154.

%B Biological Journal of the Linnean Society %V 101 %P 130-154 %8 Sep %@ 0024-4066 %G English %M WOS:000281210500012 %! Biol J Linn SocBiol J Linn Soc %0 Journal Article %J Biodiversity and Conservation %D 2010 %T Local people value environmental services provided by forested parks %A Sodhi, N. S. %A Lee, T. M. %A Sekercioglu, C. H. %A Webb, E. L. %A Prawiradilaga, D. M. %A Lohman, D. J. %A Pierce, N. E. %A Diesmos, A. C. %A Rao, M. %A Ehrlich, P. R. %K Biodiversity %K Conservation %K conservation education %K deforestation %K Ecosystem %K ecosystem services %K livelihoods %K protected areas %K rationale %K southeast asia %K tropical forest %X

Garnering support from local people is critical for maintaining ecologically viable and functional protected areas. However, empirical data illustrating local people's awareness of the importance of nature's services is limited; hence possibly impeding effective ecosystem (environmental)-services based conservation efforts. Using data from five protected forests in four developing Southeast Asian countries, we provide evidence that local people living near parks value a wide range of environmental services, including cultural, provisioning, and regulating services, provided by the forests. Local people with longer residency valued environmental services more. Educated as well as poor people valued forest ecosystem services more. Conservation education has some influence on people's environmental awareness. For conservation endeavors to be successful, large-scale transmigration programs should be avoided and local people must be provided with alternative sustenance opportunities and basic education in addition to environmental outreach to reduce their reliance on protected forests and to enhance conservation support.

%B Biodiversity and Conservation %V 19 %P 1175-1188 %8 Apr %@ 0960-3115 %G English %M WOS:000275536700017 %! Biodivers ConservBiodivers Conserv %0 Journal Article %J Molecular Phylogenetics and Evolution %D 2010 %T A molecular phylogeny of the stingless bee genus Melipona (Hymenoptera: Apidae) %A Ramirez, S. R. %A Nieh, J. C. %A Quental, T. B. %A Roubik, D. W. %A Imperatriz-Fonseca, V. L. %A Pierce, N. E. %K apinae %K apis %K communication %K corbiculate bees %K dance-language %K distance %K divergence times %K eusociality %K fossil bee %K honey bees %K honey-bees %K melipona %K mitochondrial-DNA sequence %K penalized likelihood %K referential communication %K relaxed molecular clock %K stingless bees %X

Stingless bees (Meliponini) constitute a diverse group of highly eusocial insects that occur throughout tropical regions around the world. The meliponine genus Melipona is restricted to the New World tropics and has over 50 described species. Melipona, like Apis, possesses the remarkable ability to use representational communication to indicate the location of foraging patches. Although Melipona has been the subject of numerous behavioral, ecological, and genetic studies, the evolutionary history of this genus remains largely unexplored. Here, we implement a multigene phylogenetic approach based on nuclear, mitochondrial, and ribosomal loci, coupled with molecular clock methods, to elucidate the phylogenetic relationships and antiquity of subgenera and species of Melipona. Our phylogenetic analysis resolves the relationship among subgenera and tends to agree with morphology-based classification hypotheses. Our molecular clock analysis indicates that the genus Melipona shared a most recent common ancestor at least similar to 14-17 million years (My) ago. These results provide the groundwork for future comparative analyses aimed at understanding the evolution of complex communication mechanisms in eusocial Apidae. (C) 2010 Elsevier Inc. All rights reserved.

%B Molecular Phylogenetics and Evolution %V 56 %P 519-525 %8 Aug %@ 1055-7903 %G English %M WOS:000278890100002 %! Mol Phylogenet EvolMol Phylogenet Evol %0 Journal Article %J Biological Journal of the Linnean Society %D 2010 %T Phylogeny, diversification patterns and historical biogeography of euglossine orchid bees (Hymenoptera: Apidae) %A Ramirez, S. R. %A Roubik, D. W. %A Skov, C. %A Pierce, N. E. %K aglae %K amazon %K andes %K biology %K divergence times %K dominican amber %K dtt plots %K eulaema %K Evolution %K forest %K fragrances %K Miocene %K Morphology %K oligocene %K penalized likelihood %K relaxed molecular clocks %K species hymenoptera %K taxonomic notes %X

The orchid bees constitute a clade of prominent insect pollinators distributed throughout the Neotropical region. Males of all species collect fragrances from natural sources, including flowers, decaying vegetation and fungi, and store them in specialized leg pockets to later expose during courtship display. In addition, orchid bees provide pollination services to a diverse array of Neotropical angiosperms when foraging for food and nesting materials. However, despite their ecological importance, little is known about the evolutionary history of orchid bees. Here, we present a comprehensive molecular phylogenetic analysis based on similar to 4.0 kb of DNA from four loci [cytochrome oxidase (CO1), elongation factor 1-alpha (EF1-alpha), arginine kinase (ArgK) and RNA polymerase II (Pol-II)] across the entire tribe Euglossini, including all five genera, eight subgenera and 126 of the approximately 200 known species. We investigated lineage diversification using fossil-calibrated molecular clocks and the evolution of morphological traits using disparity-through-time plots. In addition, we inferred past biogeographical events by implementing model-based likelihood methods. Our dataset supports a new view on generic relationships and indicates that the cleptoparasitic genus Exaerete is sister to the remaining orchid bee genera. Our divergence time estimates indicate that extant orchid bee lineages shared a most recent common ancestor at 27-42 Mya. In addition, our analysis of morphology shows that tongue length and body size experienced rapid disparity bursts that coincide with the origin of diverse genera (Euglossa and Eufriesea). Finally, our analysis of historical biogeography indicates that early diversification episodes shared a history on both sides of Mesoamerica, where orchid bees dispersed across the Caribbean, and through a Panamanian connection, thus reinforcing the hypothesis that recent geological events (e.g. the formation of the isthmus of Panama) contributed to the diversification of the rich Neotropical biota. (C) 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100, 552-572.

%B Biological Journal of the Linnean Society %V 100 %P 552-572 %8 Jul %@ 0024-4066 %G English %M WOS:000279401800007 %! Biol J Linn SocBiol J Linn Soc %0 Journal Article %J Proceedings of the National Academy of Sciences %D 2010 %T Economic contract theory tests models of mutualism. %A Weyl, EG %A ME Frederickson %A Yu, D. W. %A Pierce, N. E. %X

Although mutualisms are common in all ecological communities and have played key roles in the diversification of life, our current understanding of the evolution of cooperation applies mostly to social behavior within a species. A central question is whether mutualisms persist because hosts have evolved costly punishment of cheaters. Here, we use the economic theory of employment contracts to formulate and distinguish between two mechanisms that have been proposed to prevent cheating in host–symbiont mutualisms, partner fidelity feedback (PFF) and host sanctions (HS). Under PFF, positive feedback between host fitness and symbiont fitness is sufficient to prevent cheating; in contrast, HS posits the necessity of costly punishment to maintain mutualism. A coevolutionary model of mutualism finds that HS are unlikely to evolve de novo, and published data on legume–rhizobia and yucca–moth mutualisms are consistent with PFF and not with HS. Thus, in systems considered to be textbook cases of HS, we find poor support for the theory that hosts have evolved to punish cheating symbionts; instead, we show that even horizontally transmitted mutualisms can be stabilized via PFF. PFF theory may place previously underappreciated constraints on the evolution of mutualism and explain why punishment is far from ubiquitous in nature.

%B Proceedings of the National Academy of Sciences %V 107 %P 15712-15716 %G eng %N 36 %0 Journal Article %J Biological Journal of the Linnean Society %D 2010 %T How common are dot-like distributions? Taxonomical oversplitting in western European Agrodiaetus (Lepidoptera: Lycaenidae) revealed by chromosomal and molecular markers %A Vila, R. %A Lukhtanov, V. A. %A Talavera, G. %A Gil, F. %A Pierce, N. E. %K blue butterflies %K Conservation %K distribution %K Ecology %K Evolution %K gene-sequences %K maximum-likelihood %K mitochondrial-DNA %K Phylogeny %K phylogeny estimation %K polyommatus %K radiation %K range %K reinforcement %K species %K Systematics %X

Approximately 50 taxa of butterflies in Western Europe have been described as new species or elevated to the level of species during the last 40 years. Many, especially those belonging to the genus Agrodiaetus, have unusually localized, 'dot-like' distributional ranges. In the present study, we use a combination of chromosomal and molecular markers to re-evaluate the species status of Agrodiaetus distributed west of the 17th meridian. The results obtained do not support the current designations of Agrodiaetus galloi, Agrodiaetus exuberans, and Agrodiaetus agenjoi as endemic species with highly restricted distribution ranges, but indicate that these taxa are more likely to be local populations of a widely distributed species, Agrodiaetus ripartii. Agrodiaetus violetae is shown to be a polytypic species consisting of at least two subspecies, including Agrodiaetus violetae subbaeticus comb. nov. and Agrodiaetus violetae violetae. Agrodiaetus violetae is genetically (but not chromosomally) distinct from Agrodiaetus fabressei and has a wider distribution in southern Spain than previously believed. Agrodiaetus humedasae from northern Italy is supported as a highly localized species that is distinct from its nearest relatives. We propose a revision of the species lists for Agrodiaetus taking these new data into account. The results reported in the present study are relevant to animal conservation efforts in Europe because of their implications for IUCN Red List priorities. (C) 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 130-154.

%B Biological Journal of the Linnean Society %V 101 %P 130-154 %8 Sep %@ 0024-4066 %G English %M WOS:000281210500012 %! Biol J Linn SocBiol J Linn Soc %0 Journal Article %J Biodiversity and Conservation %D 2010 %T Local people value environmental services provided by forested parks %A Sodhi, N. S. %A Lee, T. M. %A Sekercioglu, C. H. %A Webb, E. L. %A Prawiradilaga, D. M. %A Lohman, D. J. %A Pierce, N. E. %A Diesmos, A. C. %A Rao, M. %A Ehrlich, P. R. %K Biodiversity %K Conservation %K conservation education %K deforestation %K Ecosystem %K ecosystem services %K livelihoods %K protected areas %K rationale %K southeast asia %K tropical forest %X

Garnering support from local people is critical for maintaining ecologically viable and functional protected areas. However, empirical data illustrating local people's awareness of the importance of nature's services is limited; hence possibly impeding effective ecosystem (environmental)-services based conservation efforts. Using data from five protected forests in four developing Southeast Asian countries, we provide evidence that local people living near parks value a wide range of environmental services, including cultural, provisioning, and regulating services, provided by the forests. Local people with longer residency valued environmental services more. Educated as well as poor people valued forest ecosystem services more. Conservation education has some influence on people's environmental awareness. For conservation endeavors to be successful, large-scale transmigration programs should be avoided and local people must be provided with alternative sustenance opportunities and basic education in addition to environmental outreach to reduce their reliance on protected forests and to enhance conservation support.

%B Biodiversity and Conservation %V 19 %P 1175-1188 %8 Apr %@ 0960-3115 %G English %M WOS:000275536700017 %! Biodivers ConservBiodivers Conserv %0 Journal Article %J Molecular Phylogenetics and Evolution %D 2010 %T A molecular phylogeny of the stingless bee genus Melipona (Hymenoptera: Apidae) %A Ramirez, S. R. %A Nieh, J. C. %A Quental, T. B. %A Roubik, D. W. %A Imperatriz-Fonseca, V. L. %A Pierce, N. E. %K apinae %K apis %K communication %K corbiculate bees %K dance-language %K distance %K divergence times %K eusociality %K fossil bee %K honey bees %K honey-bees %K melipona %K mitochondrial-DNA sequence %K penalized likelihood %K referential communication %K relaxed molecular clock %K stingless bees %X

Stingless bees (Meliponini) constitute a diverse group of highly eusocial insects that occur throughout tropical regions around the world. The meliponine genus Melipona is restricted to the New World tropics and has over 50 described species. Melipona, like Apis, possesses the remarkable ability to use representational communication to indicate the location of foraging patches. Although Melipona has been the subject of numerous behavioral, ecological, and genetic studies, the evolutionary history of this genus remains largely unexplored. Here, we implement a multigene phylogenetic approach based on nuclear, mitochondrial, and ribosomal loci, coupled with molecular clock methods, to elucidate the phylogenetic relationships and antiquity of subgenera and species of Melipona. Our phylogenetic analysis resolves the relationship among subgenera and tends to agree with morphology-based classification hypotheses. Our molecular clock analysis indicates that the genus Melipona shared a most recent common ancestor at least similar to 14-17 million years (My) ago. These results provide the groundwork for future comparative analyses aimed at understanding the evolution of complex communication mechanisms in eusocial Apidae. (C) 2010 Elsevier Inc. All rights reserved.

%B Molecular Phylogenetics and Evolution %V 56 %P 519-525 %8 Aug %@ 1055-7903 %G English %M WOS:000278890100002 %! Mol Phylogenet EvolMol Phylogenet Evol %0 Journal Article %J Biological Journal of the Linnean Society %D 2010 %T Phylogeny, diversification patterns and historical biogeography of euglossine orchid bees (Hymenoptera: Apidae) %A Ramirez, S. R. %A Roubik, D. W. %A Skov, C. %A Pierce, N. E. %K aglae %K amazon %K andes %K biology %K divergence times %K dominican amber %K dtt plots %K eulaema %K Evolution %K forest %K fragrances %K Miocene %K Morphology %K oligocene %K penalized likelihood %K relaxed molecular clocks %K species hymenoptera %K taxonomic notes %X

The orchid bees constitute a clade of prominent insect pollinators distributed throughout the Neotropical region. Males of all species collect fragrances from natural sources, including flowers, decaying vegetation and fungi, and store them in specialized leg pockets to later expose during courtship display. In addition, orchid bees provide pollination services to a diverse array of Neotropical angiosperms when foraging for food and nesting materials. However, despite their ecological importance, little is known about the evolutionary history of orchid bees. Here, we present a comprehensive molecular phylogenetic analysis based on similar to 4.0 kb of DNA from four loci [cytochrome oxidase (CO1), elongation factor 1-alpha (EF1-alpha), arginine kinase (ArgK) and RNA polymerase II (Pol-II)] across the entire tribe Euglossini, including all five genera, eight subgenera and 126 of the approximately 200 known species. We investigated lineage diversification using fossil-calibrated molecular clocks and the evolution of morphological traits using disparity-through-time plots. In addition, we inferred past biogeographical events by implementing model-based likelihood methods. Our dataset supports a new view on generic relationships and indicates that the cleptoparasitic genus Exaerete is sister to the remaining orchid bee genera. Our divergence time estimates indicate that extant orchid bee lineages shared a most recent common ancestor at 27-42 Mya. In addition, our analysis of morphology shows that tongue length and body size experienced rapid disparity bursts that coincide with the origin of diverse genera (Euglossa and Eufriesea). Finally, our analysis of historical biogeography indicates that early diversification episodes shared a history on both sides of Mesoamerica, where orchid bees dispersed across the Caribbean, and through a Panamanian connection, thus reinforcing the hypothesis that recent geological events (e.g. the formation of the isthmus of Panama) contributed to the diversification of the rich Neotropical biota. (C) 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100, 552-572.

%B Biological Journal of the Linnean Society %V 100 %P 552-572 %8 Jul %@ 0024-4066 %G English %M WOS:000279401800007 %! Biol J Linn SocBiol J Linn Soc %0 Journal Article %J Proceedings of the National Academy of Sciences of the United States of America %D 2009 %T Bacterial gut symbionts are tightly linked with the evolution of herbivory in ants %A Russell, J. A. %A Moreau, C. S. %A Goldman-Huertas, B. %A Fujiwara, M. %A Lohman, D. J. %A Pierce, N. E. %K blochmannia %K community %K diversification %K diversity %K formicidae %K microbiota %K microorganisms %K nitrogen-fixation %K rain-forest %K rhizobiales %K symbiosis %K tetraponera %K trophic level %K wolbachia %X

Ants are a dominant feature of terrestrial ecosystems, yet we know little about the forces that drive their evolution. Recent findings illustrate that their diets range from herbivorous to predaceous, with "herbivores'' feeding primarily on exudates from plants and sap-feeding insects. Persistence on these nitrogen-poor food sources raises the question of how ants obtain sufficient nutrition. To investigate the potential role of symbiotic microbes, we have surveyed 283 species from 18 of the 21 ant subfamilies using molecular techniques. Our findings uncovered a wealth of bacteria from across the ants. Notable among the surveyed hosts were herbivorous "turtle ants'' from the related genera Cephalotes and Procryptocerus (tribe Cephalotini). These commonly harbored bacteria from ant-specific clades within the Burkholderiales, Pseudomonadales, Rhizobiales, Verrucomicrobiales, and Xanthomonadales, and studies of lab-reared Cephalotes varians characterized these microbes as symbiotic residents of ant guts. Although most of these symbionts were confined to turtle ants, bacteria from an ant-specific clade of Rhizobiales were more broadly distributed. Statistical analyses revealed a strong relationship between herbivory and the prevalence of Rhizobiales gut symbionts within ant genera. Furthermore, a consideration of the ant phylogeny identified at least five independent origins of symbioses between herbivorous ants and related Rhizobiales. Combined with previous findings and the potential for symbiotic nitrogen fixation, our results strongly support the hypothesis that bacteria have facilitated convergent evolution of herbivory across the ants, further implicating symbiosis as a major force in ant evolution.

%B Proceedings of the National Academy of Sciences of the United States of America %V 106 %P 21236-21241 %8 Dec 15 %@ 0027-8424 %G English %M WOS:000272795300042 %! P Natl Acad Sci USAP Natl Acad Sci USA %0 Journal Article %J Ecological Entomology %D 2009 %T The double cloak of invisibility: phenotypic plasticity and larval decoration in a geometrid moth, Synchlora frondaria, across three diet treatments %A Canfield, M. R. %A Chang, S. %A Pierce, N. E. %K camouflage %K crypsis %K decoration %K Evolution %K growth %K lepidoptera %K Phenotypic plasticity %K phytophagous insects %K polyphenism %X

1. Crypsis is one of the main defences that insects use to avoid predators, and both the juveniles and adults of many geometrid moths are remarkable in their ability to blend into different host backgrounds. The larvae of Synchlora frondaria have two methods to achieve crypsis: phenotypic plasticity in colouration that enable them to hide more effectively on their host plants, and a self-decorating behaviour whereby the larvae camouflage themselves with materials from their host plants.2. Larvae of Synchlora frondaria reared on three different host plants showed systematic differences in relative growth rate, survivorship and larval colouration.3. Larval colouration varied across diet treatments in a way that was consistent with diet-induced phenotypic plasticity, and larvae also exhibited characteristic decorating behaviour on all three hosts.4. Larvae showed highest survivorship on Heterotheca subaxillaris (Asteraceae), and had significantly higher relative growth rates on H. subaxillaris (Asteraceae) and Lantana camara (Verbenaceae) than on Bejaria racemosa (Ericaceae).5. Synchlora frondaria provides an example of a species where both decorating behaviour and phenotypic plasticity in larval colouration produce a cryptic form that is remarkably responsive to its background.

%B Ecological Entomology %V 34 %P 412-414 %8 Jun %@ 0307-6946 %G English %M WOS:000266015900015 %! Ecol EntomolEcol Entomol %0 Journal Article %J Evolution %D 2009 %T Specialization and Geographic Isolation among Wolbachia Symbionts from Ants and Lycaenid Butterflies %A Russell, J. A. %A Goldman-Huertas, B. %A Moreau, C. S. %A Baldo, L. %A Stahlhut, J. K. %A Werren, J. H. %A Pierce, N. E. %K Bacteria %K coevolution %K drosophila-simulans diptera %K ephestia-kuehniella %K evolutionary history %K gene-sequences %K horizontal transfer %K Insects %K mediterranean flour moth %K mitochondrial-DNA %K muscidifurax-uniraptor %K phylogenetics %K specialization %K symbiont %K trichogramma-wasps %K wsp gene %X

Wolbachia are the most prevalent and influential bacteria described among the insects to date. But despite their significance, we lack an understanding of their evolutionary histories. To describe the evolution of symbioses between Wolbachia and their hosts, we surveyed global collections of two diverse families of insects, the ants and lycaenid butterflies. In total, 54 Wolbachia isolates were typed using a Multi Locus Sequence Typing (MLST) approach, in which five unlinked loci were sequenced and analyzed to decipher evolutionary patterns. AMOVA and phylogenetic analyses demonstrated that related Wolbachia commonly infect related hosts, revealing a pattern of host association that was strongest among strains from the ants. A review of the literature indicated that horizontal transfer is most successful when Wolbachia move between related hosts, suggesting that patterns of host association are driven by specialization on a common physiological background. Aside from providing the broadest and strongest evidence to date for Wolbachia specialization, our findings also reveal that strains from New World ants differ markedly from those in ants from other locations. We, therefore, conclude that both geographic and phylogenetic barriers have promoted evolutionary divergence among these influential symbionts.

%B Evolution %V 63 %P 624-640 %8 Mar %@ 0014-3820 %G English %M WOS:000263679800005 %! EvolutionEvolution %0 Journal Article %J Proceedings of the National Academy of Sciences of the United States of America %D 2009 %T Bacterial gut symbionts are tightly linked with the evolution of herbivory in ants %A Russell, J. A. %A Moreau, C. S. %A Goldman-Huertas, B. %A Fujiwara, M. %A Lohman, D. J. %A Pierce, N. E. %K blochmannia %K community %K diversification %K diversity %K formicidae %K microbiota %K microorganisms %K nitrogen-fixation %K rain-forest %K rhizobiales %K symbiosis %K tetraponera %K trophic level %K wolbachia %X

Ants are a dominant feature of terrestrial ecosystems, yet we know little about the forces that drive their evolution. Recent findings illustrate that their diets range from herbivorous to predaceous, with "herbivores'' feeding primarily on exudates from plants and sap-feeding insects. Persistence on these nitrogen-poor food sources raises the question of how ants obtain sufficient nutrition. To investigate the potential role of symbiotic microbes, we have surveyed 283 species from 18 of the 21 ant subfamilies using molecular techniques. Our findings uncovered a wealth of bacteria from across the ants. Notable among the surveyed hosts were herbivorous "turtle ants'' from the related genera Cephalotes and Procryptocerus (tribe Cephalotini). These commonly harbored bacteria from ant-specific clades within the Burkholderiales, Pseudomonadales, Rhizobiales, Verrucomicrobiales, and Xanthomonadales, and studies of lab-reared Cephalotes varians characterized these microbes as symbiotic residents of ant guts. Although most of these symbionts were confined to turtle ants, bacteria from an ant-specific clade of Rhizobiales were more broadly distributed. Statistical analyses revealed a strong relationship between herbivory and the prevalence of Rhizobiales gut symbionts within ant genera. Furthermore, a consideration of the ant phylogeny identified at least five independent origins of symbioses between herbivorous ants and related Rhizobiales. Combined with previous findings and the potential for symbiotic nitrogen fixation, our results strongly support the hypothesis that bacteria have facilitated convergent evolution of herbivory across the ants, further implicating symbiosis as a major force in ant evolution.

%B Proceedings of the National Academy of Sciences of the United States of America %V 106 %P 21236-21241 %8 Dec 15 %@ 0027-8424 %G English %M WOS:000272795300042 %! P Natl Acad Sci USAP Natl Acad Sci USA %0 Journal Article %J Ecological Entomology %D 2009 %T The double cloak of invisibility: phenotypic plasticity and larval decoration in a geometrid moth, Synchlora frondaria, across three diet treatments %A Canfield, M. R. %A Chang, S. %A Pierce, N. E. %K camouflage %K crypsis %K decoration %K Evolution %K growth %K lepidoptera %K Phenotypic plasticity %K phytophagous insects %K polyphenism %X

1. Crypsis is one of the main defences that insects use to avoid predators, and both the juveniles and adults of many geometrid moths are remarkable in their ability to blend into different host backgrounds. The larvae of Synchlora frondaria have two methods to achieve crypsis: phenotypic plasticity in colouration that enable them to hide more effectively on their host plants, and a self-decorating behaviour whereby the larvae camouflage themselves with materials from their host plants.2. Larvae of Synchlora frondaria reared on three different host plants showed systematic differences in relative growth rate, survivorship and larval colouration.3. Larval colouration varied across diet treatments in a way that was consistent with diet-induced phenotypic plasticity, and larvae also exhibited characteristic decorating behaviour on all three hosts.4. Larvae showed highest survivorship on Heterotheca subaxillaris (Asteraceae), and had significantly higher relative growth rates on H. subaxillaris (Asteraceae) and Lantana camara (Verbenaceae) than on Bejaria racemosa (Ericaceae).5. Synchlora frondaria provides an example of a species where both decorating behaviour and phenotypic plasticity in larval colouration produce a cryptic form that is remarkably responsive to its background.

%B Ecological Entomology %V 34 %P 412-414 %8 Jun %@ 0307-6946 %G English %M WOS:000266015900015 %! Ecol EntomolEcol Entomol %0 Journal Article %J Evolution %D 2009 %T Specialization and Geographic Isolation among Wolbachia Symbionts from Ants and Lycaenid Butterflies %A Russell, J. A. %A Goldman-Huertas, B. %A Moreau, C. S. %A Baldo, L. %A Stahlhut, J. K. %A Werren, J. H. %A Pierce, N. E. %K Bacteria %K coevolution %K drosophila-simulans diptera %K ephestia-kuehniella %K evolutionary history %K gene-sequences %K horizontal transfer %K Insects %K mediterranean flour moth %K mitochondrial-DNA %K muscidifurax-uniraptor %K phylogenetics %K specialization %K symbiont %K trichogramma-wasps %K wsp gene %X

Wolbachia are the most prevalent and influential bacteria described among the insects to date. But despite their significance, we lack an understanding of their evolutionary histories. To describe the evolution of symbioses between Wolbachia and their hosts, we surveyed global collections of two diverse families of insects, the ants and lycaenid butterflies. In total, 54 Wolbachia isolates were typed using a Multi Locus Sequence Typing (MLST) approach, in which five unlinked loci were sequenced and analyzed to decipher evolutionary patterns. AMOVA and phylogenetic analyses demonstrated that related Wolbachia commonly infect related hosts, revealing a pattern of host association that was strongest among strains from the ants. A review of the literature indicated that horizontal transfer is most successful when Wolbachia move between related hosts, suggesting that patterns of host association are driven by specialization on a common physiological background. Aside from providing the broadest and strongest evidence to date for Wolbachia specialization, our findings also reveal that strains from New World ants differ markedly from those in ants from other locations. We, therefore, conclude that both geographic and phylogenetic barriers have promoted evolutionary divergence among these influential symbionts.

%B Evolution %V 63 %P 624-640 %8 Mar %@ 0014-3820 %G English %M WOS:000263679800005 %! EvolutionEvolution %0 Journal Article %J Trends in Ecology & Evolution %D 2008 %T Delicious poison: genetics of Drosophila host plant preference %A Whiteman, N. K. %A Pierce, N. E. %K Behavior %K Evolution %K expression %K family %K genomes %K Humans %K resistance %K sechellia %K selection %K specialization %X

Insects use chemical cues to identify host plants, which suggests that chemosensory perception could be a target of natural selection during host specialization. Five papers using data from the 12 recently sequenced Drosophila genomes examined chemosensory gene function and evolution across specialist and generalist species. A functional study identifies odorant binding proteins that mediate loss of toxin avoidance in a specialist, and targeted genomic studies indicate specialists and island endemics lose chemosensory genes more rapidly than generalist and mainland relatives. Together, these studies suggest a mode of chemoreceptor evolution dominated by birth/death dynamics, coupled with a low level of potential positive selection.

%B Trends in Ecology & Evolution %V 23 %P 473-478 %8 Sep %@ 0169-5347 %G English %M WOS:000259283600002 %! Trends Ecol EvolTrends Ecol Evol %0 Journal Article %J Molecular Phylogenetics and Evolution %D 2008 %T Exploring phenotypic plasticity and biogeography in emerald moths: A phylogeny of the genus Nemoria (Lepidoptera: Geometridae) %A Canfield, MR %A E Greene %A CS Moreau %A N Chen %A Pierce, N. E. %X

The moth genus Nemoria (Lepidoptera: Geometridae) includes 134 described species whose larvae and adults display a considerable range of phenotypic plasticity in coloration and morphology. We reconstructed the phylogeny of 54 species of Nemoria and seven outgroups using characters from the mitochondrial genes, Cytochrome Oxidase I and II (COI and COII), and the nuclear gene, Elongation Factor-a (EF-1a). Maximum parsimony, maximum likelihood and Bayesian inference were used to infer the phylogeny. The 54 ingroup species represented 13 of the 15 recognized species groups of Nemoria [Ferguson, D.C., 1985. Fasc. 18.1, Geometroidea: Geometridae (in part). In: Dominick, R.B. (Ed.), The Moths of America North of Mexico, Fasc. 18.1. Wedge Entomological Research Foundation, Washington; Pitkin, L.M., 1993. Neotropical emerald moths of the genera Nemoria, Lissochlora and Chavarriella, with particular reference to the species of Costa Rica (Lepidoptera: Geometridae, Geometrinae). Bull. Br. Mus. Nat. Hist. 62, 39–159], and the seven outgroups came from four tribes of Geometrinae. These data support Nemoria as a monophyletic group and largely recover the species groupings proposed in previous taxonomic analyses using morphological characters. Phenotypic plasticity of larvae is not correlated with plasticity of adults among those species of Nemoria where life histories are known, and appears to be evolutionarily labile for both life history stages: Species exhibiting larval phenotypic plasticity, such as N. arizonaria and N. outina, are placed in several distinct clades, suggesting that this trait has evolved multiple times, and species displaying adult phenotypic plasticity are likewise distributed throughout the phylogeny. A comparative analysis of the biogeographic history of Nemoria supports a South American origin for the genus with multiple introductions into North America, and an application of published substitution rates to the phylogram provides an age estimate of 7.5 million years.

%B Molecular Phylogenetics and Evolution %V 49 %P 477-487 %G eng %0 Journal Article %J Tropical Lepidoptera Research %D 2008 %T A novel organ and mechanism for larval sound production in butterfly caterpillars: Eurybia elvina(Lepidoptera: Riodinidae). %A MA Travassos %A PJ Devries %A Pierce, N. E. %X

Abstract – Eurybia elvina larvae produce substrate-borne vibrations by grating a cervical membrane studded with teeth against hemispherical protuberances scattered along the surface of the head.

%B Tropical Lepidoptera Research %V 18 %P 20-23 %G eng %0 Journal Article %J Bmc Evolutionary Biology %D 2008 %T Phylogeography and genetic diversity of a widespread Old World butterfly, Lampides boeticus (Lepidoptera: Lycaenidae) %A Lohman, D. J. %A Peggie, D. %A Pierce, N. E. %A Meier, R. %K evolutionary %K helicoverpa-armigera lepidoptera %K identification %K maximum-likelihood %K mitochondrial-DNA %K parameters %K polymorphism %K population-growth %K sequences %K wolbachia %X

Background: Evolutionary genetics provides a rich theoretical framework for empirical studies of phylogeography. Investigations of intraspecific genetic variation can uncover new putative species while allowing inference into the evolutionary origin and history of extant populations. With a distribution on four continents ranging throughout most of the Old World, Lampides boeticus ( Lepidoptera: Lycaenidae) is one of the most widely distributed species of butterfly. It is placed in a monotypic genus with no commonly accepted subspecies. Here, we investigate the demographic history and taxonomic status of this widespread species, and screen for the presence or absence of the bacterial endosymbiont Wolbachia.Results: We performed phylogenetic, population genetic, and phylogeographic analyses using 1799 bp of mitochondrial sequence data from 57 specimens collected throughout the species' range. Most of the samples(> 90%) were nearly genetically identical, with uncorrected pairwise sequence differences of 0 0.5% across geographic distances > 9,000 km. However, five samples from central Thailand, Madagascar, northern Australia and the Moluccas formed two divergent clades differing from the majority of samples by uncorrected pairwise distances ranging from 1.79-2.21%. Phylogenetic analyses suggest that L. boeticus is almost certainly monophyletic, with all sampled genes coalescing well after the divergence from three closely related taxa included for outgroup comparisons. Analyses of molecular diversity indicate that most L. boeticus individuals in extant populations are descended from one or two relatively recent population bottlenecks.Conclusion: The combined analyses suggest a scenario in which the most recent common ancestor of L. boeticus and its sister taxon lived in the African region approximately 7 Mya; extant lineages of L. boeticus began spreading throughout the Old World at least 1.5 Mya. More recently, expansion after population bottlenecks approximately 1.4 Mya seem to have displaced most of the ancestral polymorphism throughout its range, though at least two early-branching lineages still persist. One of these lineages, in northern Australia and the Moluccas, may have experienced accelerated differentiation due to infection with the bacterial endosymbiont Wolbachia, which affects reproduction. Examination of a haplotype network suggests that Australia has been colonized by the species several times. While there is little evidence for the existence of morphologically cryptic species, these results suggest a complex history affected by repeated dispersal events.

%B Bmc Evolutionary Biology %V 8 %8 Oct 30 %@ 1471-2148 %G English %M WOS:000262180200001 %! Bmc Evol BiolBmc Evol Biol %0 Journal Article %J Tropical Lepidoptera Research %D 2008 %T The singing reaper: diet, morphology and vibrational signaling in the Nearctic species Feniseca tarquinius. %A Mathew, J %A MA Travassos %A M Canfield %A D Murawski %A RL Kitching %A Pierce, N. E. %X

Abstract – A survey at fourteen sites in Eastern North America of populations of the carnivorous lycaenid butterfly, Feniseca tarquinius, confirmed that the sole prey item on Alnus rugosa (Betulaceae) for this species in these regions was Paraprociphilus tessellatus (Homoptera: Aphidoidea: Pemphigidae). Overwhelmingly, these aphids were tended by ants in the subfamily Formicinae. These results are compiled with all earlier records of prey aphids, their host plants and attendant ants for this species. SEM examination of a 4th instar larva of F. tarquinius supported Cottrell’s (1984) observation that the dorsal nectary organ and tentacle organs are absent in the 4th instar of virtually all Miletinae. Larvae of F. tarquinius were found to produce substrate-borne vibrations that possess a long pulse length and narrow bandwidth when compared with other lycaenid calls. The possible function of these calls is briefly discussed.

%B Tropical Lepidoptera Research %V 18 %P 24-29 %G eng %0 Journal Article %J Trends in Ecology & Evolution %D 2008 %T Social insect symbionts: evolution in homeostatic fortresses %A Hughes, D. P. %A Pierce, N. E. %A Boomsma, J. J. %K apis-mellifera %K cooperation %K Ecology %K fungus-growing ants %K latitudinal diversity gradient %K life-history %K mutualism %K Parasites %K pathogens %K Virulence %X

The massive environmentally buffered nests of some social insects can contain millions of individuals and a wide variety of parasites, commensals and mutualists. We suggest that the ways in which these homeostatic fortress environments affect the evolution of social insect symbionts are relevant for epidemiology, evolutionary biology and macroecology. We contend that specialized parasites will tend to become less virulent and mutualists less cooperative, compared to those associated with solitary or small-colony hosts. These processes are expected to contribute to the very high symbiont diversity observed in these nests. We hypothesize that biodiversity gradients in these hotspots might be less affected by abiotic latitudinal clines than gradients in neighboring 'control' habitats. We suggest several research lines to test these ideas.

%B Trends in Ecology & Evolution %V 23 %P 672-677 %8 Dec %@ 0169-5347 %G English %M WOS:000261532300005 %! Trends Ecol EvolTrends Ecol Evol %0 Journal Article %J Molecular Phylogenetics and Evolution %D 2008 %T Exploring phenotypic plasticity and biogeography in emerald moths: A phylogeny of the genus Nemoria (Lepidoptera: Geometridae) %A Canfield, MR %A E Greene %A CS Moreau %A N Chen %A Pierce, N. E. %X

The moth genus Nemoria (Lepidoptera: Geometridae) includes 134 described species whose larvae and adults display a considerable range of phenotypic plasticity in coloration and morphology. We reconstructed the phylogeny of 54 species of Nemoria and seven outgroups using characters from the mitochondrial genes, Cytochrome Oxidase I and II (COI and COII), and the nuclear gene, Elongation Factor-a (EF-1a). Maximum parsimony, maximum likelihood and Bayesian inference were used to infer the phylogeny. The 54 ingroup species represented 13 of the 15 recognized species groups of Nemoria [Ferguson, D.C., 1985. Fasc. 18.1, Geometroidea: Geometridae (in part). In: Dominick, R.B. (Ed.), The Moths of America North of Mexico, Fasc. 18.1. Wedge Entomological Research Foundation, Washington; Pitkin, L.M., 1993. Neotropical emerald moths of the genera Nemoria, Lissochlora and Chavarriella, with particular reference to the species of Costa Rica (Lepidoptera: Geometridae, Geometrinae). Bull. Br. Mus. Nat. Hist. 62, 39–159], and the seven outgroups came from four tribes of Geometrinae. These data support Nemoria as a monophyletic group and largely recover the species groupings proposed in previous taxonomic analyses using morphological characters. Phenotypic plasticity of larvae is not correlated with plasticity of adults among those species of Nemoria where life histories are known, and appears to be evolutionarily labile for both life history stages: Species exhibiting larval phenotypic plasticity, such as N. arizonaria and N. outina, are placed in several distinct clades, suggesting that this trait has evolved multiple times, and species displaying adult phenotypic plasticity are likewise distributed throughout the phylogeny. A comparative analysis of the biogeographic history of Nemoria supports a South American origin for the genus with multiple introductions into North America, and an application of published substitution rates to the phylogram provides an age estimate of 7.5 million years.

%B Molecular Phylogenetics and Evolution %V 49 %P 477-487 %G eng %0 Journal Article %J Tropical Lepidoptera Research %D 2008 %T A novel organ and mechanism for larval sound production in butterfly caterpillars: Eurybia elvina(Lepidoptera: Riodinidae). %A MA Travassos %A PJ Devries %A Pierce, N. E. %X

Abstract – Eurybia elvina larvae produce substrate-borne vibrations by grating a cervical membrane studded with teeth against hemispherical protuberances scattered along the surface of the head.

%B Tropical Lepidoptera Research %V 18 %P 20-23 %G eng %0 Journal Article %J Tropical Lepidoptera Research %D 2008 %T The singing reaper: diet, morphology and vibrational signaling in the Nearctic species Feniseca tarquinius. %A Mathew, J %A MA Travassos %A M Canfield %A D Murawski %A RL Kitching %A Pierce, N. E. %X

Abstract – A survey at fourteen sites in Eastern North America of populations of the carnivorous lycaenid butterfly, Feniseca tarquinius, confirmed that the sole prey item on Alnus rugosa (Betulaceae) for this species in these regions was Paraprociphilus tessellatus (Homoptera: Aphidoidea: Pemphigidae). Overwhelmingly, these aphids were tended by ants in the subfamily Formicinae. These results are compiled with all earlier records of prey aphids, their host plants and attendant ants for this species. SEM examination of a 4th instar larva of F. tarquinius supported Cottrell’s (1984) observation that the dorsal nectary organ and tentacle organs are absent in the 4th instar of virtually all Miletinae. Larvae of F. tarquinius were found to produce substrate-borne vibrations that possess a long pulse length and narrow bandwidth when compared with other lycaenid calls. The possible function of these calls is briefly discussed.

%B Tropical Lepidoptera Research %V 18 %P 24-29 %G eng %0 Journal Article %J Trends in Ecology & Evolution %D 2008 %T Delicious poison: genetics of Drosophila host plant preference %A Whiteman, N. K. %A Pierce, N. E. %K Behavior %K Evolution %K expression %K family %K genomes %K Humans %K resistance %K sechellia %K selection %K specialization %X

Insects use chemical cues to identify host plants, which suggests that chemosensory perception could be a target of natural selection during host specialization. Five papers using data from the 12 recently sequenced Drosophila genomes examined chemosensory gene function and evolution across specialist and generalist species. A functional study identifies odorant binding proteins that mediate loss of toxin avoidance in a specialist, and targeted genomic studies indicate specialists and island endemics lose chemosensory genes more rapidly than generalist and mainland relatives. Together, these studies suggest a mode of chemoreceptor evolution dominated by birth/death dynamics, coupled with a low level of potential positive selection.

%B Trends in Ecology & Evolution %V 23 %P 473-478 %8 Sep %@ 0169-5347 %G English %M WOS:000259283600002 %! Trends Ecol EvolTrends Ecol Evol %0 Journal Article %J Bmc Evolutionary Biology %D 2008 %T Phylogeography and genetic diversity of a widespread Old World butterfly, Lampides boeticus (Lepidoptera: Lycaenidae) %A Lohman, D. J. %A Peggie, D. %A Pierce, N. E. %A Meier, R. %K evolutionary %K helicoverpa-armigera lepidoptera %K identification %K maximum-likelihood %K mitochondrial-DNA %K parameters %K polymorphism %K population-growth %K sequences %K wolbachia %X

Background: Evolutionary genetics provides a rich theoretical framework for empirical studies of phylogeography. Investigations of intraspecific genetic variation can uncover new putative species while allowing inference into the evolutionary origin and history of extant populations. With a distribution on four continents ranging throughout most of the Old World, Lampides boeticus ( Lepidoptera: Lycaenidae) is one of the most widely distributed species of butterfly. It is placed in a monotypic genus with no commonly accepted subspecies. Here, we investigate the demographic history and taxonomic status of this widespread species, and screen for the presence or absence of the bacterial endosymbiont Wolbachia.Results: We performed phylogenetic, population genetic, and phylogeographic analyses using 1799 bp of mitochondrial sequence data from 57 specimens collected throughout the species' range. Most of the samples(> 90%) were nearly genetically identical, with uncorrected pairwise sequence differences of 0 0.5% across geographic distances > 9,000 km. However, five samples from central Thailand, Madagascar, northern Australia and the Moluccas formed two divergent clades differing from the majority of samples by uncorrected pairwise distances ranging from 1.79-2.21%. Phylogenetic analyses suggest that L. boeticus is almost certainly monophyletic, with all sampled genes coalescing well after the divergence from three closely related taxa included for outgroup comparisons. Analyses of molecular diversity indicate that most L. boeticus individuals in extant populations are descended from one or two relatively recent population bottlenecks.Conclusion: The combined analyses suggest a scenario in which the most recent common ancestor of L. boeticus and its sister taxon lived in the African region approximately 7 Mya; extant lineages of L. boeticus began spreading throughout the Old World at least 1.5 Mya. More recently, expansion after population bottlenecks approximately 1.4 Mya seem to have displaced most of the ancestral polymorphism throughout its range, though at least two early-branching lineages still persist. One of these lineages, in northern Australia and the Moluccas, may have experienced accelerated differentiation due to infection with the bacterial endosymbiont Wolbachia, which affects reproduction. Examination of a haplotype network suggests that Australia has been colonized by the species several times. While there is little evidence for the existence of morphologically cryptic species, these results suggest a complex history affected by repeated dispersal events.

%B Bmc Evolutionary Biology %V 8 %8 Oct 30 %@ 1471-2148 %G English %M WOS:000262180200001 %! Bmc Evol BiolBmc Evol Biol %0 Journal Article %J Trends in Ecology & Evolution %D 2008 %T Social insect symbionts: evolution in homeostatic fortresses %A Hughes, D. P. %A Pierce, N. E. %A Boomsma, J. J. %K apis-mellifera %K cooperation %K Ecology %K fungus-growing ants %K latitudinal diversity gradient %K life-history %K mutualism %K Parasites %K pathogens %K Virulence %X

The massive environmentally buffered nests of some social insects can contain millions of individuals and a wide variety of parasites, commensals and mutualists. We suggest that the ways in which these homeostatic fortress environments affect the evolution of social insect symbionts are relevant for epidemiology, evolutionary biology and macroecology. We contend that specialized parasites will tend to become less virulent and mutualists less cooperative, compared to those associated with solitary or small-colony hosts. These processes are expected to contribute to the very high symbiont diversity observed in these nests. We hypothesize that biodiversity gradients in these hotspots might be less affected by abiotic latitudinal clines than gradients in neighboring 'control' habitats. We suggest several research lines to test these ideas.

%B Trends in Ecology & Evolution %V 23 %P 672-677 %8 Dec %@ 0169-5347 %G English %M WOS:000261532300005 %! Trends Ecol EvolTrends Ecol Evol %0 Journal Article %J Nature %D 2007 %T Dating the origin of the Orchidaceae from a fossil orchid with its pollinator %A Ramirez, S. R. %A Gravendeel, B. %A Singer, R. B. %A Marshall, C. R. %A Pierce, N. E. %K age %K amber %K angiosperms %K Evolution %K flowering plants %K Morphology %K sequences %X

Since the time of Darwin(1), evolutionary biologists have been fascinated by the spectacular adaptations to insect pollination exhibited by orchids. However, despite being the most diverse plant family on Earth(2), the Orchidaceae lack a definitive fossil record and thus many aspects of their evolutionary history remain obscure. Here we report an exquisitely preserved orchid pollinarium (of Meliorchis caribea gen. et sp. nov.) attached to the mesoscutellum of an extinct stingless bee, Proplebeia dominicana, recovered from Miocene amber in the Dominican Republic, that is 15-20 million years (Myr) old(3). This discovery constitutes both the first unambiguous fossil of Orchidaceae(4) and an unprecedented direct fossil observation of a plant-pollinator interaction(5,6). By applying cladistic methods to a morphological character matrix, we resolve the phylogenetic position of M. caribea within the extant subtribe Goodyerinae (subfamily Orchidoideae). We use the ages of other fossil monocots and M. caribea to calibrate a molecular phylogenetic tree of the Orchidaceae. Our results indicate that the most recent common ancestor of extant orchids lived in the Late Cretaceous (76-84 Myr ago), and also suggest that the dramatic radiation of orchids began shortly after the mass extinctions at the K/T boundary. These results further support the hypothesis of an ancient origin for Orchidaceae.

%B Nature %V 448 %P 1042-1045 %8 Aug 30 %@ 0028-0836 %G English %M WOS:000249097600036 %! NatureNature %0 Journal Article %J Molecular Ecology %D 2007 %T The geography of diversification in mutualistic ants: a gene's-eye view into the Neogene history of Sundaland rain forests %A Quek, S. P. %A Davies, S. J. %A Ashton, P. S. %A Itino, T. %A Pierce, N. E. %K association %K biogeography %K cospeciation %K crematogaster %K cytochrome oxidase %K formicidae %K macaranga %K macaranga euphorbiaceae %K mitochondrial-DNA %K molecular clock %K polymorphism %K population-growth %K rain forest %K southeast asia %K southeast-asia %X

We investigate the geographical and historical context of diversification in a complex of mutualistic Crematogaster ants living in Macaranga trees in the equatorial rain forests of Southeast Asia. Using mitochondrial DNA from 433 ant colonies collected from 32 locations spanning Borneo, Malaya and Sumatra, we infer branching relationships, patterns of genetic diversity and population history. We reconstruct a time frame for the ants' diversification and demographic expansions, and identify areas that might have been refugia or centres of diversification. Seventeen operational lineages are identified, most of which can be distinguished by host preference and geographical range. The ants first diversified 16-20 Ma, not long after the onset of the everwet forests in Sundaland, and achieved most of their taxonomic diversity during the Pliocene. Pleistocene demographic expansions are inferred for several of the younger lineages. Phylogenetic relationships suggest a Bornean cradle and major axis of diversification. Taxonomic diversity tends to be associated with mountain ranges; in Borneo, it is greatest in the Crocker Range of Sabah and concentrated also in other parts of the northern northwest coast. Within-lineage genetic diversity in Malaya and Sumatra tends to also coincide with mountain ranges. A series of disjunct and restricted distributions spanning northern northwest Borneo and the major mountain ranges of Malaya and Sumatra, seen in three pairs of sister lineages, further suggests that these regions were rain-forest refuges during drier climatic phases of the Pleistocene. Results are discussed in the context of the history of Sundaland's rain forests.

%B Molecular Ecology %V 16 %P 2045-2062 %8 May %@ 0962-1083 %G English %M WOS:000246366800006 %! Mol EcolMol Ecol %0 Journal Article %J American Naturalist %D 2007 %T Host plant specialization driven by sexual selection %A Quental, T. B. %A Patten, M. M. %A Pierce, N. E. %K danaid butterflies %K defense-mechanisms %K diet breadth %K enemy-free space %K Evolution %K female choice %K host shift %K lepidoptera %K maternal effects %K moth utetheisa-ornatrix %K pheromone %K phytophagous insects %K pyrrolizidine alkaloids %K specialization %X

We propose a new mechanism based on sexual selection to explain the evolution of diet breadth in insects. More specifically, we show that mate choice in females for certain diet-derived male pheromones can be exploited by maternal effect genes that preferentially place offspring on a specific host plant, resulting in specialization. Our analytical model also suggests that the process is more likely to occur with species that show male-congregating mating strategies, such as lekking and hilltopping. The model offers a new explanation for the similarity between the composition of male lepidopteran pheromones and the chemistry of their host plants and also suggests a novel mechanism of host plant shift. This is the first time that sexual selection has been proposed to drive host plant specialization and the first time that a mechanism with selection acting solely on the adult stage has been shown to be capable of determining larval feeding habits.

%B American Naturalist %V 169 %P 830-836 %8 Jun %@ 0003-0147 %G English %M WOS:000246553900013 %! Am NatAm Nat %0 Journal Article %J Evolution %D 2007 %T Karyotypic diversity and speciation in Agrodiaetus butterflies %A Kandul, N. R. %A Lukhtanov, V. A. %A Pierce, N. E. %K agrodiaetus %K beetles coleoptera %K chromosomal evolution %K chromosomal speciation %K comparative phylogenetic analysis %K differential fusion %K divergence times %K genetic approach %K maximum-likelihood %K metaphase-i bivalents %K mitochondrial-DNA %K postzygotic isolation %K rapid speciation %K species richness %X

That chromosomal rearrangements may play an important role in maintaining postzygotic isolation between well-established species is part of the standard theory of speciation. However, little evidence exists on the role of karyotypic change in speciation itself-in the establishment of reproductive barriers between previously interbreeding populations. The large genus' Agrodiaetus (Lepidoptera: Lycaenidae) provides a model system to study this question. Agrodiaetus butterflies exhibit unusual interspecific diversity in chromosome number, from n = 10 to n = 134; in contrast, the majority of lycaenid butterflies have n = 23/24. We analyzed the evolution of karyotypic diversity by mapping chromosome numbers on a thoroughly sampled mitochondrial phylogeny of the genus. Karyotypic differences accumulate gradually between allopatric sister taxa, but more rapidly between sympatric sister taxa. Overall, sympatric sister taxa have a higher average karyotypic diversity than allopatric sister taxa. Differential fusion of diverged populations may account for this pattern because the degree of karyotypic difference acquired between allopatric populations may determine whether they will persist as nascent biological species in secondary sympatry. This study therefore finds evidence of a direct role for chromosomal rearrangements in the final stages of animal speciation. Rapid karyotypic diversification is likely to have contributed to the explosive speciation rate observed in Agrodiaetus, 1.6 species per million years.

%B Evolution %V 61 %P 546-559 %8 Mar %@ 0014-3820 %G English %M WOS:000245133900006 %! EvolutionEvolution %0 Journal Article %J Biological Journal of the Linnean Society %D 2007 %T Phylogeny and historical biogeography of the subtribe Aporiina (Lepidoptera : Pieridae): implications for the origin of Australian butterflies %A Braby, M. F. %A Pierce, N. E. %A Vila, R. %K australian origins %K cretaceous %K divergence times %K elongation factor-1-alpha %K eucheira-socialis %K gene-sequences %K gondwana %K greater india %K laurasia %K long-distance dispersal %K Madagascar %K molecular phylogenies %K nymphalidae lepidoptera %K oceanic dispersal %K passerine birds %K permutation tests %K Tertiary %K vicariance %K wingless gene %X

The Australian fauna is composed of several major biogeographical elements reflecting different spatial and temporal histories. Two groups of particular interest are the Gondwanan Element, reflecting an ancient origin in Gondwana or southern Gondwana (southern vicariance hypothesis), and the Asian Element, reflecting a more recent origin in Asia, Eurasia or Laurasia (northern dispersal hypothesis). Theories regarding the origin and evolution of butterflies (Hesperioidea, Papilionoidea) in Australia are controversial, with no clear consensus. Here, we investigate the phylogenetic and historical biogeographical relationships of the subtribe Aporiina, a widespread taxon with disjunct distributions in each of the major zoogeographical regions. Attention is paid to origins of the subtribe in the Australian Region for which several conflicting hypotheses have been proposed for the Old World genus Delias Hubner. Our phylogenetic reconstruction was based on analysis of fragments of two nuclear genes (elongation factor-1 alpha, wingless) and one mitochondrial gene (cytochrome oxidase subunit 1) for 30 taxa. Phylogenetic analyses based on maximum parsimony, maximum likelihood and Bayesian inference of the combined data set (2729 bp; 917 parsimony informative characters) recovered six major lineages within the monophyletic Aporiina, with the following topology: (Cepora + Prioneris + (Mylothris + (Aporia + Delias group + Catasticta group))). Given a probable age of origin of the stem-group near the Cretaceous/Tertiary boundary (69-54 Mya), followed by diversification of the crown-group in the early to mid Tertiary (57-45 Mya), we show that an origin of the Aporiina in either southern Gondwana or Laurasia is equally parsimonious, and that dispersal has played a major role in shaping the underlying phylogenetic pattern. We tentatively conclude that an origin in southern Gondwanan is more likely; however, neither hypothesis satisfactorily explains the present-day distribution, and additional lower-level phylogenies are needed to determine the directionality of dispersal events of several taxa and to reject one hypothesis over the other. Dispersal is inferred to have occurred primarily during cooler periods when land bridges or stepping-stones were available between many of the zoogeographical regions. (c) 2007 The Linnean Society of London.

%B Biological Journal of the Linnean Society %V 90 %P 413-440 %8 Mar %@ 0024-4066 %G English %M WOS:000245107100003 %! Biol J Linn SocBiol J Linn Soc %0 Journal Article %J Systematic Entomology %D 2007 %T Systematics, biogeography and diversification of the Indo-Australian genus Delias Hubner (Lepidoptera : Pieridae): phylogenetic evidence supports an 'out-of-Australia' origin %A Braby, M. F. %A Pierce, N. E. %K butterflies %K elongation factor-1-alpha %K Evolution %K historical biogeography %K limits %K mitochondrial nd5 gene %K molecular phylogeny %K permutation tests %K sequence data %K substitution %X

Two alternative hypotheses for the origin of butterflies in the Australian Region, that elements dispersed relatively recently from the Oriental Region into Australia (northern dispersal hypothesis) or descended from ancient stocks in Gondwana (southern vicariance hypothesis), were tested using methods of cladistic vicariance biogeography for the Delias group, a diverse and widespread clade in the Indo-Australian Region. A phylogenetic hypothesis of the twenty-four species-groups recognized currently in Delias and its sister genus Leuciacria is inferred from molecular characters generated from the nuclear gene elongation factor-1 alpha (EF-1 alpha) and the mitochondrial genes cytochrome oxidase subunits I and II (COI/COII) and NADH dehydrogenase 5 (ND5). Phylogenetic analyses based on maximum parsimony, maximum likelihood and Bayesian inference of the combined dataset (3888 bp, 1014 parsimony informative characters) confirmed the monophyly of Delias and recovered eight major lineages within the genus, informally designated the singhapura, belladonna, hyparete, chrysomelaena, eichhorni, cuningputi, belisama and nigrina clades. Species-group relationships within these clades are, in general, concordant with current systematic arrangements based on morphology. The major discrepancies concern the placement of the aganippe, belisama and chrysomelaena groups, as well as several species-groups endemic to mainland New Guinea. Two species (D. harpalyce (Donovan), D. messalina Arora) of uncertain group status are currently misplaced based on strong evidence of paraphyly, and are accordingly transferred to the nigrina and kummeri groups, respectively. Based on this phylogeny, a revised systematic classification is presented at the species-group level. An historical biogeographical analysis of the Delias group revealed that the most parsimonious reconstruction is an origin in the Australian Region, with at least seven dispersal events across Wallacea to the Oriental Region. The eight major clades of Delias appear to have diverged rapidly following complete separation of the Australian plate from Gondwana and its collision with the Asian plate in the late Oligocene. Further diversification and dispersal of Delias in the Miocene-Pliocene are associated with major geological and climatic changes that occurred in Australia-New Guinea during the late Tertiary. The 'out-of-Australia' hypothesis for the Delias group supports an origin of the Aporiina in southern Gondwana (southern vicariance hypothesis), which proposes that the ancestor of Delias + Leuciacria differentiated vicariantly on the Australian plate.

%B Systematic Entomology %V 32 %P 2-25 %8 Jan %@ 0307-6970 %G English %M WOS:000242992800002 %! Syst EntomolSyst Entomol %0 Journal Article %J Nature %D 2007 %T Dating the origin of the Orchidaceae from a fossil orchid with its pollinator %A Ramirez, S. R. %A Gravendeel, B. %A Singer, R. B. %A Marshall, C. R. %A Pierce, N. E. %K age %K amber %K angiosperms %K Evolution %K flowering plants %K Morphology %K sequences %X

Since the time of Darwin(1), evolutionary biologists have been fascinated by the spectacular adaptations to insect pollination exhibited by orchids. However, despite being the most diverse plant family on Earth(2), the Orchidaceae lack a definitive fossil record and thus many aspects of their evolutionary history remain obscure. Here we report an exquisitely preserved orchid pollinarium (of Meliorchis caribea gen. et sp. nov.) attached to the mesoscutellum of an extinct stingless bee, Proplebeia dominicana, recovered from Miocene amber in the Dominican Republic, that is 15-20 million years (Myr) old(3). This discovery constitutes both the first unambiguous fossil of Orchidaceae(4) and an unprecedented direct fossil observation of a plant-pollinator interaction(5,6). By applying cladistic methods to a morphological character matrix, we resolve the phylogenetic position of M. caribea within the extant subtribe Goodyerinae (subfamily Orchidoideae). We use the ages of other fossil monocots and M. caribea to calibrate a molecular phylogenetic tree of the Orchidaceae. Our results indicate that the most recent common ancestor of extant orchids lived in the Late Cretaceous (76-84 Myr ago), and also suggest that the dramatic radiation of orchids began shortly after the mass extinctions at the K/T boundary. These results further support the hypothesis of an ancient origin for Orchidaceae.

%B Nature %V 448 %P 1042-1045 %8 Aug 30 %@ 0028-0836 %G English %M WOS:000249097600036 %! NatureNature %0 Journal Article %J Molecular Ecology %D 2007 %T The geography of diversification in mutualistic ants: a gene's-eye view into the Neogene history of Sundaland rain forests %A Quek, S. P. %A Davies, S. J. %A Ashton, P. S. %A Itino, T. %A Pierce, N. E. %K association %K biogeography %K cospeciation %K crematogaster %K cytochrome oxidase %K formicidae %K macaranga %K macaranga euphorbiaceae %K mitochondrial-DNA %K molecular clock %K polymorphism %K population-growth %K rain forest %K southeast asia %K southeast-asia %X

We investigate the geographical and historical context of diversification in a complex of mutualistic Crematogaster ants living in Macaranga trees in the equatorial rain forests of Southeast Asia. Using mitochondrial DNA from 433 ant colonies collected from 32 locations spanning Borneo, Malaya and Sumatra, we infer branching relationships, patterns of genetic diversity and population history. We reconstruct a time frame for the ants' diversification and demographic expansions, and identify areas that might have been refugia or centres of diversification. Seventeen operational lineages are identified, most of which can be distinguished by host preference and geographical range. The ants first diversified 16-20 Ma, not long after the onset of the everwet forests in Sundaland, and achieved most of their taxonomic diversity during the Pliocene. Pleistocene demographic expansions are inferred for several of the younger lineages. Phylogenetic relationships suggest a Bornean cradle and major axis of diversification. Taxonomic diversity tends to be associated with mountain ranges; in Borneo, it is greatest in the Crocker Range of Sabah and concentrated also in other parts of the northern northwest coast. Within-lineage genetic diversity in Malaya and Sumatra tends to also coincide with mountain ranges. A series of disjunct and restricted distributions spanning northern northwest Borneo and the major mountain ranges of Malaya and Sumatra, seen in three pairs of sister lineages, further suggests that these regions were rain-forest refuges during drier climatic phases of the Pleistocene. Results are discussed in the context of the history of Sundaland's rain forests.

%B Molecular Ecology %V 16 %P 2045-2062 %8 May %@ 0962-1083 %G English %M WOS:000246366800006 %! Mol EcolMol Ecol %0 Journal Article %J American Naturalist %D 2007 %T Host plant specialization driven by sexual selection %A Quental, T. B. %A Patten, M. M. %A Pierce, N. E. %K danaid butterflies %K defense-mechanisms %K diet breadth %K enemy-free space %K Evolution %K female choice %K host shift %K lepidoptera %K maternal effects %K moth utetheisa-ornatrix %K pheromone %K phytophagous insects %K pyrrolizidine alkaloids %K specialization %X

We propose a new mechanism based on sexual selection to explain the evolution of diet breadth in insects. More specifically, we show that mate choice in females for certain diet-derived male pheromones can be exploited by maternal effect genes that preferentially place offspring on a specific host plant, resulting in specialization. Our analytical model also suggests that the process is more likely to occur with species that show male-congregating mating strategies, such as lekking and hilltopping. The model offers a new explanation for the similarity between the composition of male lepidopteran pheromones and the chemistry of their host plants and also suggests a novel mechanism of host plant shift. This is the first time that sexual selection has been proposed to drive host plant specialization and the first time that a mechanism with selection acting solely on the adult stage has been shown to be capable of determining larval feeding habits.

%B American Naturalist %V 169 %P 830-836 %8 Jun %@ 0003-0147 %G English %M WOS:000246553900013 %! Am NatAm Nat %0 Journal Article %J Evolution %D 2007 %T Karyotypic diversity and speciation in Agrodiaetus butterflies %A Kandul, N. R. %A Lukhtanov, V. A. %A Pierce, N. E. %K agrodiaetus %K beetles coleoptera %K chromosomal evolution %K chromosomal speciation %K comparative phylogenetic analysis %K differential fusion %K divergence times %K genetic approach %K maximum-likelihood %K metaphase-i bivalents %K mitochondrial-DNA %K postzygotic isolation %K rapid speciation %K species richness %X

That chromosomal rearrangements may play an important role in maintaining postzygotic isolation between well-established species is part of the standard theory of speciation. However, little evidence exists on the role of karyotypic change in speciation itself-in the establishment of reproductive barriers between previously interbreeding populations. The large genus' Agrodiaetus (Lepidoptera: Lycaenidae) provides a model system to study this question. Agrodiaetus butterflies exhibit unusual interspecific diversity in chromosome number, from n = 10 to n = 134; in contrast, the majority of lycaenid butterflies have n = 23/24. We analyzed the evolution of karyotypic diversity by mapping chromosome numbers on a thoroughly sampled mitochondrial phylogeny of the genus. Karyotypic differences accumulate gradually between allopatric sister taxa, but more rapidly between sympatric sister taxa. Overall, sympatric sister taxa have a higher average karyotypic diversity than allopatric sister taxa. Differential fusion of diverged populations may account for this pattern because the degree of karyotypic difference acquired between allopatric populations may determine whether they will persist as nascent biological species in secondary sympatry. This study therefore finds evidence of a direct role for chromosomal rearrangements in the final stages of animal speciation. Rapid karyotypic diversification is likely to have contributed to the explosive speciation rate observed in Agrodiaetus, 1.6 species per million years.

%B Evolution %V 61 %P 546-559 %8 Mar %@ 0014-3820 %G English %M WOS:000245133900006 %! EvolutionEvolution %0 Journal Article %J Biological Journal of the Linnean Society %D 2007 %T Phylogeny and historical biogeography of the subtribe Aporiina (Lepidoptera : Pieridae): implications for the origin of Australian butterflies %A Braby, M. F. %A Pierce, N. E. %A Vila, R. %K australian origins %K cretaceous %K divergence times %K elongation factor-1-alpha %K eucheira-socialis %K gene-sequences %K gondwana %K greater india %K laurasia %K long-distance dispersal %K Madagascar %K molecular phylogenies %K nymphalidae lepidoptera %K oceanic dispersal %K passerine birds %K permutation tests %K Tertiary %K vicariance %K wingless gene %X

The Australian fauna is composed of several major biogeographical elements reflecting different spatial and temporal histories. Two groups of particular interest are the Gondwanan Element, reflecting an ancient origin in Gondwana or southern Gondwana (southern vicariance hypothesis), and the Asian Element, reflecting a more recent origin in Asia, Eurasia or Laurasia (northern dispersal hypothesis). Theories regarding the origin and evolution of butterflies (Hesperioidea, Papilionoidea) in Australia are controversial, with no clear consensus. Here, we investigate the phylogenetic and historical biogeographical relationships of the subtribe Aporiina, a widespread taxon with disjunct distributions in each of the major zoogeographical regions. Attention is paid to origins of the subtribe in the Australian Region for which several conflicting hypotheses have been proposed for the Old World genus Delias Hubner. Our phylogenetic reconstruction was based on analysis of fragments of two nuclear genes (elongation factor-1 alpha, wingless) and one mitochondrial gene (cytochrome oxidase subunit 1) for 30 taxa. Phylogenetic analyses based on maximum parsimony, maximum likelihood and Bayesian inference of the combined data set (2729 bp; 917 parsimony informative characters) recovered six major lineages within the monophyletic Aporiina, with the following topology: (Cepora + Prioneris + (Mylothris + (Aporia + Delias group + Catasticta group))). Given a probable age of origin of the stem-group near the Cretaceous/Tertiary boundary (69-54 Mya), followed by diversification of the crown-group in the early to mid Tertiary (57-45 Mya), we show that an origin of the Aporiina in either southern Gondwana or Laurasia is equally parsimonious, and that dispersal has played a major role in shaping the underlying phylogenetic pattern. We tentatively conclude that an origin in southern Gondwanan is more likely; however, neither hypothesis satisfactorily explains the present-day distribution, and additional lower-level phylogenies are needed to determine the directionality of dispersal events of several taxa and to reject one hypothesis over the other. Dispersal is inferred to have occurred primarily during cooler periods when land bridges or stepping-stones were available between many of the zoogeographical regions. (c) 2007 The Linnean Society of London.

%B Biological Journal of the Linnean Society %V 90 %P 413-440 %8 Mar %@ 0024-4066 %G English %M WOS:000245107100003 %! Biol J Linn SocBiol J Linn Soc %0 Journal Article %J Systematic Entomology %D 2007 %T Systematics, biogeography and diversification of the Indo-Australian genus Delias Hubner (Lepidoptera : Pieridae): phylogenetic evidence supports an 'out-of-Australia' origin %A Braby, M. F. %A Pierce, N. E. %K butterflies %K elongation factor-1-alpha %K Evolution %K historical biogeography %K limits %K mitochondrial nd5 gene %K molecular phylogeny %K permutation tests %K sequence data %K substitution %X

Two alternative hypotheses for the origin of butterflies in the Australian Region, that elements dispersed relatively recently from the Oriental Region into Australia (northern dispersal hypothesis) or descended from ancient stocks in Gondwana (southern vicariance hypothesis), were tested using methods of cladistic vicariance biogeography for the Delias group, a diverse and widespread clade in the Indo-Australian Region. A phylogenetic hypothesis of the twenty-four species-groups recognized currently in Delias and its sister genus Leuciacria is inferred from molecular characters generated from the nuclear gene elongation factor-1 alpha (EF-1 alpha) and the mitochondrial genes cytochrome oxidase subunits I and II (COI/COII) and NADH dehydrogenase 5 (ND5). Phylogenetic analyses based on maximum parsimony, maximum likelihood and Bayesian inference of the combined dataset (3888 bp, 1014 parsimony informative characters) confirmed the monophyly of Delias and recovered eight major lineages within the genus, informally designated the singhapura, belladonna, hyparete, chrysomelaena, eichhorni, cuningputi, belisama and nigrina clades. Species-group relationships within these clades are, in general, concordant with current systematic arrangements based on morphology. The major discrepancies concern the placement of the aganippe, belisama and chrysomelaena groups, as well as several species-groups endemic to mainland New Guinea. Two species (D. harpalyce (Donovan), D. messalina Arora) of uncertain group status are currently misplaced based on strong evidence of paraphyly, and are accordingly transferred to the nigrina and kummeri groups, respectively. Based on this phylogeny, a revised systematic classification is presented at the species-group level. An historical biogeographical analysis of the Delias group revealed that the most parsimonious reconstruction is an origin in the Australian Region, with at least seven dispersal events across Wallacea to the Oriental Region. The eight major clades of Delias appear to have diverged rapidly following complete separation of the Australian plate from Gondwana and its collision with the Asian plate in the late Oligocene. Further diversification and dispersal of Delias in the Miocene-Pliocene are associated with major geological and climatic changes that occurred in Australia-New Guinea during the late Tertiary. The 'out-of-Australia' hypothesis for the Delias group supports an origin of the Aporiina in southern Gondwana (southern vicariance hypothesis), which proposes that the ancestor of Delias + Leuciacria differentiated vicariantly on the Australian plate.

%B Systematic Entomology %V 32 %P 2-25 %8 Jan %@ 0307-6970 %G English %M WOS:000242992800002 %! Syst EntomolSyst Entomol %0 Journal Article %J Ecological Entomology %D 2006 %T Convergence of chemical mimicry in a guild of aphid predators %A Lohman, D. J. %A Liao, Q. %A Pierce, N. E. %K ants hymenoptera %K aphidophagous hoverflies %K butterfly maculinea-rebeli %K chemical camouflage %K chemical mimicry %K colony odor %K cuticular hydrocarbons %K formicidae %K harvester ants %K host-specificity %K life history %K myrmecophily %K nestmate recognition %K obligate predator %K pheromone %K predation %K semiochemical %K tritrophic interaction %K trophobiont %X

1. A variety of insects prey on honeydew-producing Homoptera and many do so even in the presence of ants that tend, and endeavour to protect, these trophobionts from natural enemies. Few studies have explored the semiochemical mechanisms by which these predators circumvent attack by otherwise aggressive ants.2. Ants use specific mixtures of cuticular hydrocarbons (CHCs) as recognition labels, but this simple mechanism is frequently circumvented by nest parasites that engage in 'chemical mimicry' of their host ants by producing or acquiring a critical suite of these CHCs.3. Analysis of the CHCs from the North American woolly alder aphid, Prociphilus tessellatus (Homoptera: Aphididae), their tending ants, and aphid predators from three insect orders, Feniseca tarquinius (Lepidoptera: Lycaenidae), Chrysopa slossonae (Neuroptera: Chrysopidae), and Syrphus ribesii (Diptera: Syrphidae), showed that while the CHC profile of each predatory species was distinct, each was chemically more similar to the aphids than to either tending ant species. Further, the CHCs of each predator species were a subset of the compounds found in the aphids' profile.4. These results implicate CHCs as a recognition cue used by ants to discriminate trophobionts from potential prey and a probable mechanism by which trophobiont predators circumvent detection by aphids and their tending ants.5. Although several features of the aphids' CHC profile are shared among the chemically mimetic taxa, variation in the precision of mimicry among the members of this predatory guild demonstrates that a chemical mimic need not replicate every feature of its model.

%B Ecological Entomology %V 31 %P 41-51 %8 Feb %@ 0307-6946 %G English %M WOS:000236666900006 %! Ecol EntomolEcol Entomol %0 Journal Article %J Ecological Entomology %D 2006 %T Convergence of chemical mimicry in a guild of aphid predators. %A DJ Lohman %A Q Liao %A Pierce, N. E. %X

Abstract. 1. A variety of insects prey on honeydew-producing Homoptera and many do so even in the presence of ants that tend, and endeavour to protect, these trophobionts from natural enemies. Few studies have explored the semiochemical mechanisms by which these predators circumvent attack by otherwise aggressive ants. 2. Ants use specific mixtures of cuticular hydrocarbons (CHCs) as recognition labels, but this simple mechanism is frequently circumvented by nest parasites that engage in ‘chemical mimicry’ of their host ants by producing or acquiring a critical suite of these CHCs. 3. Analysis of the CHCs from the North American woolly alder aphid, Prociphilus tessellatus (Homoptera: Aphididae), their tending ants, and aphid predators from three insect orders, Feniseca tarquinius (Lepidoptera: Lycaenidae), Chrysopa slossonae (Neuroptera: Chrysopidae), and Syrphus ribesii (Diptera: Syrphidae), showed that while the CHC profile of each predatory species was distinct, each was chemically more similar to the aphids than to either tending ant species. Further, the CHCs of each predator species were a subset of the compounds found in the aphids’ profile. 4. These results implicate CHCs as a recognition cue used by ants to discriminate trophobionts from potential prey and a probable mechanism by which trophobiont predators circumvent detection by aphids and their tending ants. 5. Although several features of the aphids’ CHC profile are shared among the chemically mimetic taxa, variation in the precision of mimicry among the members of this predatory guild demonstrates that a chemical mimic need not replicate every feature of its model.

%B Ecological Entomology %V 31 %P 41-51 %G eng %0 Journal Article %J Zoological Journal of the Linnaean Society %D 2006 %T Molecular phylogeny and systematics of the Pieridae (Lepidoptera: Papilionoidea): higher classification and biogeography %A Braby, MF %A Villa, R. %A Pierce, N. E. %X

The systematic relationships of the butterfly family Pieridae are poorly understood. Much of our current understanding is based primarily on detailed morphological observations made 50–70 years ago. However, the family and its putative four subfamilies and two tribes, have rarely been subjected to rigorous phylogenetic analysis. Here we present results based on an analysis of molecular characters used to reconstruct the phylogeny of the Pieridae in order to infer higher-level classification above the generic level and patterns of historical biogeography. Our sample contained 90 taxa representing 74 genera and six subgenera, or 89% of all genera recognized in the family. Three complementary approaches were employed: (1) a combined analysis of a 30 taxon subset for sequences from four gene regions, including elongation factor-1 alpha (EF-1α), wingless, cytochrome oxidase subunit I (COI), and 28S (3675 bp, 1031 parsimony-informative characters), mainly to establish higher-level relationships, (2) a single-gene analysis of the 90 taxon data set for sequences from EF-1α (1066 bp, 364 parsimony-informative characters), mainly to establish lower-level relationships, and (3) an all available data analysis of the entire data set for sequences from the four genes, to recover both deep and shallow nodes. Analyses using maximum parsimony, maximum likelihood and Bayesian inference provided similar results. All supported monophyly for the four subfamilies but not for the two tribes, with the Anthocharidini polyphyletic and the Pierini paraphyletic. The combined and all available data analyses support the following relationships among the subfamilies: ((Pseudopontiinae + Dismorphiinae) + (Coliadinae + Pierinae)), corroborating Ehrlich’s 1958 phenetic hypothesis. On the basis of these analyses, and additional morphological and life history evidence, we propose a reclassification of the subfamily Pierinae into two tribes (Anthocharidini s.s., Pierini s.s.) and two informal groups (Colotis group, Leptosia), with the tribe Pierini s.s. subdivided into three subtribes (Appiadina, Pierina, Aporiina) and three genera (Elodina, Dixeia, Belenois) of uncertain status (incertae sedis). The combined and all available data analyses support the following relationships among the Pierinae: (Colotis group + Anthocharidini s.s. + Leptosia + (Elodina + ((Dixeia + Belenois) + Appiadina + Pierina + Aporiina))). Application of a molecular clock calibrated using fossil evidence and semiparametric rate smoothing suggests that divergence between the Pierina and Aporiina occurred no later than the Palaeocene (> 60 Myr). The minimum estimate for the age of the crown-group of the Pieridae was 112–82 Myr, with a mean of 95 Myr. A historical biogeographical hypothesis is proposed to explain the present-day distribution of the clade Pseudopontiinae + Dismorphiinae, which argues for an origin of the two subfamilies in western Gondwana (Africa + South America) during the Late Cretaceous.

%B Zoological Journal of the Linnaean Society %V 147 %P 239-275 %G eng %0 Journal Article %J Zoological Journal of the Linnean Society %D 2006 %T Molecular phylogeny and systematics of the Pieridae (Lepidoptera : Papilionoidea): higher classification and biogeography (vol 147, pg 239, 2006) %A Braby, M. F. %A Vila, R. %A Pierce, N. E. %B Zoological Journal of the Linnean Society %V 147 %P 417-417 %8 Jul %@ 0024-4082 %G English %M WOS:000238393900005 %! Zool J Linn Soc-LondZool J Linn Soc-Lond %0 Journal Article %J Science %D 2006 %T Phylogeny of the ants: Diversification in the age of angiosperms %A Moreau, C. S. %A Bell, C. D. %A Vila, R. %A Archibald, S. B. %A Pierce, N. E. %K beetles %K formicidae %K hymenoptera %K origin %K rdna sequences %X

We present a large-scale molecular phylogeny of the ants (Hymenoptera: Formicidae), based on 4.5 kilobases of sequence data from six gene regions extracted from 139 of the 288 described extant genera, representing 19 of the 20 subfamilies. All but two subfamilies are recovered as monophyletic. Divergence time estimates calibrated by minimum age constraints from 43 fossils indicate that most of the subfamilies representing extant ants arose much earlier than previously proposed but only began to diversify during the Late Cretaceous to Early Eocene. This period also witnessed the rise of angiosperms and most herbivorous insects.

%B Science %V 312 %P 101-104 %8 Apr 7 %@ 0036-8075 %G English %M WOS:000236584400040 %! ScienceScience %0 Journal Article %J Ecological Entomology %D 2006 %T Convergence of chemical mimicry in a guild of aphid predators. %A DJ Lohman %A Q Liao %A Pierce, N. E. %X

Abstract. 1. A variety of insects prey on honeydew-producing Homoptera and many do so even in the presence of ants that tend, and endeavour to protect, these trophobionts from natural enemies. Few studies have explored the semiochemical mechanisms by which these predators circumvent attack by otherwise aggressive ants. 2. Ants use specific mixtures of cuticular hydrocarbons (CHCs) as recognition labels, but this simple mechanism is frequently circumvented by nest parasites that engage in ‘chemical mimicry’ of their host ants by producing or acquiring a critical suite of these CHCs. 3. Analysis of the CHCs from the North American woolly alder aphid, Prociphilus tessellatus (Homoptera: Aphididae), their tending ants, and aphid predators from three insect orders, Feniseca tarquinius (Lepidoptera: Lycaenidae), Chrysopa slossonae (Neuroptera: Chrysopidae), and Syrphus ribesii (Diptera: Syrphidae), showed that while the CHC profile of each predatory species was distinct, each was chemically more similar to the aphids than to either tending ant species. Further, the CHCs of each predator species were a subset of the compounds found in the aphids’ profile. 4. These results implicate CHCs as a recognition cue used by ants to discriminate trophobionts from potential prey and a probable mechanism by which trophobiont predators circumvent detection by aphids and their tending ants. 5. Although several features of the aphids’ CHC profile are shared among the chemically mimetic taxa, variation in the precision of mimicry among the members of this predatory guild demonstrates that a chemical mimic need not replicate every feature of its model.

%B Ecological Entomology %V 31 %P 41-51 %G eng %0 Journal Article %J Evolution %D 2006 %T Do ants enhance diversification in lycaenid butterflies? Phylogeographic evidence from a model myrmecophile, Jalmenus evagoras. %A Eastwood, R. %A Pierce, N. E. %A RL Kitching %A JM Hughes %X

Abstract. The ant-tended Australian butterfly, Jalmenus evagoras, has been a model system for studying the ecology and evolution of mutualism. A phylogeographic analysis of mitochondrial DNA cytochrome oxidase I sequences from 242 butterflies (615 bp) and 66 attendant ants (585 bp) from 22 populations was carried out to explore the relationship between ant association and butterfly population structure. This analysis revealed 12 closely related butterfly haplotypes in three distinct clades roughly corresponding to three allopatric subpopulations of the butterflies. Minimal genetic diversity and widespread haplotypes within biogeographical regions suggest high levels of matrilineal gene flow. Attendant ants are significantly more diverse than was previously thought, with at least seven well-defined clades corresponding to independent morphological determinations, distributed throughout the range of the butterflies. Nested analysis of molecular variance showed that biogeography, host plant, and ant associate all contribute significantly in explaining variation in butterfly genetic diversity, but these variables are not independent of one another. Major influences appear to come from fragmentation due to large-scale biogeographical barriers, and diversification following a shift in habitat preference. A consequence of such a shift could be codiversification of the butterfly with habitatadapted ants, resulting in apparent phylogenetic concordance between butterflies and ants. The implications of these results are discussed in terms of possible effects of ant attendance on the diversification of Lycaenidae as a whole.

%B Evolution %V 60 %P 315-327 %G eng %0 Journal Article %J Zoological Journal of the Linnaean Society %D 2006 %T Molecular phylogeny and systematics of the Pieridae (Lepidoptera: Papilionoidea): higher classification and biogeography %A Braby, MF %A Villa, R. %A Pierce, N. E. %X

The systematic relationships of the butterfly family Pieridae are poorly understood. Much of our current understanding is based primarily on detailed morphological observations made 50–70 years ago. However, the family and its putative four subfamilies and two tribes, have rarely been subjected to rigorous phylogenetic analysis. Here we present results based on an analysis of molecular characters used to reconstruct the phylogeny of the Pieridae in order to infer higher-level classification above the generic level and patterns of historical biogeography. Our sample contained 90 taxa representing 74 genera and six subgenera, or 89% of all genera recognized in the family. Three complementary approaches were employed: (1) a combined analysis of a 30 taxon subset for sequences from four gene regions, including elongation factor-1 alpha (EF-1α), wingless, cytochrome oxidase subunit I (COI), and 28S (3675 bp, 1031 parsimony-informative characters), mainly to establish higher-level relationships, (2) a single-gene analysis of the 90 taxon data set for sequences from EF-1α (1066 bp, 364 parsimony-informative characters), mainly to establish lower-level relationships, and (3) an all available data analysis of the entire data set for sequences from the four genes, to recover both deep and shallow nodes. Analyses using maximum parsimony, maximum likelihood and Bayesian inference provided similar results. All supported monophyly for the four subfamilies but not for the two tribes, with the Anthocharidini polyphyletic and the Pierini paraphyletic. The combined and all available data analyses support the following relationships among the subfamilies: ((Pseudopontiinae + Dismorphiinae) + (Coliadinae + Pierinae)), corroborating Ehrlich’s 1958 phenetic hypothesis. On the basis of these analyses, and additional morphological and life history evidence, we propose a reclassification of the subfamily Pierinae into two tribes (Anthocharidini s.s., Pierini s.s.) and two informal groups (Colotis group, Leptosia), with the tribe Pierini s.s. subdivided into three subtribes (Appiadina, Pierina, Aporiina) and three genera (Elodina, Dixeia, Belenois) of uncertain status (incertae sedis). The combined and all available data analyses support the following relationships among the Pierinae: (Colotis group + Anthocharidini s.s. + Leptosia + (Elodina + ((Dixeia + Belenois) + Appiadina + Pierina + Aporiina))). Application of a molecular clock calibrated using fossil evidence and semiparametric rate smoothing suggests that divergence between the Pierina and Aporiina occurred no later than the Palaeocene (> 60 Myr). The minimum estimate for the age of the crown-group of the Pieridae was 112–82 Myr, with a mean of 95 Myr. A historical biogeographical hypothesis is proposed to explain the present-day distribution of the clade Pseudopontiinae + Dismorphiinae, which argues for an origin of the two subfamilies in western Gondwana (Africa + South America) during the Late Cretaceous.

%B Zoological Journal of the Linnaean Society %V 147 %P 239-275 %G eng %0 Journal Article %J Ecological Entomology %D 2006 %T Convergence of chemical mimicry in a guild of aphid predators %A Lohman, D. J. %A Liao, Q. %A Pierce, N. E. %K ants hymenoptera %K aphidophagous hoverflies %K butterfly maculinea-rebeli %K chemical camouflage %K chemical mimicry %K colony odor %K cuticular hydrocarbons %K formicidae %K harvester ants %K host-specificity %K life history %K myrmecophily %K nestmate recognition %K obligate predator %K pheromone %K predation %K semiochemical %K tritrophic interaction %K trophobiont %X

1. A variety of insects prey on honeydew-producing Homoptera and many do so even in the presence of ants that tend, and endeavour to protect, these trophobionts from natural enemies. Few studies have explored the semiochemical mechanisms by which these predators circumvent attack by otherwise aggressive ants.2. Ants use specific mixtures of cuticular hydrocarbons (CHCs) as recognition labels, but this simple mechanism is frequently circumvented by nest parasites that engage in 'chemical mimicry' of their host ants by producing or acquiring a critical suite of these CHCs.3. Analysis of the CHCs from the North American woolly alder aphid, Prociphilus tessellatus (Homoptera: Aphididae), their tending ants, and aphid predators from three insect orders, Feniseca tarquinius (Lepidoptera: Lycaenidae), Chrysopa slossonae (Neuroptera: Chrysopidae), and Syrphus ribesii (Diptera: Syrphidae), showed that while the CHC profile of each predatory species was distinct, each was chemically more similar to the aphids than to either tending ant species. Further, the CHCs of each predator species were a subset of the compounds found in the aphids' profile.4. These results implicate CHCs as a recognition cue used by ants to discriminate trophobionts from potential prey and a probable mechanism by which trophobiont predators circumvent detection by aphids and their tending ants.5. Although several features of the aphids' CHC profile are shared among the chemically mimetic taxa, variation in the precision of mimicry among the members of this predatory guild demonstrates that a chemical mimic need not replicate every feature of its model.

%B Ecological Entomology %V 31 %P 41-51 %8 Feb %@ 0307-6946 %G English %M WOS:000236666900006 %! Ecol EntomolEcol Entomol %0 Journal Article %J Zoological Journal of the Linnean Society %D 2006 %T Molecular phylogeny and systematics of the Pieridae (Lepidoptera : Papilionoidea): higher classification and biogeography (vol 147, pg 239, 2006) %A Braby, M. F. %A Vila, R. %A Pierce, N. E. %B Zoological Journal of the Linnean Society %V 147 %P 417-417 %8 Jul %@ 0024-4082 %G English %M WOS:000238393900005 %! Zool J Linn Soc-LondZool J Linn Soc-Lond %0 Journal Article %J Science %D 2006 %T Phylogeny of the ants: Diversification in the age of angiosperms %A Moreau, C. S. %A Bell, C. D. %A Vila, R. %A Archibald, S. B. %A Pierce, N. E. %K beetles %K formicidae %K hymenoptera %K origin %K rdna sequences %X

We present a large-scale molecular phylogeny of the ants (Hymenoptera: Formicidae), based on 4.5 kilobases of sequence data from six gene regions extracted from 139 of the 288 described extant genera, representing 19 of the 20 subfamilies. All but two subfamilies are recovered as monophyletic. Divergence time estimates calibrated by minimum age constraints from 43 fossils indicate that most of the subfamilies representing extant ants arose much earlier than previously proposed but only began to diversify during the Late Cretaceous to Early Eocene. This period also witnessed the rise of angiosperms and most herbivorous insects.

%B Science %V 312 %P 101-104 %8 Apr 7 %@ 0036-8075 %G English %M WOS:000236584400040 %! ScienceScience %0 Journal Article %J Proceedings of the National Academy of Sciences of the United States of America %D 2005 %T Pseudomonas syringae manipulates systemic plant defenses against pathogens and herbivores %A Cui, J. %A Bahrami, A. K. %A Pringle, E. G. %A Hernandez-Guzman, G. %A Bender, C. L. %A Pierce, N. E. %A Ausubel, F. M. %K acquired-resistance %K arabidopsis %K arabidopsis-thaliana %K coronatine %K cross-talk %K gene-expression %K induced disease resistance %K induced susceptibility %K jasmonic acid %K methyl jasmonate %K npr1-independent expression %K phytotoxin coronatine %K salicylic-acid %K trichoplusia ni %X

Many pathogens are virulent because they specifically interfere with host defense responses and therefore can proliferate. Here, we report that virulent strains of the bacterial phytopathogen Pseudomonas syringae induce systemic susceptibility to secondary A syringae infection in the host plant Arabidopsis thaliana. This systemic induced susceptibility (SIS) is in direct contrast to the well studied avirulence/R gene-dependent resistance response known as the hypersensitive response that elicits systemic acquired resistance. We show that A syringae-elicited SIS is caused by the production of coronatine (COR), a pathogen -derived functional and structural mimic of the phytohormone jasmonic acid (JA). These data suggest that SIS may be a consequence of the previously described mutually antagonistic interaction between the salicylic acid and JA signaling pathways. Virulent P. syringae also has the potential to induce net systemic susceptibility to herbivory by an insect (Trichoplusia ni, cabbage looper), but this susceptibility is not caused by COR. Rather, consistent with its role as a A mimic, COR induces systemic resistance to T. ni. These data highlight the complexity of defense signaling interactions among plants, pathogens, and herbivores.

%B Proceedings of the National Academy of Sciences of the United States of America %V 102 %P 1791-1796 %8 Feb 1 %@ 0027-8424 %G English %M WOS:000226877300095 %! P Natl Acad Sci USAP Natl Acad Sci USA %0 Journal Article %J Nature %D 2005 %T Reinforcement of pre-zygotic isolation and karyotype evolution in Agrodiaetus butterflies %A Lukhtanov, V. A. %A Kandul, N. P. %A Plotkin, J. B. %A Dantchenko, A. V. %A Haig, D. %A Pierce, N. E. %K driven %K Drosophila %K lepidoptera %K patterns %K phylogenies %K sexual selection %K Speciation %X

The reinforcement model of evolution argues that natural selection enhances pre-zygotic isolation between divergent populations or species by selecting against unfit hybrids(1,2) or costly interspecific matings(3). Reinforcement is distinguished from other models that consider the formation of reproductive isolation to be a by-product of divergent evolution(4,5). Although theory has shown that reinforcement is a possible mechanism that can lead to speciation(6-8), empirical evidence has been sufficiently scarce to raise doubts about the importance of reinforcement in nature(6,9,10). Agrodiaetus butterflies ( Lepidoptera: Lycaenidae) exhibit unusual variability in chromosome number. Whereas their genitalia and other morphological characteristics are largely uniform, different species vary considerably in male wing colour, and provide a model system to study the role of reinforcement in speciation. Using comparative phylogenetic methods, we show that the sympatric distribution of 15 relatively young sister taxa of Agrodiaetus strongly correlates with differences in male wing colour, and that this pattern is most likely the result of reinforcement. We find little evidence supporting sympatric speciation: rather, in Agrodiaetus, karyotypic changes accumulate gradually in allopatry, prompting reinforcement when karyotypically divergent races come into contact.

%B Nature %V 436 %P 385-389 %8 Jul 21 %@ 0028-0836 %G English %M WOS:000230623400041 %! NatureNature %0 Journal Article %J Proceedings of the Royal Society B-Biological Sciences %D 2005 %T Synergistic effects of combining morphological and molecular data in resolving the phylogeny of butterflies and skippers %A Wahlberg, N. %A Braby, M. F. %A Brower, A. V. Z. %A de Jong, R. %A Lee, M. M. %A Nylin, S. %A Pierce, N. E. %A Sperling, F. A. H. %A Vila, R. %A Warren, A. D. %A Zakharov, E. %K data sets %K family %K gene-sequences %K insecta %K molecular systematics %K nuclear gene %K nymphalidae lepidoptera %K papilionoidea %K reconstruction %K riodinidae lepidoptera %K Systematics %K total evidence %K wingless gene %X

Phylogenetic relationships among major clades of butterflies and skippers have long been controversial, with no general consensus even today. Such lack of resolution is a substantial impediment to using the otherwise well studied butterflies as a model group in biology. Here we report the results of a combined analysis of DNA sequences from three genes and a morphological data matrix for 57 taxa (3258 characters, 1290 parsimony informative) representing all major lineages from the three putative butterfly super-families (Hedyloidea, Hesperioidea and Papilionoidea), plus out-groups representing other ditrysian Lepidoptera families. Recently, the utility of morphological data as a source of phylogenetic evidence has been debated. We present the first well supported phylogenetic hypothesis for the butterflies and skippers based on a total-evidence analysis of both traditional morphological characters and new molecular characters from three gene regions (GOI, EF-1 alpha and wingless). All four data partitions show substantial hidden support for the deeper nodes, which emerges only in a combined analysis in which the addition of morphological data plays a crucial role. With the exception of Nymphalidae, the traditionally recognized families are found to be strongly supported monophyletic clades with the following relationships: (Hesperiidae+(Papilionidae+(Pieridae+(Nymphalidae+(Lycaenidae+Riodinidae))))). Nymphalidae is recovered as a monophyletic clade but this clade does not have strong support. Lycaenidae and Riodinidae are sister groups with strong support and we suggest that the latter be given family rank. The position of Pieridae as the sister taxon to nymphalids, lycaenids and riodinids is supported by morphology and the EF-1 alpha data but conflicted by the COI and wingless data. Hedylidae are more likely to be related to butterflies and skippers than geometrid moths and appear to be the sister group to Papilionoidea + Hesperioidea.

%B Proceedings of the Royal Society B-Biological Sciences %V 272 %P 1577-1586 %8 Aug 7 %@ 0962-8452 %G English %M WOS:000231504300008 %! P Roy Soc B-Biol SciP Roy Soc B-Biol Sci %0 Journal Article %J Proceedings of the National Academy of Sciences of the United States of America %D 2005 %T Pseudomonas syringae manipulates systemic plant defenses against pathogens and herbivores %A Cui, J. %A Bahrami, A. K. %A Pringle, E. G. %A Hernandez-Guzman, G. %A Bender, C. L. %A Pierce, N. E. %A Ausubel, F. M. %K acquired-resistance %K arabidopsis %K arabidopsis-thaliana %K coronatine %K cross-talk %K gene-expression %K induced disease resistance %K induced susceptibility %K jasmonic acid %K methyl jasmonate %K npr1-independent expression %K phytotoxin coronatine %K salicylic-acid %K trichoplusia ni %X

Many pathogens are virulent because they specifically interfere with host defense responses and therefore can proliferate. Here, we report that virulent strains of the bacterial phytopathogen Pseudomonas syringae induce systemic susceptibility to secondary A syringae infection in the host plant Arabidopsis thaliana. This systemic induced susceptibility (SIS) is in direct contrast to the well studied avirulence/R gene-dependent resistance response known as the hypersensitive response that elicits systemic acquired resistance. We show that A syringae-elicited SIS is caused by the production of coronatine (COR), a pathogen -derived functional and structural mimic of the phytohormone jasmonic acid (JA). These data suggest that SIS may be a consequence of the previously described mutually antagonistic interaction between the salicylic acid and JA signaling pathways. Virulent P. syringae also has the potential to induce net systemic susceptibility to herbivory by an insect (Trichoplusia ni, cabbage looper), but this susceptibility is not caused by COR. Rather, consistent with its role as a A mimic, COR induces systemic resistance to T. ni. These data highlight the complexity of defense signaling interactions among plants, pathogens, and herbivores.

%B Proceedings of the National Academy of Sciences of the United States of America %V 102 %P 1791-1796 %8 Feb 1 %@ 0027-8424 %G English %M WOS:000226877300095 %! P Natl Acad Sci USAP Natl Acad Sci USA %0 Journal Article %J Nature %D 2005 %T Reinforcement of pre-zygotic isolation and karyotype evolution in Agrodiaetus butterflies %A Lukhtanov, V. A. %A Kandul, N. P. %A Plotkin, J. B. %A Dantchenko, A. V. %A Haig, D. %A Pierce, N. E. %K driven %K Drosophila %K lepidoptera %K patterns %K phylogenies %K sexual selection %K Speciation %X

The reinforcement model of evolution argues that natural selection enhances pre-zygotic isolation between divergent populations or species by selecting against unfit hybrids(1,2) or costly interspecific matings(3). Reinforcement is distinguished from other models that consider the formation of reproductive isolation to be a by-product of divergent evolution(4,5). Although theory has shown that reinforcement is a possible mechanism that can lead to speciation(6-8), empirical evidence has been sufficiently scarce to raise doubts about the importance of reinforcement in nature(6,9,10). Agrodiaetus butterflies ( Lepidoptera: Lycaenidae) exhibit unusual variability in chromosome number. Whereas their genitalia and other morphological characteristics are largely uniform, different species vary considerably in male wing colour, and provide a model system to study the role of reinforcement in speciation. Using comparative phylogenetic methods, we show that the sympatric distribution of 15 relatively young sister taxa of Agrodiaetus strongly correlates with differences in male wing colour, and that this pattern is most likely the result of reinforcement. We find little evidence supporting sympatric speciation: rather, in Agrodiaetus, karyotypic changes accumulate gradually in allopatry, prompting reinforcement when karyotypically divergent races come into contact.

%B Nature %V 436 %P 385-389 %8 Jul 21 %@ 0028-0836 %G English %M WOS:000230623400041 %! NatureNature %0 Journal Article %J Proceedings of the Royal Society B-Biological Sciences %D 2005 %T Synergistic effects of combining morphological and molecular data in resolving the phylogeny of butterflies and skippers %A Wahlberg, N. %A Braby, M. F. %A Brower, A. V. Z. %A de Jong, R. %A Lee, M. M. %A Nylin, S. %A Pierce, N. E. %A Sperling, F. A. H. %A Vila, R. %A Warren, A. D. %A Zakharov, E. %K data sets %K family %K gene-sequences %K insecta %K molecular systematics %K nuclear gene %K nymphalidae lepidoptera %K papilionoidea %K reconstruction %K riodinidae lepidoptera %K Systematics %K total evidence %K wingless gene %X

Phylogenetic relationships among major clades of butterflies and skippers have long been controversial, with no general consensus even today. Such lack of resolution is a substantial impediment to using the otherwise well studied butterflies as a model group in biology. Here we report the results of a combined analysis of DNA sequences from three genes and a morphological data matrix for 57 taxa (3258 characters, 1290 parsimony informative) representing all major lineages from the three putative butterfly super-families (Hedyloidea, Hesperioidea and Papilionoidea), plus out-groups representing other ditrysian Lepidoptera families. Recently, the utility of morphological data as a source of phylogenetic evidence has been debated. We present the first well supported phylogenetic hypothesis for the butterflies and skippers based on a total-evidence analysis of both traditional morphological characters and new molecular characters from three gene regions (GOI, EF-1 alpha and wingless). All four data partitions show substantial hidden support for the deeper nodes, which emerges only in a combined analysis in which the addition of morphological data plays a crucial role. With the exception of Nymphalidae, the traditionally recognized families are found to be strongly supported monophyletic clades with the following relationships: (Hesperiidae+(Papilionidae+(Pieridae+(Nymphalidae+(Lycaenidae+Riodinidae))))). Nymphalidae is recovered as a monophyletic clade but this clade does not have strong support. Lycaenidae and Riodinidae are sister groups with strong support and we suggest that the latter be given family rank. The position of Pieridae as the sister taxon to nymphalids, lycaenids and riodinids is supported by morphology and the EF-1 alpha data but conflicted by the COI and wingless data. Hedylidae are more likely to be related to butterflies and skippers than geometrid moths and appear to be the sister group to Papilionoidea + Hesperioidea.

%B Proceedings of the Royal Society B-Biological Sciences %V 272 %P 1577-1586 %8 Aug 7 %@ 0962-8452 %G English %M WOS:000231504300008 %! P Roy Soc B-Biol SciP Roy Soc B-Biol Sci %0 Journal Article %J Evolution %D 2004 %T Codiversification in an ant-plant mutualism: Stem texture and the evolution of host use in Crematogaster (Formicidae : Myrmicinae) inhabitants of Macaranga (Euphorbiaceae) %A Quek, S. P. %A Davies, S. J. %A Itino, T. %A Pierce, N. E. %K coevoltion %K coevolution %K cospeciation %K cytochrome-oxidase-i %K decacrema %K fig-pollinating wasps %K inferring phylogenies %K macaranga %K mitochondrial-DNA %K molecular phylogeny %K mtdna variation %K myrmecophyte %K nuclear-gene trees %K phytoecy %K phytophagous insects %K southeast asia %K species trees %X

We investigate the evolution of host association in a cryptic complex of mutualistic Crematogaster (Decacrema) ants that inhabits and defends Macaranga trees in Southeast Asia. Previous phylogenetic studies based on limited samplings of Decacrema present conflicting reconstructions of the evolutionary history of the association, inferring both cospeciation and the predominance of host shifts. We use cytochrome oxidase I (COI) to reconstruct phylogenetic relationships in a comprehensive sampling of the Decacrema inhabitants of Macaranga. Using a published Macaranga phylogeny, we test whether the ants and plants have cospeciated. The COI phylogeny reveals 10 well-supported lineages and an absence of cospeciation. Host shifts, however, have been constrained by stem traits that are themselves correlated with Macaranga phylogeny. Earlier lineages of Decacrema exclusively inhabit waxy stems, a basal state in the Pachystemon clade within Macaranga, whereas younger species of Pachystemon, characterized by nonwaxy stems, are inhabited only by younger lineages of Decacrema. Despite the absence of cospeciation, the correlated succession of stem texture in both phylogenies suggests that Decacrema and Pachystemon have diversified in association, or codiversified. Subsequent to the colonization of the Pachystemon clade, Decacrema expanded onto a second clade within Macaranga, inducing the development of myrmecophytism in the Pruinosae group. Confinement to the aseasonal wet climate zone of western Malesia suggests myrmecophytic Macaranga are no older than the wet forest community in Southeast Asia, estimated to be about 20 million years old (early Miocene). Our calculation of COI divergence rates from several published arthropod studies that relied on tenable calibrations indicates a generally conserved rate of approximately 1.5% per million years. Applying this rate to a rate-smoothed Bayesian chronogram of the ants, the Decacrema from Macaranga are inferred to be at least 12 million years old (mid-Miocene). However, using the extremes of rate variation in COI produces an age as recent as 6 million years. Our inferred timeline based on 1.5% per million years concurs with independent biogeographical events in the region reconstructed from palynological data, thus suggesting that the evolutionary histories of Decacrema and their Pachystemon hosts have been contemporaneous since the mid-Miocene. The evolution of myrmecophytism enabled Macaranga to radiate into enemy-free space, while the ants' diversification has been shaped by stem traits, host specialization, and geographic factors. We discuss the possibility that the ancient and exclusive association between Decacrema and Macaranga was facilitated by an impoverished diversity of myrmecophytes and phytoecious (obligately plant inhabiting) ants in the region.

%B Evolution %V 58 %P 554-570 %8 Mar %@ 0014-3820 %G English %M WOS:000220704800010 %! EvolutionEvolution %0 Journal Article %J Nature %D 2004 %T The evolution of alternative parasitic life histories in large blue butterflies %A Als, T. D. %A Vila, R. %A Kandul, N. P. %A Nash, D. R. %A Yen, S. H. %A Hsu, Y. F. %A Mignault, A. A. %A Boomsma, J. J. %A Pierce, N. E. %K adoption %K alcon %K ant-nests %K formicidae %K host-specificity %K lepidoptera %K lycaenidae %K maculinea butterflies %X

Large blue (Maculinea) butterflies are highly endangered throughout the Palaearctic region, and have been the focus of intense conservation research(1-3). In addition, their extraordinary parasitic lifestyles make them ideal for studies of life history evolution. Early instars consume flower buds of specific host plants, but later instars live in ant nests where they either devour the brood (predators), or are fed mouth-to-mouth by the adult ants (cuckoos). Here we present the phylogeny for the group, which shows that it is a monophyletic clade nested within Phengaris, a rare Oriental genus whose species have similar life histories(4,5). Cuckoo species are likely to have evolved from predatory ancestors. As early as five million years ago, two Maculinea clades diverged, leading to the different parasitic strategies seen in the genus today. Contrary to current belief, the two recognized cuckoo species show little genetic divergence and are probably a single ecologically differentiated species(6-10). On the other hand, some of the predatory morphospecies exhibit considerable genetic divergence and may contain cryptic species. These findings have important implications for conservation and reintroduction efforts.

%B Nature %V 432 %P 386-390 %8 Nov 18 %@ 0028-0836 %G English %M WOS:000225161400054 %! NatureNature %0 Journal Article %J Systematic Entomology %D 2004 %T Molecular phylogeny of the Oriental butterfly genus Arhopala (Lycaenidae, Theclinae) inferred from mitochondrial and nuclear genes %A Megens, H. J. %A Van Nes, W. J. %A Van Moorsel, C. H. M. %A Pierce, N. E. %A de Jong, R. %K compositional bias %K cytochrome-oxidase-i %K DNA %K Evolution %K Heliconius butterflies %K inference %K lepidoptera %K nucleotide-sequence %K nymphalidae %K wingless gene %X

We present a phylogeny for a selection of species of the butterfly genus Arhopala Boisduval, 1832 based on molecular characters. We sequenced 1778 bases of the mitochondrial genes Cytochrome Oxidase 1 and 2 including tRNA(Leu), and a 393-bp fragment of the nuclear wingless gene for a total of 42 specimens of 33 species, representing all major species groups. Analyses of mtDNA and wingless genes show congruent phylogenetic signal. The phylogeny presented here confirms the monophyly of the centaurus, eumolphus, camdeo and epimuta groups and the amphimuta subgroup. It confirms close relationships between species within the agelastus group, that together with the amphimuta subgroup, centaurus and camdeo groups form a monophyletic group. However, incongruencies with previous taxonomic studies also occur; the amphimuta and silhetensis groups are not monophyletic, as is the genus Arhopala itself. One enigmatic species, A. kinabala, was evaluated further for topology and the support for basal placement of this species is due mainly to the wingless gene. However, in the Parsimony analysis, and subsequent Maximum Likelihood evaluations, certain nodes could not be resolved due to insufficient support. The mtDNA shows extreme AT bias with compositional heterogeneity at 3rd codon positions, which may result in saturation. By contrast, the wingless gene does not show compositional bias, suggesting that poor support is not due solely to saturation. The evaluation of morphological characters used in previous studies on Arhopala systematics on the molecular tree indicates that the macular pattern and the absence of tails at the hind wings show extensive homoplasy. A significant phylogenetic signal (as indicated by T-PTP tests) is present in several of these morphological characters, which are nevertheless of limited use in phylogenetic studies due to their labile nature.

%B Systematic Entomology %V 29 %P 115-131 %8 Jan %@ 0307-6970 %G English %M WOS:000188849100007 %! Syst EntomolSyst Entomol %0 Journal Article %J Systematic Biology %D 2004 %T Phylogeny of Agrodiaetus Hubner 1822 (Lepidoptera : Lycaenidae) inferred from mtDNA sequences of COI and COII and nuclear sequences of EF1-alpha: Karyotype diversification and species radiation %A Kandul, N. P. %A Lukhtanov, V. A. %A Dantchenko, A. V. %A Coleman, J. W. S. %A Sekercioglu, C. H. %A Haig, D. %A Pierce, N. E. %K agrodiaetus %K bayesian-inference %K chromosomal features %K cytochrome oxidase i %K cytochrome oxidase ii %K drosophila-melanogaster %K elongation factor 1 alpha %K elongation factor-1-alpha %K evolutionary genetics %K karyotype diversification %K length difference test %K maximum-likelihood %K metaphase-i bivalents %K mitochondrial-DNA %K Phylogeny %K polyommatus %K Speciation %K transposable element %X

Butterflies in the large Palearctic genus Agrodiaetus ( Lepidoptera: Lycaenidae) are extremely uniform and exhibit few distinguishing morphological characters. However, these insects are distinctive in one respect: as a group they possess among the greatest interspecific karyotype diversity in the animal kingdom, with chromosome numbers (n) ranging from 10 to 125. The monophyly of Agrodiaetus and its systematic position relative to other groups within the section Polyommatus have been controversial. Characters from the mitochondrial genes for cytochrome oxidases I and II and from the nuclear gene for elongation factor 1alpha were used to reconstruct the phylogeny of Agrodiaetus using maximum parsimony and Bayesian phylogenetic methods. Ninety-one individuals, encompassing most of the taxonomic diversity of Agrodiaetus, and representatives of 14 related genera were included in this analysis. Our data indicate that Agrodiaetus is monophyletic. Representatives of the genus Polyommatus ( sensu stricto) are the closest relatives. The sequences of the Agrodiaetus taxa in this analysis are tentatively arranged into 12 clades, only 1 of which corresponds to a species group traditionally recognized in Agrodiaetus. Heterogeneous substitution rates across a recovered topology were homogenized with a nonparametric rate-smoothing algorithm before the application of a molecular clock. Two published estimates of substitution rates dated the origin of Agrodiaetus between 2.51 and 3.85 million years ago. During this time, there was heterogeneity in the rate and direction of karyotype evolution among lineages within the genus. Karyotype instability has evolved independently three times in the section Polyommatus, within the lineages Agrodiaetus, Lysandra, and Plebicula. Rapid karyotype diversification may have played a significant role in the radiation of the genus Agrodiaetus.

%B Systematic Biology %V 53 %P 278-298 %8 Apr %@ 1063-5157 %G English %M WOS:000220800500006 %! Syst BiolSyst Biol %0 Journal Article %J Molecular Phylogenetics and Evolution %D 2004 %T Tempo of speciation in a butterfly genus from the Southeast Asian tropics, inferred from mitochondrial and nuclear DNA sequence data %A Megens, H. J. %A Van Moorsel, C. H. M. %A Piel, W. H. %A Pierce, N. E. %A de Jong, R. %K arhopala %K canary-islands %K cytochrome-oxidase-i %K diversification %K Evolution %K hard polytomy %K Heliconius butterflies %K molecular clock %K molecular phylogenetics %K null model %K phylogenetic-relationships %K rapid radiation %K substitution rates %K tropical diversity %X

Molecular systematics is frequently beset with phylogenetic results that are not fully resolved. Researchers either state that the absence of resolution is due to character conflict, explosive speciation, or some combination of the two, but seldom do they carefully examine their data to distinguish between these causes. In this study, we exhaustively analyze a set of nuclear and mitochondrial nucleotide data for the Asian tropical butterfly genus Arhopala so as to highlight the causes of polytomies in the phylogenetic trees, and, as a result, to infer important biological events in the history of this genus. We began by using non-parametric statistical methods to determine whether the ambiguously resolved regions in these trees represent hard or soft polytomies. In addition we determined how this correlated to number of inferred changes on branches, using parametric maximum likelihood estimations. Based on congruent patterns in both mitochondrial and nuclear DNA sequences, we concluded that at two stages in the history of Arhopala there have been accelerated instances of speciation. One event, at the base of the phylogeny, generated many of the groups and subgroups currently recognized in this genus, while a later event generated another major clade consisting of both Oriental and Papuan species groups. Based on comparisons of closely related taxa, the ratio of instantaneous rate of evolution between mitochondrial and nuclear DNA evolution is established at approximately 3:1. The earliest radiation is dated between 7 and 11 Ma by a molecular clock analysis, setting the events generating much of the diversity of Arhopala at well before the Pleistocene. Periodical flooding of the Sunda plateau during interglacial periods was, therefore, not responsible for generating the major divisions in the genus Arhopala. Instead, we hypothesize that large-scale climatic changes taking place in the Miocene have induced the early acceleration in speciation. (C) 2003 Elsevier Inc. All rights reserved.

%B Molecular Phylogenetics and Evolution %V 31 %P 1181-1196 %8 Jun %@ 1055-7903 %G English %M WOS:000221443400029 %! Mol Phylogenet EvolMol Phylogenet Evol %0 Journal Article %J Evolution %D 2004 %T Codiversification in an ant-plant mutualism: Stem texture and the evolution of host use in Crematogaster (Formicidae : Myrmicinae) inhabitants of Macaranga (Euphorbiaceae) %A Quek, S. P. %A Davies, S. J. %A Itino, T. %A Pierce, N. E. %K coevoltion %K coevolution %K cospeciation %K cytochrome-oxidase-i %K decacrema %K fig-pollinating wasps %K inferring phylogenies %K macaranga %K mitochondrial-DNA %K molecular phylogeny %K mtdna variation %K myrmecophyte %K nuclear-gene trees %K phytoecy %K phytophagous insects %K southeast asia %K species trees %X

We investigate the evolution of host association in a cryptic complex of mutualistic Crematogaster (Decacrema) ants that inhabits and defends Macaranga trees in Southeast Asia. Previous phylogenetic studies based on limited samplings of Decacrema present conflicting reconstructions of the evolutionary history of the association, inferring both cospeciation and the predominance of host shifts. We use cytochrome oxidase I (COI) to reconstruct phylogenetic relationships in a comprehensive sampling of the Decacrema inhabitants of Macaranga. Using a published Macaranga phylogeny, we test whether the ants and plants have cospeciated. The COI phylogeny reveals 10 well-supported lineages and an absence of cospeciation. Host shifts, however, have been constrained by stem traits that are themselves correlated with Macaranga phylogeny. Earlier lineages of Decacrema exclusively inhabit waxy stems, a basal state in the Pachystemon clade within Macaranga, whereas younger species of Pachystemon, characterized by nonwaxy stems, are inhabited only by younger lineages of Decacrema. Despite the absence of cospeciation, the correlated succession of stem texture in both phylogenies suggests that Decacrema and Pachystemon have diversified in association, or codiversified. Subsequent to the colonization of the Pachystemon clade, Decacrema expanded onto a second clade within Macaranga, inducing the development of myrmecophytism in the Pruinosae group. Confinement to the aseasonal wet climate zone of western Malesia suggests myrmecophytic Macaranga are no older than the wet forest community in Southeast Asia, estimated to be about 20 million years old (early Miocene). Our calculation of COI divergence rates from several published arthropod studies that relied on tenable calibrations indicates a generally conserved rate of approximately 1.5% per million years. Applying this rate to a rate-smoothed Bayesian chronogram of the ants, the Decacrema from Macaranga are inferred to be at least 12 million years old (mid-Miocene). However, using the extremes of rate variation in COI produces an age as recent as 6 million years. Our inferred timeline based on 1.5% per million years concurs with independent biogeographical events in the region reconstructed from palynological data, thus suggesting that the evolutionary histories of Decacrema and their Pachystemon hosts have been contemporaneous since the mid-Miocene. The evolution of myrmecophytism enabled Macaranga to radiate into enemy-free space, while the ants' diversification has been shaped by stem traits, host specialization, and geographic factors. We discuss the possibility that the ancient and exclusive association between Decacrema and Macaranga was facilitated by an impoverished diversity of myrmecophytes and phytoecious (obligately plant inhabiting) ants in the region.

%B Evolution %V 58 %P 554-570 %8 Mar %@ 0014-3820 %G English %M WOS:000220704800010 %! EvolutionEvolution %0 Journal Article %J Nature %D 2004 %T The evolution of alternative parasitic life histories in large blue butterflies %A Als, T. D. %A Vila, R. %A Kandul, N. P. %A Nash, D. R. %A Yen, S. H. %A Hsu, Y. F. %A Mignault, A. A. %A Boomsma, J. J. %A Pierce, N. E. %K adoption %K alcon %K ant-nests %K formicidae %K host-specificity %K lepidoptera %K lycaenidae %K maculinea butterflies %X

Large blue (Maculinea) butterflies are highly endangered throughout the Palaearctic region, and have been the focus of intense conservation research(1-3). In addition, their extraordinary parasitic lifestyles make them ideal for studies of life history evolution. Early instars consume flower buds of specific host plants, but later instars live in ant nests where they either devour the brood (predators), or are fed mouth-to-mouth by the adult ants (cuckoos). Here we present the phylogeny for the group, which shows that it is a monophyletic clade nested within Phengaris, a rare Oriental genus whose species have similar life histories(4,5). Cuckoo species are likely to have evolved from predatory ancestors. As early as five million years ago, two Maculinea clades diverged, leading to the different parasitic strategies seen in the genus today. Contrary to current belief, the two recognized cuckoo species show little genetic divergence and are probably a single ecologically differentiated species(6-10). On the other hand, some of the predatory morphospecies exhibit considerable genetic divergence and may contain cryptic species. These findings have important implications for conservation and reintroduction efforts.

%B Nature %V 432 %P 386-390 %8 Nov 18 %@ 0028-0836 %G English %M WOS:000225161400054 %! NatureNature %0 Journal Article %J Systematic Entomology %D 2004 %T Molecular phylogeny of the Oriental butterfly genus Arhopala (Lycaenidae, Theclinae) inferred from mitochondrial and nuclear genes %A Megens, H. J. %A Van Nes, W. J. %A Van Moorsel, C. H. M. %A Pierce, N. E. %A de Jong, R. %K compositional bias %K cytochrome-oxidase-i %K DNA %K Evolution %K Heliconius butterflies %K inference %K lepidoptera %K nucleotide-sequence %K nymphalidae %K wingless gene %X

We present a phylogeny for a selection of species of the butterfly genus Arhopala Boisduval, 1832 based on molecular characters. We sequenced 1778 bases of the mitochondrial genes Cytochrome Oxidase 1 and 2 including tRNA(Leu), and a 393-bp fragment of the nuclear wingless gene for a total of 42 specimens of 33 species, representing all major species groups. Analyses of mtDNA and wingless genes show congruent phylogenetic signal. The phylogeny presented here confirms the monophyly of the centaurus, eumolphus, camdeo and epimuta groups and the amphimuta subgroup. It confirms close relationships between species within the agelastus group, that together with the amphimuta subgroup, centaurus and camdeo groups form a monophyletic group. However, incongruencies with previous taxonomic studies also occur; the amphimuta and silhetensis groups are not monophyletic, as is the genus Arhopala itself. One enigmatic species, A. kinabala, was evaluated further for topology and the support for basal placement of this species is due mainly to the wingless gene. However, in the Parsimony analysis, and subsequent Maximum Likelihood evaluations, certain nodes could not be resolved due to insufficient support. The mtDNA shows extreme AT bias with compositional heterogeneity at 3rd codon positions, which may result in saturation. By contrast, the wingless gene does not show compositional bias, suggesting that poor support is not due solely to saturation. The evaluation of morphological characters used in previous studies on Arhopala systematics on the molecular tree indicates that the macular pattern and the absence of tails at the hind wings show extensive homoplasy. A significant phylogenetic signal (as indicated by T-PTP tests) is present in several of these morphological characters, which are nevertheless of limited use in phylogenetic studies due to their labile nature.

%B Systematic Entomology %V 29 %P 115-131 %8 Jan %@ 0307-6970 %G English %M WOS:000188849100007 %! Syst EntomolSyst Entomol %0 Journal Article %J Systematic Biology %D 2004 %T Phylogeny of Agrodiaetus Hubner 1822 (Lepidoptera : Lycaenidae) inferred from mtDNA sequences of COI and COII and nuclear sequences of EF1-alpha: Karyotype diversification and species radiation %A Kandul, N. P. %A Lukhtanov, V. A. %A Dantchenko, A. V. %A Coleman, J. W. S. %A Sekercioglu, C. H. %A Haig, D. %A Pierce, N. E. %K agrodiaetus %K bayesian-inference %K chromosomal features %K cytochrome oxidase i %K cytochrome oxidase ii %K drosophila-melanogaster %K elongation factor 1 alpha %K elongation factor-1-alpha %K evolutionary genetics %K karyotype diversification %K length difference test %K maximum-likelihood %K metaphase-i bivalents %K mitochondrial-DNA %K Phylogeny %K polyommatus %K Speciation %K transposable element %X

Butterflies in the large Palearctic genus Agrodiaetus ( Lepidoptera: Lycaenidae) are extremely uniform and exhibit few distinguishing morphological characters. However, these insects are distinctive in one respect: as a group they possess among the greatest interspecific karyotype diversity in the animal kingdom, with chromosome numbers (n) ranging from 10 to 125. The monophyly of Agrodiaetus and its systematic position relative to other groups within the section Polyommatus have been controversial. Characters from the mitochondrial genes for cytochrome oxidases I and II and from the nuclear gene for elongation factor 1alpha were used to reconstruct the phylogeny of Agrodiaetus using maximum parsimony and Bayesian phylogenetic methods. Ninety-one individuals, encompassing most of the taxonomic diversity of Agrodiaetus, and representatives of 14 related genera were included in this analysis. Our data indicate that Agrodiaetus is monophyletic. Representatives of the genus Polyommatus ( sensu stricto) are the closest relatives. The sequences of the Agrodiaetus taxa in this analysis are tentatively arranged into 12 clades, only 1 of which corresponds to a species group traditionally recognized in Agrodiaetus. Heterogeneous substitution rates across a recovered topology were homogenized with a nonparametric rate-smoothing algorithm before the application of a molecular clock. Two published estimates of substitution rates dated the origin of Agrodiaetus between 2.51 and 3.85 million years ago. During this time, there was heterogeneity in the rate and direction of karyotype evolution among lineages within the genus. Karyotype instability has evolved independently three times in the section Polyommatus, within the lineages Agrodiaetus, Lysandra, and Plebicula. Rapid karyotype diversification may have played a significant role in the radiation of the genus Agrodiaetus.

%B Systematic Biology %V 53 %P 278-298 %8 Apr %@ 1063-5157 %G English %M WOS:000220800500006 %! Syst BiolSyst Biol %0 Journal Article %J Molecular Phylogenetics and Evolution %D 2004 %T Tempo of speciation in a butterfly genus from the Southeast Asian tropics, inferred from mitochondrial and nuclear DNA sequence data %A Megens, H. J. %A Van Moorsel, C. H. M. %A Piel, W. H. %A Pierce, N. E. %A de Jong, R. %K arhopala %K canary-islands %K cytochrome-oxidase-i %K diversification %K Evolution %K hard polytomy %K Heliconius butterflies %K molecular clock %K molecular phylogenetics %K null model %K phylogenetic-relationships %K rapid radiation %K substitution rates %K tropical diversity %X

Molecular systematics is frequently beset with phylogenetic results that are not fully resolved. Researchers either state that the absence of resolution is due to character conflict, explosive speciation, or some combination of the two, but seldom do they carefully examine their data to distinguish between these causes. In this study, we exhaustively analyze a set of nuclear and mitochondrial nucleotide data for the Asian tropical butterfly genus Arhopala so as to highlight the causes of polytomies in the phylogenetic trees, and, as a result, to infer important biological events in the history of this genus. We began by using non-parametric statistical methods to determine whether the ambiguously resolved regions in these trees represent hard or soft polytomies. In addition we determined how this correlated to number of inferred changes on branches, using parametric maximum likelihood estimations. Based on congruent patterns in both mitochondrial and nuclear DNA sequences, we concluded that at two stages in the history of Arhopala there have been accelerated instances of speciation. One event, at the base of the phylogeny, generated many of the groups and subgroups currently recognized in this genus, while a later event generated another major clade consisting of both Oriental and Papuan species groups. Based on comparisons of closely related taxa, the ratio of instantaneous rate of evolution between mitochondrial and nuclear DNA evolution is established at approximately 3:1. The earliest radiation is dated between 7 and 11 Ma by a molecular clock analysis, setting the events generating much of the diversity of Arhopala at well before the Pleistocene. Periodical flooding of the Sunda plateau during interglacial periods was, therefore, not responsible for generating the major divisions in the genus Arhopala. Instead, we hypothesize that large-scale climatic changes taking place in the Miocene have induced the early acceleration in speciation. (C) 2003 Elsevier Inc. All rights reserved.

%B Molecular Phylogenetics and Evolution %V 31 %P 1181-1196 %8 Jun %@ 1055-7903 %G English %M WOS:000221443400029 %! Mol Phylogenet EvolMol Phylogenet Evol %0 Book Section %B Butterflies as Model Systems %D 2003 %T Phylogenetic relationships of the Riodinidae: Implications for the evolution of ant association. %A Campbell, DL %A Pierce, N. E. %B Butterflies as Model Systems %I Chicago University Press %P 395-408 %G eng %0 Journal Article %J Annual Review of Entomology %D 2002 %T The ecology and evolution of ant association in the Lycaenidae (Lepidoptera) %A Pierce, N. E. %A Braby, M. F. %A Heath, A. %A Lohman, D. J. %A Mathew, J. %A Rand, D. B. %A Travassos, M. A. %K attendant ants %K biogeography %K butterfly %K butterfly maculinea-rebeli %K caterpillars lepidoptera %K host-specificity %K jalmenus-evagoras %K larvae lepidoptera %K myrmecophilous relationships %K myrmecophily %K parasitism %K plant-selection %K polymorphic growth %K social parasite %K symbiosis %X

The estimated 6000 species of Lycaenidae account for about one third of all Papilionoidea. The majority of lycaenids have associations with ants that can be facultative or obligate and range from mutualism to parasitism. Lycaenid larvae and pupae employ complex chemical and acoustical signals to manipulate ants. Cost/benefit analyses have demonstrated multiple trade-offs involved in myrmecophily. Both demographic and phylogenetic evidence indicate that ant association has shaped the evolution of obligately associated groups. Parasitism typically arises from mutualism with ants, arid entomophagous species are disproportionately common in the Lycaenidae compared with other Lepidoptera. Obligate associations are more common in the Southern Hemisphere, in part because highly ant-associated lineages make up a larger proportion of the fauna in these, regions. Further research on phylogeny and natural history, particularly of the Neotropical fauna, will be necessary to understand the rote ant association has played in the evolution of the Lycaenidae.

%B Annual Review of Entomology %V 47 %P 733-771 %@ 0066-4170 %G English %M WOS:000173421900024 %! Annu Rev EntomolAnnu Rev Entomol %0 Journal Article %J Journal of Evolutionary Biology %D 2002 %T Oviposition tests of ant preference in a myrmecophilous butterfly %A Fraser, A. M. %A Tregenza, T. %A Wedell, N. %A Elgar, M. A. %A Pierce, N. E. %K Associations %K Behavior %K diversification %K Evolution %K formicidae %K genetic-variation %K geographical specialization %K host selection %K host shift %K jalmenus-evagoras %K lepidoptera %K local adaptation %K lycaenid butterflies %K lycaenidae %K myrmecophily %K oviposition preference %K plant-selection %K Speciation %K specificity %X

Butterflies in the family Lycaenidae that have obligate associations with ants frequently exhibit ant-dependent egg laying behaviour. In a series of field and laboratory choice tests, we assessed oviposition preference of the Australian lycaenid Jalmenus evagoras in response to different species and populations of ants. Females discriminated between attendant and nonattendant ant species, between attendant ant species, and to some extent, between populations of a single ant species. When preferences were found, ovipositing butterflies preferred their locally predominant attendant ant species and geographically proximate attendant ant populations. A reciprocal choice test using adults from a generation of butterflies reared in the absence of ants indicated a genetic component to oviposition preference. Individual females were flexible with respect to oviposition site choice, often ovipositing on more than one treatment during a trial. Preferences arose from a hierarchical ranking of ant treatments. These results are discussed in terms of local adaptation and its possible significance in the diversification of ant-associated lycaenids.

%B Journal of Evolutionary Biology %V 15 %P 861-870 %8 Sep %@ 1010-061X %G English %M WOS:000177589000019 %! J Evolution BiolJ Evolution Biol %0 Journal Article %J Plant Physiology %D 2002 %T Signals involved in Arabidopsis resistance to Trichoplusia ni caterpillars induced by virulent and avirulent strains of the phytopathogen Pseudomonas syringae %A Cui, J. P. %A Jander, G. %A Racki, L. R. %A Kim, P. D. %A Pierce, N. E. %A Ausubel, F. M. %K cell-death %K enhanced disease susceptibility %K gene-expression %K hypersensitive response %K jasmonic acid %K nucleotide-binding site %K plant defense responses %K salicylic-acid %K systemic acquired-resistance %K wound response %X

Plants have evolved different but interconnected strategies to defend themselves against herbivorous insects and microbial pathogens. We used an Arabidopsis/Pseudomonas syringae pathosystem to investigate the impact of pathogen-induced defense responses on cabbage looper (Trichoplusia ni) larval feeding. Arabidopsis mutants [npr1, pad4, eds5, and sid2(eds16)] or transgenic plants (nahG) that are more susceptible to microbial pathogens and are compromised in salicylic acid (SA)-dependent defense responses exhibited reduced levels of feeding by T. ni compared with wild-type plants. Consistent with these results, Arabidopsis mutants that are more resistant to microbial pathogens and have elevated levels of SA (cpr1 and cpr6) exhibited enhanced levels of T. ni feeding. These experiments suggested an inverse relationship between an active SA defense pathway and insect feeding. In contrast to these results, there was increased resistance to T. ni in wild-type Arabidopsis ecotype Columbia plants that were infected with P. syringae pv. maculicola strain ES4326 (Psm ES4326) expressing the avirulence genes avrRpt2 or avrB, which elicit a hypersensitive response, high levels of SA accumulation, and systemic acquired resistance to bacterial infection. Similar results were obtained with other ecotypes, including Landsberg erecta, Cape Verdi Islands, and Shakdara. When infected with Psm ES4326(avrRpt2) or Psm ES4326(avrB), nahG transgenic and npr1 mutant plants (which are more susceptible to virulent and avirulent P. syringae strains) failed to show the increased insect resistance exhibited by wild-type plants. It was surprising that wild-type plants, as well as nahG and npr1 plants, infected with Psm ES4326 not expressing avrRpt2 or avrB, which elicits disease, became more susceptible to T. ni. Our results suggest two potentially novel systemic signaling pathways: a systemic response elicited by HR that leads to enhanced T. ni resistance and overrides the SA-mediated increase in T. ni susceptibility, and a SA-independent systemic response induced by virulent pathogens that leads to enhanced susceptibility to T. ni.

%B Plant Physiology %V 129 %P 551-564 %8 Jun %@ 0032-0889 %G English %M WOS:000176363800015 %! Plant PhysiolPlant Physiol %0 Journal Article %J Annual Review of Entomology %D 2002 %T The ecology and evolution of ant association in the Lycaenidae (Lepidoptera) %A Pierce, N. E. %A Braby, M. F. %A Heath, A. %A Lohman, D. J. %A Mathew, J. %A Rand, D. B. %A Travassos, M. A. %K attendant ants %K biogeography %K butterfly %K butterfly maculinea-rebeli %K caterpillars lepidoptera %K host-specificity %K jalmenus-evagoras %K larvae lepidoptera %K myrmecophilous relationships %K myrmecophily %K parasitism %K plant-selection %K polymorphic growth %K social parasite %K symbiosis %X

The estimated 6000 species of Lycaenidae account for about one third of all Papilionoidea. The majority of lycaenids have associations with ants that can be facultative or obligate and range from mutualism to parasitism. Lycaenid larvae and pupae employ complex chemical and acoustical signals to manipulate ants. Cost/benefit analyses have demonstrated multiple trade-offs involved in myrmecophily. Both demographic and phylogenetic evidence indicate that ant association has shaped the evolution of obligately associated groups. Parasitism typically arises from mutualism with ants, arid entomophagous species are disproportionately common in the Lycaenidae compared with other Lepidoptera. Obligate associations are more common in the Southern Hemisphere, in part because highly ant-associated lineages make up a larger proportion of the fauna in these, regions. Further research on phylogeny and natural history, particularly of the Neotropical fauna, will be necessary to understand the rote ant association has played in the evolution of the Lycaenidae.

%B Annual Review of Entomology %V 47 %P 733-771 %@ 0066-4170 %G English %M WOS:000173421900024 %! Annu Rev EntomolAnnu Rev Entomol %0 Journal Article %J Journal of Evolutionary Biology %D 2002 %T Oviposition tests of ant preference in a myrmecophilous butterfly %A Fraser, A. M. %A Tregenza, T. %A Wedell, N. %A Elgar, M. A. %A Pierce, N. E. %K Associations %K Behavior %K diversification %K Evolution %K formicidae %K genetic-variation %K geographical specialization %K host selection %K host shift %K jalmenus-evagoras %K lepidoptera %K local adaptation %K lycaenid butterflies %K lycaenidae %K myrmecophily %K oviposition preference %K plant-selection %K Speciation %K specificity %X

Butterflies in the family Lycaenidae that have obligate associations with ants frequently exhibit ant-dependent egg laying behaviour. In a series of field and laboratory choice tests, we assessed oviposition preference of the Australian lycaenid Jalmenus evagoras in response to different species and populations of ants. Females discriminated between attendant and nonattendant ant species, between attendant ant species, and to some extent, between populations of a single ant species. When preferences were found, ovipositing butterflies preferred their locally predominant attendant ant species and geographically proximate attendant ant populations. A reciprocal choice test using adults from a generation of butterflies reared in the absence of ants indicated a genetic component to oviposition preference. Individual females were flexible with respect to oviposition site choice, often ovipositing on more than one treatment during a trial. Preferences arose from a hierarchical ranking of ant treatments. These results are discussed in terms of local adaptation and its possible significance in the diversification of ant-associated lycaenids.

%B Journal of Evolutionary Biology %V 15 %P 861-870 %8 Sep %@ 1010-061X %G English %M WOS:000177589000019 %! J Evolution BiolJ Evolution Biol %0 Journal Article %J Plant Physiology %D 2002 %T Signals involved in Arabidopsis resistance to Trichoplusia ni caterpillars induced by virulent and avirulent strains of the phytopathogen Pseudomonas syringae %A Cui, J. P. %A Jander, G. %A Racki, L. R. %A Kim, P. D. %A Pierce, N. E. %A Ausubel, F. M. %K cell-death %K enhanced disease susceptibility %K gene-expression %K hypersensitive response %K jasmonic acid %K nucleotide-binding site %K plant defense responses %K salicylic-acid %K systemic acquired-resistance %K wound response %X

Plants have evolved different but interconnected strategies to defend themselves against herbivorous insects and microbial pathogens. We used an Arabidopsis/Pseudomonas syringae pathosystem to investigate the impact of pathogen-induced defense responses on cabbage looper (Trichoplusia ni) larval feeding. Arabidopsis mutants [npr1, pad4, eds5, and sid2(eds16)] or transgenic plants (nahG) that are more susceptible to microbial pathogens and are compromised in salicylic acid (SA)-dependent defense responses exhibited reduced levels of feeding by T. ni compared with wild-type plants. Consistent with these results, Arabidopsis mutants that are more resistant to microbial pathogens and have elevated levels of SA (cpr1 and cpr6) exhibited enhanced levels of T. ni feeding. These experiments suggested an inverse relationship between an active SA defense pathway and insect feeding. In contrast to these results, there was increased resistance to T. ni in wild-type Arabidopsis ecotype Columbia plants that were infected with P. syringae pv. maculicola strain ES4326 (Psm ES4326) expressing the avirulence genes avrRpt2 or avrB, which elicit a hypersensitive response, high levels of SA accumulation, and systemic acquired resistance to bacterial infection. Similar results were obtained with other ecotypes, including Landsberg erecta, Cape Verdi Islands, and Shakdara. When infected with Psm ES4326(avrRpt2) or Psm ES4326(avrB), nahG transgenic and npr1 mutant plants (which are more susceptible to virulent and avirulent P. syringae strains) failed to show the increased insect resistance exhibited by wild-type plants. It was surprising that wild-type plants, as well as nahG and npr1 plants, infected with Psm ES4326 not expressing avrRpt2 or avrB, which elicits disease, became more susceptible to T. ni. Our results suggest two potentially novel systemic signaling pathways: a systemic response elicited by HR that leads to enhanced T. ni resistance and overrides the SA-mediated increase in T. ni susceptibility, and a SA-independent systemic response induced by virulent pathogens that leads to enhanced susceptibility to T. ni.

%B Plant Physiology %V 129 %P 551-564 %8 Jun %@ 0032-0889 %G English %M WOS:000176363800015 %! Plant PhysiolPlant Physiol %0 Journal Article %J Animal Behaviour %D 2001 %T The cavity-dwelling ant Leptothorax curvispinosus uses nest geometry to discriminate between potential homes %A Pratt, S. C. %A Pierce, N. E. %K community %K construction %K division-of-labor %K emigration %K habitat selection %K limitation %K resilience %K site selection %K social-organization %K unifasciatus latr %X

Nest site selection is a frequent context for decision making in ants, but little is known of the criteria used to make a choice. We tested the nest site preferences of Leptothorax curvispinosus, both by measuring hollow acorn nests occupied in nature, and by inducing laboratory colonies to choose between artificial nests of different design. Three criteria were examined. (1) Entrance size: the ants preferred small entrance holes, presumably for their greater defensibility and crypsis. Natural nest entrances were small, and 52% of them were reduced still further by the addition of rims of soil and leaf litter. In choice tests, colonies selected nest entrances near the median size of rimmed natural holes, rejecting those near the larger end of the distribution of raw natural holes. (2) Cavity volume: acorn cavity volume was weakly correlated with the size of the occupying colony. In choice tests, colonies rejected cavities near the median size of natural nests, preferring instead larger cavities near the upper end of the natural size distribution. This may reflect active size matching of colonies to nests, because the colonies used in the choice test were bigger than those from the natural nest sample. Alternatively, all colonies may prefer big nests, but face limited availability of large cavities in nature. (3) Cavity shape: colonies preferred shapes roughly similar to that of an acorn interior, rejecting thin crevices in favour of compact, high-ceilinged cavities. (C) 2001 The Association for the Study of Animal Behaviour.

%B Animal Behaviour %V 62 %P 281-287 %8 Aug %@ 0003-3472 %G English %M WOS:000171088000012 %! Anim BehavAnim Behav %0 Journal Article %J Ecology %D 2001 %T An empirical model of species coexistence in a spatially structured environment %A Yu, D. W. %A Wilson, H. B. %A Pierce, N. E. %K allomerus %K ant-plant mutualism %K ant-plant symbiosis %K azteca %K Community structure %K competition %K competition-colonization trade-off %K cordia nodosa %K dispersal-fecundity trade-off %K Evolution %K habitat destruction and selection hypotheses %K lottery model %K metapopulation %K Parasite %K recruitment limitation %K seed size %K source-sink dynamics %K space %K storage effect %K trade-off %K variability %K yucca moths %X

Ecological theory has long supported the idea that species coexistence in a homogeneous habitat is promoted by spatial structure, but empirical evidence for this hypothesis has lagged behind theory. Here we describe a Neotropical ant-plant symbiosis that is ideally suited for testing spatial models of coexistence. Two genera of ants, Allomerus cf. demerarae and three species of Azteca are specialized to live on a single species of ant-plant, Cordia nodosa, in a Western Amazonian tropical rain forest. Empirically, using census data from widely separated localities, we show that the relative colonization abilities of the two ant genera are a function of plant density. A parameterized model shows that this pattern alone is sufficiently robust to explain coexistence in;the system. Census and experimental data suggest that Azteca queens are better long-distance flyers, but that Allomerus colonies are more fecund. Thus, Azteca can dominate in areas where host-plant densities are low land parent colony-sapling distances are long), and Allomerus can dominate in areas where host-plant densities are high. Existing spatial heterogeneity in host-plant densities therefore can allow regional coexistence, and intersite dispersal can produce local mixing. In conclusion, a dispersal-fecundity trade-off appears to allow the two genera to treat spatial heterogeneity in patch density as a niche axis. This study further suggests that a spatially structured approach is essential in understanding the persistence of some mutualisms in the presence of parasites.

%B Ecology %V 82 %P 1761-1771 %8 Jun %@ 0012-9658 %G English %M WOS:000169521200021 %! EcologyEcology %0 Journal Article %J Biological Journal of the Linnean Society %D 2001 %T Molecular evolution of a long wavelength-sensitive opsin in mimetic Heliconius butterflies (Lepidoptera : Nymphalidae) %A Hsu, R. %A Briscoe, A. D. %A Chang, B. S. W. %A Pierce, N. E. %K bovine rhodopsin %K butterfly %K identification %K invertebrate vision %K lepidopteran %K mimicry %K mitochondrial-DNA %K papilio-glaucus %K passion-vine butterflies %K Phylogeny %K protein-coupled receptor %K red %K schiff-base counterion %K visual pigment %K visual pigments %X

This study examines the pattern of opsin nucleotide and amino acid substitution among mimetic species 'rings' of Heliconius butterflies that are characterized by divergent wing colour patterns. A long wavelength opsin gene, OPS1, was sequenced from each of seven species of Heliconius and one species of Dryas (Lepidoptera: Nymphalidae). A parsimony analysis of OPS1 nucleotide and amino acid sequences resulted in a phylogeny that was consistent with that presented by Brewer & Egan in 1997, which was based on mitochondrial cytochrome oxidase I and II as well as nuclear wingless genes. Nodes in the OPS1 phylogeny were well supported by bootstrap analysis and decay indices. An analysis of specific sites within the gene indicates that the accumulation of amino acid substitutions has occurred independently of the morphological diversification of Heliconius wing colour patterns. Amino acid substitutions were examined with respect to their location within the opsin protein and their possible interactions with the chromophore and the G-protein. Of the 15 amino acid substitutions identified among the eight species, one nonconservative replacement (A226Q) was identified in a position that may be involved in binding with the G-protein. (C) 2001 The Linnean Society of London.

%B Biological Journal of the Linnean Society %V 72 %P 435-449 %8 Mar %@ 0024-4066 %G English %M WOS:000168187100008 %! Biol J Linn SocBiol J Linn Soc %0 Journal Article %J Molecular Phylogenetics and Evolution %D 2001 %T Phylogeny of Bicyclus (Lepidoptera : Nymphalidae) inferred from COI, COII, and EF-1 alpha gene sequences %A Monteiro, A. %A Pierce, N. E. %K anynana %K bicyclus %K butterfly wings %K coi %K coii %K Development %K ef-1 alpha %K elongation factor-1-alpha %K Evolution %K eyespot patterns %K eyespots %K lepidoptera %K likelihood %K maximum %K mitochondrial-DNA %K nucleotide-sequence %K nymphalidae %K parsimony %K Phenotypic plasticity %K Phylogeny %K satyrinae %K seasonal polyphenism %K synonymous substitution %K wing pattern %X

Despite the fact that Bicyclus anynana has become an important model species for wing-pattern developmental biology and studies of phenotypic plasticity, little is known of the evolutionary history of the genus Bicyclus and the position of B. anynana. Understanding the evolution of development as well as the evolution of plasticity can be attempted in this species-rich genus that displays a large range of wing patterns with variable degrees of phenotypic responses to the environment. A context to guide extrapolations from population genetic studies within B. anynana to those between closely related species has been long overdue. A phylogeny of 54 of the 80 known Bicyclus species is presented based on the combined 3000-bp sequences of two mitochondrial genes, cytochrome oxidase I and II, and the nuclear gene, elongation factor I alpha. A series of tree topologies, constructed either from the individual genes or from the combined data, using heuristic searches under a variety of weighting schemes were compared under the best maximum-likelihood models fitted for each gene separately. The most likely tree topology to have generated the three data sets was found to be a tree resulting from a combined MP analysis with equal weights. Most phylogenetic signal for the analysis comes from silent substitutions at the third position, and despite the faster rate of evolution and higher levels of homoplasy of the mitochondrial genes relative to the nuclear gene, the latter does not show substantially stronger support for basal clades. Finally, moving branches from the chosen tree topology to other positions on the tree so as to comply better with a previous morphological study did not significantly affect tree length. (C) 2001 Academic Press.

%B Molecular Phylogenetics and Evolution %V 18 %P 264-281 %8 Feb %@ 1055-7903 %G English %M WOS:000166821700009 %! Mol Phylogenet EvolMol Phylogenet Evol %0 Journal Article %J Plant Physiology %D 2001 %T The TASTY locus on chromosome 1 of Arabidopsis affects feeding of the insect herbivore Trichoplusia ni %A Jander, G. %A Cui, J. P. %A Nhan, B. %A Pierce, N. E. %A Ausubel, F. M. %K cabbage-looper %K defense %K deterrency %K lepidoptera %K populations %K quantitative genetic-variation %K recombinant inbred lines %K resistance %K thaliana %K trait loci %X

The generalist insect herbivore Trichoplusia ni (cabbage looper) readily consumes Arabidopsis and can complete its entire life cycle on this plant. Natural isolates (ecotypes) of Arabidopsis are not equally susceptible to T. ni feeding. While some are hardly touched by T. ni, others are eaten completely to the ground. Comparison of two commonly studied Arabidopsis ecotypes in choice experiments showed that Columbia is considerably more resistant than Landsberg erecta. In no-choice experiments, where larvae were confined on one or the other ecotype, weight gain was more rapid on Landsberg erects than on Columbia. Genetic mapping of this difference in insect susceptibility using recombinant inbred lines resulted in the discovery of the TASTY locus near 85 cM on chromosome 1 of Arabidopsis. The resistant allele of this locus is in the Columbia ecotype, and an F, hybrid has a sensitive phenotype that is similar to that of Landsberg erecta. The TASTY locus is distinct from known genetic differences between Columbia and Landsberg erecta that affect glucosinolate content, trichome density, disease resistance, and flowering time.

%B Plant Physiology %V 126 %P 890-898 %8 Jun %@ 0032-0889 %G English %M WOS:000169297000044 %! Plant PhysiolPlant Physiol %0 Book Section %B Model systems in behavioral ecology %D 2001 %T Peeling the onion: Symbioses between ants and blue butterflies %A Pierce, N. E. %B Model systems in behavioral ecology %I Princeton University Press %C Princeton %P 41-56 %G eng %0 Journal Article %J Oecologia %D 2001 %T Assessing the quality of different ant species as partners of a myrmecophilous butterfly %A Fraser, A. M. %A Axen, A. H. %A Pierce, N. E. %K association %K attendant ants %K benefits %K conditional mutualism %K Cost-Benefit Analysis %K jalmenus-evagoras %K larvae %K lepidoptera %K lycaenid butterflies %K lycaenid butterfly-ant interactions %K mutualistic interactions %K neotropical herb %K parasitism %K Pollination %K species-specific effects %X

We assessed the quality of different ant species as partners of the facultatively myrmecophilous lycaenid butterfly Glaucopsyche lygdamus. We compared disappearance and parasitism rates of G. lygdamus larvae in the field, and development of non-feeding pre-pupae in the laboratory, when individuals were untended or tended by one of four ant species. Formica podzolica was the only ant species to provide a clear benefit to G. lygdamus, in the form of reduced larval parasitism relative to untended larvae. F. 'neogagates' (F. neogagates + F. lasioides) and Tapinoma sessile were essentially neutral partners, providing no significant cost or benefit for any of the parameters measured. Relative to untended individuals, association with F. obscuripes significantly increased larval disappearance and significantly decreased pupal mass. Thus, F. obscuripes may act as a parasite of the general association between G. lygdamus and ants under certain conditions. Ant species also differed in their persistence as tenders of G. lygdamus larvae once an interaction was established. Over the lifetime of a larva, F. podzolica and F. obscuripes usually remained as the attendant ant species on plants over consecutive census dates, while F. 'neogagates' and T. sessile were frequently replaced, most commonly by F. obscuripes. It remains to be determined if disappearance and developmental outcomes reported here reflect true fitness costs (i.e. reduced survivorship and lower reproductive success) for G. lygdamus. The potential and limitations for specialization in association between G. lygdamus and high quality ant partners are discussed.

%B Oecologia %V 129 %P 452-460 %8 Nov %@ 0029-8549 %G English %M WOS:000172418100016 %! OecologiaOecologia %0 Journal Article %J Animal Behaviour %D 2001 %T The cavity-dwelling ant Leptothorax curvispinosus uses nest geometry to discriminate between potential homes %A Pratt, S. C. %A Pierce, N. E. %K community %K construction %K division-of-labor %K emigration %K habitat selection %K limitation %K resilience %K site selection %K social-organization %K unifasciatus latr %X

Nest site selection is a frequent context for decision making in ants, but little is known of the criteria used to make a choice. We tested the nest site preferences of Leptothorax curvispinosus, both by measuring hollow acorn nests occupied in nature, and by inducing laboratory colonies to choose between artificial nests of different design. Three criteria were examined. (1) Entrance size: the ants preferred small entrance holes, presumably for their greater defensibility and crypsis. Natural nest entrances were small, and 52% of them were reduced still further by the addition of rims of soil and leaf litter. In choice tests, colonies selected nest entrances near the median size of rimmed natural holes, rejecting those near the larger end of the distribution of raw natural holes. (2) Cavity volume: acorn cavity volume was weakly correlated with the size of the occupying colony. In choice tests, colonies rejected cavities near the median size of natural nests, preferring instead larger cavities near the upper end of the natural size distribution. This may reflect active size matching of colonies to nests, because the colonies used in the choice test were bigger than those from the natural nest sample. Alternatively, all colonies may prefer big nests, but face limited availability of large cavities in nature. (3) Cavity shape: colonies preferred shapes roughly similar to that of an acorn interior, rejecting thin crevices in favour of compact, high-ceilinged cavities. (C) 2001 The Association for the Study of Animal Behaviour.

%B Animal Behaviour %V 62 %P 281-287 %8 Aug %@ 0003-3472 %G English %M WOS:000171088000012 %! Anim BehavAnim Behav %0 Journal Article %J Ecology %D 2001 %T An empirical model of species coexistence in a spatially structured environment %A Yu, D. W. %A Wilson, H. B. %A Pierce, N. E. %K allomerus %K ant-plant mutualism %K ant-plant symbiosis %K azteca %K Community structure %K competition %K competition-colonization trade-off %K cordia nodosa %K dispersal-fecundity trade-off %K Evolution %K habitat destruction and selection hypotheses %K lottery model %K metapopulation %K Parasite %K recruitment limitation %K seed size %K source-sink dynamics %K space %K storage effect %K trade-off %K variability %K yucca moths %X

Ecological theory has long supported the idea that species coexistence in a homogeneous habitat is promoted by spatial structure, but empirical evidence for this hypothesis has lagged behind theory. Here we describe a Neotropical ant-plant symbiosis that is ideally suited for testing spatial models of coexistence. Two genera of ants, Allomerus cf. demerarae and three species of Azteca are specialized to live on a single species of ant-plant, Cordia nodosa, in a Western Amazonian tropical rain forest. Empirically, using census data from widely separated localities, we show that the relative colonization abilities of the two ant genera are a function of plant density. A parameterized model shows that this pattern alone is sufficiently robust to explain coexistence in;the system. Census and experimental data suggest that Azteca queens are better long-distance flyers, but that Allomerus colonies are more fecund. Thus, Azteca can dominate in areas where host-plant densities are low land parent colony-sapling distances are long), and Allomerus can dominate in areas where host-plant densities are high. Existing spatial heterogeneity in host-plant densities therefore can allow regional coexistence, and intersite dispersal can produce local mixing. In conclusion, a dispersal-fecundity trade-off appears to allow the two genera to treat spatial heterogeneity in patch density as a niche axis. This study further suggests that a spatially structured approach is essential in understanding the persistence of some mutualisms in the presence of parasites.

%B Ecology %V 82 %P 1761-1771 %8 Jun %@ 0012-9658 %G English %M WOS:000169521200021 %! EcologyEcology %0 Journal Article %J Biological Journal of the Linnean Society %D 2001 %T Molecular evolution of a long wavelength-sensitive opsin in mimetic Heliconius butterflies (Lepidoptera : Nymphalidae) %A Hsu, R. %A Briscoe, A. D. %A Chang, B. S. W. %A Pierce, N. E. %K bovine rhodopsin %K butterfly %K identification %K invertebrate vision %K lepidopteran %K mimicry %K mitochondrial-DNA %K papilio-glaucus %K passion-vine butterflies %K Phylogeny %K protein-coupled receptor %K red %K schiff-base counterion %K visual pigment %K visual pigments %X

This study examines the pattern of opsin nucleotide and amino acid substitution among mimetic species 'rings' of Heliconius butterflies that are characterized by divergent wing colour patterns. A long wavelength opsin gene, OPS1, was sequenced from each of seven species of Heliconius and one species of Dryas (Lepidoptera: Nymphalidae). A parsimony analysis of OPS1 nucleotide and amino acid sequences resulted in a phylogeny that was consistent with that presented by Brewer & Egan in 1997, which was based on mitochondrial cytochrome oxidase I and II as well as nuclear wingless genes. Nodes in the OPS1 phylogeny were well supported by bootstrap analysis and decay indices. An analysis of specific sites within the gene indicates that the accumulation of amino acid substitutions has occurred independently of the morphological diversification of Heliconius wing colour patterns. Amino acid substitutions were examined with respect to their location within the opsin protein and their possible interactions with the chromophore and the G-protein. Of the 15 amino acid substitutions identified among the eight species, one nonconservative replacement (A226Q) was identified in a position that may be involved in binding with the G-protein. (C) 2001 The Linnean Society of London.

%B Biological Journal of the Linnean Society %V 72 %P 435-449 %8 Mar %@ 0024-4066 %G English %M WOS:000168187100008 %! Biol J Linn SocBiol J Linn Soc %0 Journal Article %J Molecular Phylogenetics and Evolution %D 2001 %T Phylogeny of Bicyclus (Lepidoptera : Nymphalidae) inferred from COI, COII, and EF-1 alpha gene sequences %A Monteiro, A. %A Pierce, N. E. %K anynana %K bicyclus %K butterfly wings %K coi %K coii %K Development %K ef-1 alpha %K elongation factor-1-alpha %K Evolution %K eyespot patterns %K eyespots %K lepidoptera %K likelihood %K maximum %K mitochondrial-DNA %K nucleotide-sequence %K nymphalidae %K parsimony %K Phenotypic plasticity %K Phylogeny %K satyrinae %K seasonal polyphenism %K synonymous substitution %K wing pattern %X

Despite the fact that Bicyclus anynana has become an important model species for wing-pattern developmental biology and studies of phenotypic plasticity, little is known of the evolutionary history of the genus Bicyclus and the position of B. anynana. Understanding the evolution of development as well as the evolution of plasticity can be attempted in this species-rich genus that displays a large range of wing patterns with variable degrees of phenotypic responses to the environment. A context to guide extrapolations from population genetic studies within B. anynana to those between closely related species has been long overdue. A phylogeny of 54 of the 80 known Bicyclus species is presented based on the combined 3000-bp sequences of two mitochondrial genes, cytochrome oxidase I and II, and the nuclear gene, elongation factor I alpha. A series of tree topologies, constructed either from the individual genes or from the combined data, using heuristic searches under a variety of weighting schemes were compared under the best maximum-likelihood models fitted for each gene separately. The most likely tree topology to have generated the three data sets was found to be a tree resulting from a combined MP analysis with equal weights. Most phylogenetic signal for the analysis comes from silent substitutions at the third position, and despite the faster rate of evolution and higher levels of homoplasy of the mitochondrial genes relative to the nuclear gene, the latter does not show substantially stronger support for basal clades. Finally, moving branches from the chosen tree topology to other positions on the tree so as to comply better with a previous morphological study did not significantly affect tree length. (C) 2001 Academic Press.

%B Molecular Phylogenetics and Evolution %V 18 %P 264-281 %8 Feb %@ 1055-7903 %G English %M WOS:000166821700009 %! Mol Phylogenet EvolMol Phylogenet Evol %0 Journal Article %J Plant Physiology %D 2001 %T The TASTY locus on chromosome 1 of Arabidopsis affects feeding of the insect herbivore Trichoplusia ni %A Jander, G. %A Cui, J. P. %A Nhan, B. %A Pierce, N. E. %A Ausubel, F. M. %K cabbage-looper %K defense %K deterrency %K lepidoptera %K populations %K quantitative genetic-variation %K recombinant inbred lines %K resistance %K thaliana %K trait loci %X

The generalist insect herbivore Trichoplusia ni (cabbage looper) readily consumes Arabidopsis and can complete its entire life cycle on this plant. Natural isolates (ecotypes) of Arabidopsis are not equally susceptible to T. ni feeding. While some are hardly touched by T. ni, others are eaten completely to the ground. Comparison of two commonly studied Arabidopsis ecotypes in choice experiments showed that Columbia is considerably more resistant than Landsberg erecta. In no-choice experiments, where larvae were confined on one or the other ecotype, weight gain was more rapid on Landsberg erects than on Columbia. Genetic mapping of this difference in insect susceptibility using recombinant inbred lines resulted in the discovery of the TASTY locus near 85 cM on chromosome 1 of Arabidopsis. The resistant allele of this locus is in the Columbia ecotype, and an F, hybrid has a sensitive phenotype that is similar to that of Landsberg erecta. The TASTY locus is distinct from known genetic differences between Columbia and Landsberg erecta that affect glucosinolate content, trichome density, disease resistance, and flowering time.

%B Plant Physiology %V 126 %P 890-898 %8 Jun %@ 0032-0889 %G English %M WOS:000169297000044 %! Plant PhysiolPlant Physiol %0 Journal Article %J Animal Behaviour %D 2000 %T Acoustics, context and function of vibrational signalling in a lycaenid butterfly-ant mutualism %A Travassos, M. A. %A Pierce, N. E. %K benefits %K communication %K formicidae %K hymenoptera %K jalmenus-evagoras %K larvae %K lepidoptera %K myrmecophilous organs %K Recruitment %X

Juveniles of the Australian common imperial blue butterfly, Jalmenus evagoras, produce substrate-borne vibrational signals in the form of two kinds of pupal calls and three larval calls. Pupae stridulate in the presence of conspecific larvae, when attended by an ant guard, and as a reaction against perturbation. Using pupal pairs in which one member was experimentally muted, pupal calls were shown to be important in ant attraction and the maintenance of an ant guard. A pupa may use-calls to regulate levels of its attendant ants and to signal its potential value in these mutualistic interactions. Therefore substrate-borne vibrations play a significant role in the communication between J. evagoras and its attendant ants and pupal calls appear to be more than just signals acting as a predator deterrent. Similarly, caterpillars make more sound when attended by Iridomyrmex anceps, suggesting that larval calls may be important in mediating ant symbioses. One larval call has the same mean dominant frequency, pulse rate, bandwidth and pulse length as the primary signal of a pupa, suggesting a similarity in function. (C) 2000 The Association for the Study of Animal Behaviour.

%B Animal Behaviour %V 60 %P 13-26 %8 Jul %@ 0003-3472 %G English %M WOS:000088673400002 %! Anim BehavAnim Behav %0 Journal Article %J Behavioral Ecology and Sociobiology %D 2000 %T Effects of mating history on ejaculate size, fecundity, longevity, and copulation duration in the ant-tended lycaenid butterfly, Jalmenus evagoras %A Hughes, L. %A Chang, B. S. W. %A Wagner, D. %A Pierce, N. E. %K copulation duration %K ejaculate size %K explosive mating system %K female reproduction %K lepidoptera %K male nutrient investment %K mate-locating behavior %K mating history %K monandry polyandry %K multiple matings %K napi l %K papilio-machaon l %K paternal investment %K pieris-rapae lepidoptera %K reproductive output %X

The mating system of the Australian lycaenid butterfly, Jalmenus evagoras, is highly unusual compared to most other Lepidoptera. Characteristics of this system, which has been termed an 'explosive mating strategy,' include the formation of an intensely competitive mating aggregation of males, a highly male biased operational sex ratio, a lack of discrimination and mate choice by both sexes, a high variance in male mating success, and female monogamy. In this study, we tested the hypothesis that multiple mating by males imposes physiological costs resulting in smaller spermatophores, and that this results in a fitness cost to females. We found that male J. evagoras transferred only 2.2% of their eclosion weight during their first mating, consistent with the hypothesis that males of monandrous species produce a relatively small investment. The wet weight of the ejaculate declined by an average of 27% at the second mating and the dry weight by 29%, and an intermating interval of 5-9 days was needed for the ejaculate to return to the size at the first mating, regardless of male size or age. Wet ejaculate mass increased proportionally with male size, though dry mass was proportionally larger in smaller males. Ejaculate mass tended to increase with male age at both first and second matings. Female characteristics, in general, did not affect ejaculate mass, although the wet weight of the ejaculate was positively associated with female weight at the second mating. Copulation duration increased from 2.4 h to approximately 3 h at the second mating, and to over 4 h at the third and fourth matings. Fecundity was positively correlated with female size but not with mating history, copulation duration, or any other characteristics measured for either males or females. Female longevity declined significantly as the number of times the male partner had previously mated increased. We conclude that despite the small male investment in ejaculate, the costs of multiple mating may nonetheless be significant, as indicated by the reduction in ejaculate mass, an increase in copulation duration, and reduction in female lifespan with increasing mating number.

%B Behavioral Ecology and Sociobiology %V 47 %P 119-128 %8 Feb %@ 0340-5443 %G English %M WOS:000085378400002 %! Behav Ecol SociobiolBehav Ecol Sociobiol %0 Journal Article %J Molecular Phylogeny and Evolution %D 2000 %T Phylogeny and life history evolution of the genus Chrysoritis within the Aphnaeini (Lepidoptera: Lycaenidae), inferred from mitochondrial cytochrome oxidase I sequences. %A Rand, DB %A A Heath %A T Suderman %A Pierce, N. E. %X

Phylogenetic relationships among 26 South African species in the tribe Aphnaeini (Lepidoptera: Lycaenidae) were inferred from DNA characters of the mitochondrial gene cytochrome oxidase I (COI), using maximum-parsimony methods. The resulting phylogenetic estimate supports the systematic hypothesis made by Heath (1997, Metamorphosis, supplement 2), based on morphological characters, that at least three preexisting genera (Chrysoritis, Poecilmitis, and Oxychaeta) should be collapsed into the single monophyletic genus Chrysoritis. Two of the species groups described by Heath within Chrysoritis are also monophyletic, while one is paraphyletic and thus unsupported by the molecular data. Strong node support and skewed transition/transversion ratios suggest that two Chrysoritis clades contain synonymous species. Aphytophagy appears as a derived feeding strategy. Evolutionary patterns of ant association indicate lability at the level of ant genus, while association with different ant subfamilies may have played an ancestral and chemically mediated role in the diversification of South African aphnaeines. ©

%B Molecular Phylogeny and Evolution %V 17 %P 85-96 %G eng %0 Journal Article %J Nature Medicine %D 2000 %T W.D. Hamilton, 1936-2000 - Obituary %A Moran, N. %A Pierce, N. %A Seger, J. %B Nature Medicine %V 6 %P 367-367 %8 Apr %@ 1078-8956 %G English %M WOS:000165474100016 %! Nat MedNat Med %0 Journal Article %J Science %D 2000 %T William Hamilton (1936-2000) %A Haig, D. %A Pierce, N. E. %A Wilson, E. O. %B Science %V 287 %P 2438-2438 %8 Mar 31 %@ 0036-8075 %G English %M WOS:000086202200034 %! ScienceScience %0 Journal Article %J Molecular Phylogeny and Evolution %D 2000 %T Phylogeny and life history evolution of the genus Chrysoritis within the Aphnaeini (Lepidoptera: Lycaenidae), inferred from mitochondrial cytochrome oxidase I sequences. %A Rand, DB %A A Heath %A T Suderman %A Pierce, N. E. %X

Phylogenetic relationships among 26 South African species in the tribe Aphnaeini (Lepidoptera: Lycaenidae) were inferred from DNA characters of the mitochondrial gene cytochrome oxidase I (COI), using maximum-parsimony methods. The resulting phylogenetic estimate supports the systematic hypothesis made by Heath (1997, Metamorphosis, supplement 2), based on morphological characters, that at least three preexisting genera (Chrysoritis, Poecilmitis, and Oxychaeta) should be collapsed into the single monophyletic genus Chrysoritis. Two of the species groups described by Heath within Chrysoritis are also monophyletic, while one is paraphyletic and thus unsupported by the molecular data. Strong node support and skewed transition/transversion ratios suggest that two Chrysoritis clades contain synonymous species. Aphytophagy appears as a derived feeding strategy. Evolutionary patterns of ant association indicate lability at the level of ant genus, while association with different ant subfamilies may have played an ancestral and chemically mediated role in the diversification of South African aphnaeines. ©

%B Molecular Phylogeny and Evolution %V 17 %P 85-96 %G eng %0 Journal Article %J Animal Behaviour %D 2000 %T Acoustics, context and function of vibrational signalling in a lycaenid butterfly-ant mutualism %A Travassos, M. A. %A Pierce, N. E. %K benefits %K communication %K formicidae %K hymenoptera %K jalmenus-evagoras %K larvae %K lepidoptera %K myrmecophilous organs %K Recruitment %X

Juveniles of the Australian common imperial blue butterfly, Jalmenus evagoras, produce substrate-borne vibrational signals in the form of two kinds of pupal calls and three larval calls. Pupae stridulate in the presence of conspecific larvae, when attended by an ant guard, and as a reaction against perturbation. Using pupal pairs in which one member was experimentally muted, pupal calls were shown to be important in ant attraction and the maintenance of an ant guard. A pupa may use-calls to regulate levels of its attendant ants and to signal its potential value in these mutualistic interactions. Therefore substrate-borne vibrations play a significant role in the communication between J. evagoras and its attendant ants and pupal calls appear to be more than just signals acting as a predator deterrent. Similarly, caterpillars make more sound when attended by Iridomyrmex anceps, suggesting that larval calls may be important in mediating ant symbioses. One larval call has the same mean dominant frequency, pulse rate, bandwidth and pulse length as the primary signal of a pupa, suggesting a similarity in function. (C) 2000 The Association for the Study of Animal Behaviour.

%B Animal Behaviour %V 60 %P 13-26 %8 Jul %@ 0003-3472 %G English %M WOS:000088673400002 %! Anim BehavAnim Behav %0 Journal Article %J Behavioral Ecology and Sociobiology %D 2000 %T Effects of mating history on ejaculate size, fecundity, longevity, and copulation duration in the ant-tended lycaenid butterfly, Jalmenus evagoras %A Hughes, L. %A Chang, B. S. W. %A Wagner, D. %A Pierce, N. E. %K copulation duration %K ejaculate size %K explosive mating system %K female reproduction %K lepidoptera %K male nutrient investment %K mate-locating behavior %K mating history %K monandry polyandry %K multiple matings %K napi l %K papilio-machaon l %K paternal investment %K pieris-rapae lepidoptera %K reproductive output %X

The mating system of the Australian lycaenid butterfly, Jalmenus evagoras, is highly unusual compared to most other Lepidoptera. Characteristics of this system, which has been termed an 'explosive mating strategy,' include the formation of an intensely competitive mating aggregation of males, a highly male biased operational sex ratio, a lack of discrimination and mate choice by both sexes, a high variance in male mating success, and female monogamy. In this study, we tested the hypothesis that multiple mating by males imposes physiological costs resulting in smaller spermatophores, and that this results in a fitness cost to females. We found that male J. evagoras transferred only 2.2% of their eclosion weight during their first mating, consistent with the hypothesis that males of monandrous species produce a relatively small investment. The wet weight of the ejaculate declined by an average of 27% at the second mating and the dry weight by 29%, and an intermating interval of 5-9 days was needed for the ejaculate to return to the size at the first mating, regardless of male size or age. Wet ejaculate mass increased proportionally with male size, though dry mass was proportionally larger in smaller males. Ejaculate mass tended to increase with male age at both first and second matings. Female characteristics, in general, did not affect ejaculate mass, although the wet weight of the ejaculate was positively associated with female weight at the second mating. Copulation duration increased from 2.4 h to approximately 3 h at the second mating, and to over 4 h at the third and fourth matings. Fecundity was positively correlated with female size but not with mating history, copulation duration, or any other characteristics measured for either males or females. Female longevity declined significantly as the number of times the male partner had previously mated increased. We conclude that despite the small male investment in ejaculate, the costs of multiple mating may nonetheless be significant, as indicated by the reduction in ejaculate mass, an increase in copulation duration, and reduction in female lifespan with increasing mating number.

%B Behavioral Ecology and Sociobiology %V 47 %P 119-128 %8 Feb %@ 0340-5443 %G English %M WOS:000085378400002 %! Behav Ecol SociobiolBehav Ecol Sociobiol %0 Journal Article %J Molecular Biology and Evolution %D 2000 %T Molecular evolution of the Wingless gene and its implications for the phylogenetic placement of the butterfly family riodinidae (Lepidoptera : Papilionoidea) %A Campbell, D. L. %A Brower, A. V. Z. %A Pierce, N. E. %K character %K divergences %K Drosophila %K gene utility %K likelihood %K lycaenidae %K maximum likelihood %K molecular phylogenetics %K nuclear gene %K nymphalidae %K pattern %K protein %K riodinidae %K sequence data %K substitution %K third codon positions %K trees %K wingless %X

The sequence evolution of the nuclear gene wingless was investigated among 34 representatives of three lepidopteran families (Riodinidae, Lycaenidae, and Nymphalidae) and four outgroups, and its utility for inferring phylogenetic relationships among these taxa was assessed. Parsimony analysis yielded a well-resolved topology supporting the monophyly of the Riodinidae and Lycaenidae, respectively, and indicating that these two groups are sister lineages, with strong nodal support based on bootstrap and decay indices. Although, wingless provides robust support for relationships within and between the riodinids and the lycaenids, it is less informative about nymphalid relationships. Wingless does not consistently recover nymphalid monophyly or traditional subfamilial relationships within the nymphalids, and nodal support for all but the most recent branches in this family is low. Much of the phylogenetic information in this data set is derived from first- and second-position substitutions. However third positions, despite showing uncorrected pairwise divergences up to 78%, also contain consistent signal at deep nodes within the family Riodinidae and at the node defining the sister relationship between the riodinids and lycaenids. Several hypotheses about how third-position signal has been retained in deep nodes are discussed. These include among-site rate variation, identified as a significant factor by maximum likelihood analyses, and nucleotide bias, a prominent feature of third positions in this data set. Understanding the mechanisms which underlie third-position signal is a first step in applying appropriate models to accommodate the specific evolutionary processes involved in each lineage.

%B Molecular Biology and Evolution %V 17 %P 684-696 %8 May %@ 0737-4038 %G English %M WOS:000086875300002 %! Mol Biol EvolMol Biol Evol %0 Journal Article %J Nature Medicine %D 2000 %T W.D. Hamilton, 1936-2000 - Obituary %A Moran, N. %A Pierce, N. %A Seger, J. %B Nature Medicine %V 6 %P 367-367 %8 Apr %@ 1078-8956 %G English %M WOS:000165474100016 %! Nat MedNat Med %0 Journal Article %J Science %D 2000 %T William Hamilton (1936-2000) %A Haig, D. %A Pierce, N. E. %A Wilson, E. O. %B Science %V 287 %P 2438-2438 %8 Mar 31 %@ 0036-8075 %G English %M WOS:000086202200034 %! ScienceScience %0 Book %D 1999 %T The Biology of Australian Butterflies (Monographs on Australian Lepidoptera Vol. 6) %A Kitching, R %A E Sheermeyer %A R Jones %A Pierce, N. E. %I CSIRO Press %C Sydney %P 395 %G eng %0 Journal Article %J International Journal of Biological Macromolecules %D 1999 %T A comparison of the composition of silk proteins produced by spiders and insects %A Craig, C. L. %A Hsu, M. %A Kaplan, D. %A Pierce, N. E. %K amino acid %K araneae %K Evolution %K hymenoptera %K Insects %K larval silk %K metabolic cost %K molecular phylogeny %K nest %K paper %K pedicel %K selection %K silk %K spiders %X

Proteins that are highly expressed and composed of amino acids that are costly to synthesize are likely to place a greater drain on an organism's energy resources than proteins that are composed of ingested amino acids or ones that are metabolically simple to produce. Silks are highly expressed proteins produced by all spiders and many insects. We compared the metabolic costs of silks spun by arthropods by calculating the amount of ATP required to produce their component amino acids. Although a definitive conclusion requires detailed information on the dietary pools of amino acids available to arthropods, on the basis of the central metabolic pathways, silks spun by herbivorous, Lepidoptera larvae require significantly less ATP to synthesize than the dragline silks spun by predatory spiders. While not enough data are available to draw a statistically based conclusion, comparison of homologous silks across ancestral and derived taxa of the Araneoidea seems to suggest an evolutionary trend towards reduced silk costs. However, comparison of the synthetic costs of dragline silks across all araneomorph spiders suggests a complicated evolutionary pattern that cannot be attributed to phylogenetic position alone. We propose that the diverse silk-producing systems of the araneoid spiders (including three types of protein glues and three types of silk fibroin), evolved through intra-organ competition and that taxon-specific differences in the composition of silks drawn from homologous glands may reflect limited or fluctuating amino acid availability. The different functional properties of spider silks may be a secondary result of selection acting on different polypeptide templates. (C) 1999 Elsevier Science B.V. All rights reserved.

%B International Journal of Biological Macromolecules %V 24 %P 109-118 %8 Mar-Apr %@ 0141-8130 %G English %M WOS:000080158000005 %! Int J Biol MacromolInt J Biol Macromol %0 Book %D 1999 %T The Biology of Australian Butterflies (Monographs on Australian Lepidoptera Vol. 6) %A Kitching, R %A E Sheermeyer %A R Jones %A Pierce, N. E. %I CSIRO Press %C Sydney %P 395 %G eng %0 Book Section %B The Biology of Australian Butterflies (Monographs on Australian Lepidoptera) %D 1999 %T The Imperial Blue, Jalmenus evagoras (Lycaenidae) %A Pierce, N. E. %A DR Nash %B The Biology of Australian Butterflies (Monographs on Australian Lepidoptera) %I CSIRO Press %C Sydney %V 6 %P 277-316 %G eng %0 Journal Article %J International Journal of Biological Macromolecules %D 1999 %T A comparison of the composition of silk proteins produced by spiders and insects %A Craig, C. L. %A Hsu, M. %A Kaplan, D. %A Pierce, N. E. %K amino acid %K araneae %K Evolution %K hymenoptera %K Insects %K larval silk %K metabolic cost %K molecular phylogeny %K nest %K paper %K pedicel %K selection %K silk %K spiders %X

Proteins that are highly expressed and composed of amino acids that are costly to synthesize are likely to place a greater drain on an organism's energy resources than proteins that are composed of ingested amino acids or ones that are metabolically simple to produce. Silks are highly expressed proteins produced by all spiders and many insects. We compared the metabolic costs of silks spun by arthropods by calculating the amount of ATP required to produce their component amino acids. Although a definitive conclusion requires detailed information on the dietary pools of amino acids available to arthropods, on the basis of the central metabolic pathways, silks spun by herbivorous, Lepidoptera larvae require significantly less ATP to synthesize than the dragline silks spun by predatory spiders. While not enough data are available to draw a statistically based conclusion, comparison of homologous silks across ancestral and derived taxa of the Araneoidea seems to suggest an evolutionary trend towards reduced silk costs. However, comparison of the synthetic costs of dragline silks across all araneomorph spiders suggests a complicated evolutionary pattern that cannot be attributed to phylogenetic position alone. We propose that the diverse silk-producing systems of the araneoid spiders (including three types of protein glues and three types of silk fibroin), evolved through intra-organ competition and that taxon-specific differences in the composition of silks drawn from homologous glands may reflect limited or fluctuating amino acid availability. The different functional properties of spider silks may be a secondary result of selection acting on different polypeptide templates. (C) 1999 Elsevier Science B.V. All rights reserved.

%B International Journal of Biological Macromolecules %V 24 %P 109-118 %8 Mar-Apr %@ 0141-8130 %G English %M WOS:000080158000005 %! Int J Biol MacromolInt J Biol Macromol %0 Journal Article %J Behavioral Ecology %D 1998 %T Aggregation as a cost-reducing strategy for lycaenid larvae %A Axen, A. H. %A Pierce, N. E. %K aggregation %K ants %K Behavior %K biological markets %K butterflies %K cooperation %K cost-benefit relationship %K food rewards %K formicidae %K hymenoptera %K iridomyrmex %K jalmenus evagoras %K jalmenus-evagoras %K lycaenidae %K mutualism %K protection %K strategic behavior %X

If a mutualistic relationship entails providing services at a cost, selection will favor individuals that maximize the net benefits of the interaction and minimize the costs. Larvae of many species of lycaenid butterflies secrete nutritious food rewards to attending ants and, in return, receive protection against predators and parasitoids. Because ants typically recruit more workers to larger resources, by forming groups the larvae may ensure more reliable access to ants and thereby gain better protection. A further consequence of aggregating should be a change of the cost-benefit relationship for individual larvae. The larger the group, the smaller a single larva's influence will be on total ant density, which could lead to a smaller investment in secretion, thus reducing the per capita cost of cooperation. In this study, the influence of ant attendance, group size, and companion quality on larval investment was investigated. The interaction between the obligately ant-dependent lycaenid, Jalmenus evagoras, and its attendant Iridomyrmex ants was manipulated and the effect on larval secretion measured. As the level of ant attendance increased, the delivery of food rewards increased, both for solitary and for aggregated larvae. When aggregated, larvae provided less food rewards to ants than when solitary, and secretion rate decreased with increasing group size. Furthermore, larvae had lower secretion rates when paired with a bigger, more attractive larva than when paired with a smaller one. The considerable reduction in secretion rates for larvae in groups suggests that gaining protection at a lower secretion cost could be one factor that promotes aggregation in myrmecophilous lycaenids.

%B Behavioral Ecology %V 9 %P 109-115 %8 Mar-Apr %@ 1045-2249 %G English %M WOS:000072758700001 %! Behav EcolBehav Ecol %0 Journal Article %J Proceedings of the Royal Society B-Biological Sciences %D 1998 %T A castration parasite of an ant-plant mutualism %A Yu, D. W. %A Pierce, N. E. %K altruism %K ant-plant %K association %K castration parasite %K cooperation %K cost of reproduction %K Ecology %K eusocial evolution %K Evolution %K inhabitants %K Insect %K mutualism %K myrmecophyte %K nitrogen %K protection %K yucca moths %X

Exploring the factors governing the maintenance and breakdown of cooperation between mutualists is an intriguing and enduring problem for evolutionary ecology, and symbioses between ants and plants can provide useful experimental models for such studies. Hundreds of tropical plant species have evolved structures to house and feed ants, and these ant-plant symbioses have long been considered classic examples of mutualism. Here, we report that the primary ant symbiont, Allomerus cf. demerarae, of the most abundant ant-plant found in south-east Peru, Cordia nodosa Lam., castrates its host plant. Allomerus workers protect new leaves and their associated domatia from herbivory, but destroy flowers, reducing fruit production to zero in most host plants. Castrated plants occupied by Allomerus provide more domatia for their associated ants than plants occupied by three species of Azteca ants that do not castrate their hosts. Allomerus colonies in larger plants have higher fecundity. As a consequence, Allomerus appears to benefit from its castration behaviour, to the detriment of C. nodosa. The C. nodosa-ant system exhibits none of the retaliatory or filtering mechanisms shown to stabilize cheating in other cooperative systems, and appears to persist because some of the plants, albeit a small;minority, are inhabited by the three species of truly mutualistic Azteca ants.

%B Proceedings of the Royal Society B-Biological Sciences %V 265 %P 375-382 %8 Mar 7 %@ 0962-8452 %G English %M WOS:000072540500004 %! P Roy Soc B-Biol SciP Roy Soc B-Biol Sci %0 Journal Article %J Journal of Neuroscience %D 1998 %T Honeybee blue-and ultraviolet-sensitive opsins: Cloning, heterologous expression in Drosophila, and physiological characterization %A Townson, S. M. %A Chang, B. S. W. %A Salcedo, E. %A Chadwell, L. V. %A Pierce, N. E. %A Britt, S. G. %K absorption-spectra %K apis mellifera %K blue-sensitive opsin %K bovine rhodopsin %K cysteine residue-110 %K drone apis-mellifera %K drosophila melanogaster %K electroretinogram %K honeybee %K human color-vision %K molecular-genetics %K photoreceptor cells %K protein expression %K schiff-base counterion %K spectral sensitivity %K spectral tuning %K ultraviolet-sensitive opsin %K visual pigment %K visual pigments %X

The honeybee (Apis mellifera) visual system contains three classes of retinal photoreceptor cells that are maximally sensitive to light at 440 nm (blue), 350 nm (ultraviolet), and 540 nm (green), We performed a PCR-based screen to identify the genes encoding the Apis blue- and ultraviolet (UV)-sensitive opsins, We obtained cDNAs that encode proteins having a high degree of sequence and structural similarity to other invertebrate and vertebrate visual pigments. The Apis blue opsin cDNA encodes a protein of 377 amino acids that is most closely related to other invertebrate visual pigments that are thought to be blue-sensitive. The UV opsin cDNA encodes a protein of 371 amino acids that is most closely related to the UV-sensitive Drosophila Rh3 and Rh4 opsins. To test whether these novel Apis opsin genes encode functional visual pigments and to determine their spectral properties, we expressed them in the R1-6 photoreceptor cells of blind ninaE mutant Drosophila, which lack the major opsin of the fly compound eye. We found that the expression of either the Apis blue- or UV-sensitive opsin in transgenic flies rescued the visual defect of ninaE mutants, indicating that both genes encode functional visual pigments. Spectral sensitivity measurements of these flies demonstrated that the blue and UV visual pigments are maximally sensitive to light at 439 and 353 nm, respectively. These maxima are in excellent agreement with those determined previously by single-cell recordings from Apis photoreceptor cells and provide definitive evidence that the genes described here encode visual pigments having blue and UV sensitivity.

%B Journal of Neuroscience %V 18 %P 2412-2422 %8 Apr 1 %@ 0270-6474 %G English %M WOS:000072778700012 %! J NeurosciJ Neurosci %0 Journal Article %J Behavioral Ecology %D 1998 %T Aggregation as a cost-reducing strategy for lycaenid larvae %A Axen, A. H. %A Pierce, N. E. %K aggregation %K ants %K Behavior %K biological markets %K butterflies %K cooperation %K cost-benefit relationship %K food rewards %K formicidae %K hymenoptera %K iridomyrmex %K jalmenus evagoras %K jalmenus-evagoras %K lycaenidae %K mutualism %K protection %K strategic behavior %X

If a mutualistic relationship entails providing services at a cost, selection will favor individuals that maximize the net benefits of the interaction and minimize the costs. Larvae of many species of lycaenid butterflies secrete nutritious food rewards to attending ants and, in return, receive protection against predators and parasitoids. Because ants typically recruit more workers to larger resources, by forming groups the larvae may ensure more reliable access to ants and thereby gain better protection. A further consequence of aggregating should be a change of the cost-benefit relationship for individual larvae. The larger the group, the smaller a single larva's influence will be on total ant density, which could lead to a smaller investment in secretion, thus reducing the per capita cost of cooperation. In this study, the influence of ant attendance, group size, and companion quality on larval investment was investigated. The interaction between the obligately ant-dependent lycaenid, Jalmenus evagoras, and its attendant Iridomyrmex ants was manipulated and the effect on larval secretion measured. As the level of ant attendance increased, the delivery of food rewards increased, both for solitary and for aggregated larvae. When aggregated, larvae provided less food rewards to ants than when solitary, and secretion rate decreased with increasing group size. Furthermore, larvae had lower secretion rates when paired with a bigger, more attractive larva than when paired with a smaller one. The considerable reduction in secretion rates for larvae in groups suggests that gaining protection at a lower secretion cost could be one factor that promotes aggregation in myrmecophilous lycaenids.

%B Behavioral Ecology %V 9 %P 109-115 %8 Mar-Apr %@ 1045-2249 %G English %M WOS:000072758700001 %! Behav EcolBehav Ecol %0 Journal Article %J Proceedings of the Royal Society B-Biological Sciences %D 1998 %T A castration parasite of an ant-plant mutualism %A Yu, D. W. %A Pierce, N. E. %K altruism %K ant-plant %K association %K castration parasite %K cooperation %K cost of reproduction %K Ecology %K eusocial evolution %K Evolution %K inhabitants %K Insect %K mutualism %K myrmecophyte %K nitrogen %K protection %K yucca moths %X

Exploring the factors governing the maintenance and breakdown of cooperation between mutualists is an intriguing and enduring problem for evolutionary ecology, and symbioses between ants and plants can provide useful experimental models for such studies. Hundreds of tropical plant species have evolved structures to house and feed ants, and these ant-plant symbioses have long been considered classic examples of mutualism. Here, we report that the primary ant symbiont, Allomerus cf. demerarae, of the most abundant ant-plant found in south-east Peru, Cordia nodosa Lam., castrates its host plant. Allomerus workers protect new leaves and their associated domatia from herbivory, but destroy flowers, reducing fruit production to zero in most host plants. Castrated plants occupied by Allomerus provide more domatia for their associated ants than plants occupied by three species of Azteca ants that do not castrate their hosts. Allomerus colonies in larger plants have higher fecundity. As a consequence, Allomerus appears to benefit from its castration behaviour, to the detriment of C. nodosa. The C. nodosa-ant system exhibits none of the retaliatory or filtering mechanisms shown to stabilize cheating in other cooperative systems, and appears to persist because some of the plants, albeit a small;minority, are inhabited by the three species of truly mutualistic Azteca ants.

%B Proceedings of the Royal Society B-Biological Sciences %V 265 %P 375-382 %8 Mar 7 %@ 0962-8452 %G English %M WOS:000072540500004 %! P Roy Soc B-Biol SciP Roy Soc B-Biol Sci %0 Journal Article %J Journal of Neuroscience %D 1998 %T Honeybee blue-and ultraviolet-sensitive opsins: Cloning, heterologous expression in Drosophila, and physiological characterization %A Townson, S. M. %A Chang, B. S. W. %A Salcedo, E. %A Chadwell, L. V. %A Pierce, N. E. %A Britt, S. G. %K absorption-spectra %K apis mellifera %K blue-sensitive opsin %K bovine rhodopsin %K cysteine residue-110 %K drone apis-mellifera %K drosophila melanogaster %K electroretinogram %K honeybee %K human color-vision %K molecular-genetics %K photoreceptor cells %K protein expression %K schiff-base counterion %K spectral sensitivity %K spectral tuning %K ultraviolet-sensitive opsin %K visual pigment %K visual pigments %X

The honeybee (Apis mellifera) visual system contains three classes of retinal photoreceptor cells that are maximally sensitive to light at 440 nm (blue), 350 nm (ultraviolet), and 540 nm (green), We performed a PCR-based screen to identify the genes encoding the Apis blue- and ultraviolet (UV)-sensitive opsins, We obtained cDNAs that encode proteins having a high degree of sequence and structural similarity to other invertebrate and vertebrate visual pigments. The Apis blue opsin cDNA encodes a protein of 377 amino acids that is most closely related to other invertebrate visual pigments that are thought to be blue-sensitive. The UV opsin cDNA encodes a protein of 371 amino acids that is most closely related to the UV-sensitive Drosophila Rh3 and Rh4 opsins. To test whether these novel Apis opsin genes encode functional visual pigments and to determine their spectral properties, we expressed them in the R1-6 photoreceptor cells of blind ninaE mutant Drosophila, which lack the major opsin of the fly compound eye. We found that the expression of either the Apis blue- or UV-sensitive opsin in transgenic flies rescued the visual defect of ninaE mutants, indicating that both genes encode functional visual pigments. Spectral sensitivity measurements of these flies demonstrated that the blue and UV visual pigments are maximally sensitive to light at 439 and 353 nm, respectively. These maxima are in excellent agreement with those determined previously by single-cell recordings from Apis photoreceptor cells and provide definitive evidence that the genes described here encode visual pigments having blue and UV sensitivity.

%B Journal of Neuroscience %V 18 %P 2412-2422 %8 Apr 1 %@ 0270-6474 %G English %M WOS:000072778700012 %! J NeurosciJ Neurosci %0 Journal Article %J Gene %D 1996 %T Cloning of the gene encoding honeybee long-wavelength rhodopsin: A new class of insect visual pigments %A Chang, B. S. W. %A Ayers, D. %A Smith, W. C. %A Pierce, N. E. %K apis mellifera %K bovine rhodopsin %K color-vision %K g protein-coupled receptor %K hymenoptera %K model %K molecular evolution %K opsin %K Phosphorylation %K photoreceptor %K Phylogeny %K protein-linked receptors %K schiff-base counterion %K shifts %K sites %K vision %K wavelength regulation %X

Rhodopsins (Rh), G-protein-coupled receptors with seven transmembrane (TM) helices, form the first step in visual transduction in most organisms. Although many long-wavelength (LW) vertebrate opsin sequences are known, less information is available for invertebrate LW sequences. By a combination of RT-PCR and cDNA library screening, we have cloned and sequenced the honeybee LW Rh gene. The deduced protein is composed of 378 amino acids (aa), appears to have seven TM regions, and contains many of the structures and key aa thought to be important for Rh function. Phylogenetic analysis of this sequence in relation to other invertebrate Rh reveals it to be a member of a new group of insect LW Rh.

%B Gene %V 173 %P 215-219 %8 Sep 16 %@ 0378-1119 %G English %M WOS:A1996VH18600015 %! GeneGene %0 Journal Article %J Insectarium %D 1996 %T A scientist of great humanity %A Pierce, N. E. %B Insectarium %V 33 %P 3 %G eng %0 Book Section %B Social competition and cooperation in insects and arachnids, Volume II: Evolution of sociality %D 1996 %T Social evolution in the Lepidoptera: ecological context and communication in larval societies. %A Costa, JT %A Pierce, N. E. %B Social competition and cooperation in insects and arachnids, Volume II: Evolution of sociality %V 2 %P 407-442 %G eng %0 Journal Article %J Insectarium %D 1996 %T A scientist of great humanity %A Pierce, N. E. %B Insectarium %V 33 %P 3 %G eng %0 Book Section %B Social competition and cooperation in insects and arachnids, Volume II: Evolution of sociality %D 1996 %T Social evolution in the Lepidoptera: ecological context and communication in larval societies. %A Costa, JT %A Pierce, N. E. %B Social competition and cooperation in insects and arachnids, Volume II: Evolution of sociality %V 2 %P 407-442 %G eng %0 Journal Article %J Gene %D 1996 %T Cloning of the gene encoding honeybee long-wavelength rhodopsin: A new class of insect visual pigments %A Chang, B. S. W. %A Ayers, D. %A Smith, W. C. %A Pierce, N. E. %K apis mellifera %K bovine rhodopsin %K color-vision %K g protein-coupled receptor %K hymenoptera %K model %K molecular evolution %K opsin %K Phosphorylation %K photoreceptor %K Phylogeny %K protein-linked receptors %K schiff-base counterion %K shifts %K sites %K vision %K wavelength regulation %X

Rhodopsins (Rh), G-protein-coupled receptors with seven transmembrane (TM) helices, form the first step in visual transduction in most organisms. Although many long-wavelength (LW) vertebrate opsin sequences are known, less information is available for invertebrate LW sequences. By a combination of RT-PCR and cDNA library screening, we have cloned and sequenced the honeybee LW Rh gene. The deduced protein is composed of 378 amino acids (aa), appears to have seven TM regions, and contains many of the structures and key aa thought to be important for Rh function. Phylogenetic analysis of this sequence in relation to other invertebrate Rh reveals it to be a member of a new group of insect LW Rh.

%B Gene %V 173 %P 215-219 %8 Sep 16 %@ 0378-1119 %G English %M WOS:A1996VH18600015 %! GeneGene %0 Journal Article %J Biological Journal of the Linnean Society %D 1996 %T The effect of ant association on the population genetics of the Australian Butterfly Jalmenus evagoras (Lepidoptera: Lycaenidae) %A Costa, J. T. %A McDonald, J. H. %A Pierce, N. E. %K attendant ants %K differentiation %K Dispersal %K extinction recolonization %K flow %K formicidae %K genetic structure %K group selection %K iridomyrmex %K larvae %K meat ants %K mutualism %K myrmecophily %K natural-populations %K oviposition preference %K plant %K wright's f-statistics %X

Populations of the myrmecophilous lycaenid Falmenus evagoras Donovan were assessed for genetic structure at three hierarchical spatial scales: sites, geographically-defined subpopulations, and subpopulations defined by species of mutualistic ant-associate. Estimates of Wright's F-ST generated from multilocus electrophoretic data revealed low, though significant, amounts of genetic structure. Most structure was observed at the level of geographic subpopulations, suggesting that adult butterflies do not exhibit preferential mating and oviposition along the lines of ant associate. The genetic structure data, together with estimates of Nei's genetic distance (D) for pairwise site and subpopulation comparisons, suggest that F. evagoras populations are spatially and temporally dynamic. These patterns are considered in the context of extinction and recolonization models. The extreme patchiness of F. evagoras populations stems from the stringent requirements of both host plant and host ant, contributing to an extinction/ recolonization process. We discuss the key parameters influencing genetic cohesion versus differentiation under an extinction/recolonization regime, including mode of butterfly dispersal, site turnover rate, and the effects of host dispersal and phenology. This system provides a model of population-level consequences of certain mutualistic interactions as well as of a class of patterns arising from an extinction/recolonization process. (C) 1996 The Linnean Society of London

%B Biological Journal of the Linnean Society %V 58 %P 287-306 %8 Jul %@ 0024-4066 %G English %M WOS:A1996UY83000003 %! Biol J Linn SocBiol J Linn Soc %0 Journal Article %J Journal of the New York Entomological Society %D 1995 %T Effects of diet quality and queen number on growth in leptothoracine ant colonies (Hymenoptera: Formicidae) %A Evans, J. D. %A Pierce, N. E. %K brood development %K Diet %K Evolution %K formicidae %K leptothorax %K myrmica %K pheromonal inhibition %K polygyny %K Reproduction %K solenopsis-invicta %X

Laboratory experiments manipulating the diet of colonies of the facultatively polygynous ant, Leptothorax curvispinosus (Mayr), demonstrated that carbohydrates and protein have synergistic effects on egg numbers and brood production in colonies of this ant. Colonies receiving insect prey and sucrose grew significantly faster than colonies reared on unlimited supplies of either of these food types alone. This study also measured the effect of queen number on colony growth rates. Because the occurrence of multiple queens might affect colony growth only under certain nutritional conditions, polygynous colonies were reared in each of the three diet treatments. Queen number did not affect colony worker production in any of the three diet treatments; thus, individual queens in polygynous colonies produced far fewer workers per queen than did queens in monogynous colonies. There were no interaction effects between queen number and diet on colony growth. Several colonies which lacked morphologically distinct queens produced workers over the course of the experiment Using artificially established colonies of unmated workers, we found no evidence for parthenogenetic (thelytokous) reproduction in these colonies.

%B Journal of the New York Entomological Society %V 103 %P 91-99 %8 Jan %@ 0028-7199 %G English %M WOS:A1995TP95900010 %! J New York Entomol SJ New York Entomol S %0 Journal Article %J Journal of Research on the Lepidoptera %D 1995 %T Exploitation of lycaenid-ant mutalisms by braconid parasitoids. %A Fiedler, K %A P Seufert %A Pierce, N. E. %A JG Pearson %A T Baumgarten %X

Abstract. Larvae of 17 Lycaenidae butterfly species from Europe, North America, South East Asia and Australia were observed to retain at least some of their adaptations related to myrmecophily even after parasitic braconid larvae have emerged from them. The myrmecophilous glandular organs and vibratory muscles of such larval carcasses remain functional for up to 8 days. The cuticle of lycaenid larvae contains extractable “adoption substances” which elicit antennal drumming in their tending ants. These adoption substances, as well, appear to persist in a functional state beyond parasitoid emergence, and the larval carcasses are hence tended much like healthy caterpillars. In all examples, the braconids may receive selective advantages through myrmecophily of their host larvae, instead of being suppressed by the ant guard. Interactions where parasitoids exploit the ant-mutualism of their lycaenid hosts have as yet been recorded only from the Apanteles group in the BraconidaeMicrogasterinae.

%B Journal of Research on the Lepidoptera %V 31 %P 153-168 %G eng %0 Journal Article %J Journal of the Lepidopterists' Society %D 1995 %T Predatory and parasitic Lepidoptera: carnivores living on plants %A Pierce, N. E. %X

Moths and butterflies whose larvae do not feed on plants represent a decided minority slice of lepidopteran diversity, yet offer insights into the ecology and evolution of feeding habits. This paper summarizes the life histories of the known predatory and parasitic lepidopteran taxa, focusing in detail on current researchin the butterfly family Lycaenidae, a group disprotionately rich in aphytophagous feeders and myrmecophilous habits.

%B Journal of the Lepidopterists' Society %V 49 %P 412-453 %G eng %0 Journal Article %J Journal of Research on the Lepidoptera %D 1995 %T Exploitation of lycaenid-ant mutalisms by braconid parasitoids. %A Fiedler, K %A P Seufert %A Pierce, N. E. %A JG Pearson %A T Baumgarten %X

Abstract. Larvae of 17 Lycaenidae butterfly species from Europe, North America, South East Asia and Australia were observed to retain at least some of their adaptations related to myrmecophily even after parasitic braconid larvae have emerged from them. The myrmecophilous glandular organs and vibratory muscles of such larval carcasses remain functional for up to 8 days. The cuticle of lycaenid larvae contains extractable “adoption substances” which elicit antennal drumming in their tending ants. These adoption substances, as well, appear to persist in a functional state beyond parasitoid emergence, and the larval carcasses are hence tended much like healthy caterpillars. In all examples, the braconids may receive selective advantages through myrmecophily of their host larvae, instead of being suppressed by the ant guard. Interactions where parasitoids exploit the ant-mutualism of their lycaenid hosts have as yet been recorded only from the Apanteles group in the BraconidaeMicrogasterinae.

%B Journal of Research on the Lepidoptera %V 31 %P 153-168 %G eng %0 Journal Article %J Journal of the Lepidopterists' Society %D 1995 %T Predatory and parasitic Lepidoptera: carnivores living on plants %A Pierce, N. E. %X

Moths and butterflies whose larvae do not feed on plants represent a decided minority slice of lepidopteran diversity, yet offer insights into the ecology and evolution of feeding habits. This paper summarizes the life histories of the known predatory and parasitic lepidopteran taxa, focusing in detail on current researchin the butterfly family Lycaenidae, a group disprotionately rich in aphytophagous feeders and myrmecophilous habits.

%B Journal of the Lepidopterists' Society %V 49 %P 412-453 %G eng %0 Journal Article %J Journal of the New York Entomological Society %D 1995 %T Effects of diet quality and queen number on growth in leptothoracine ant colonies (Hymenoptera: Formicidae) %A Evans, J. D. %A Pierce, N. E. %K brood development %K Diet %K Evolution %K formicidae %K leptothorax %K myrmica %K pheromonal inhibition %K polygyny %K Reproduction %K solenopsis-invicta %X

Laboratory experiments manipulating the diet of colonies of the facultatively polygynous ant, Leptothorax curvispinosus (Mayr), demonstrated that carbohydrates and protein have synergistic effects on egg numbers and brood production in colonies of this ant. Colonies receiving insect prey and sucrose grew significantly faster than colonies reared on unlimited supplies of either of these food types alone. This study also measured the effect of queen number on colony growth rates. Because the occurrence of multiple queens might affect colony growth only under certain nutritional conditions, polygynous colonies were reared in each of the three diet treatments. Queen number did not affect colony worker production in any of the three diet treatments; thus, individual queens in polygynous colonies produced far fewer workers per queen than did queens in monogynous colonies. There were no interaction effects between queen number and diet on colony growth. Several colonies which lacked morphologically distinct queens produced workers over the course of the experiment Using artificially established colonies of unmated workers, we found no evidence for parthenogenetic (thelytokous) reproduction in these colonies.

%B Journal of the New York Entomological Society %V 103 %P 91-99 %8 Jan %@ 0028-7199 %G English %M WOS:A1995TP95900010 %! J New York Entomol SJ New York Entomol S %0 Journal Article %J Journal of the New York Entomological Society %D 1994 %T Absence of Kin Discrimination Behavior in a Soldier-Producing Aphid, Ceratovacuna-Japonica (Hemiptera, Pemphigidae, Cerataphidini) %A Carlin, N. F. %A Gladstein, D. S. %A Berry, A. J. %A Pierce, N. E. %X

Certain aphid species produce sterile soldiers, morphologically and behaviorally specialized individuals that defend fertile colony members, some or all of which are clonemates. If the soldier morph is maintained by inclusive fitness advantages, its altruism should preferentially benefit relatives, suggesting a potential role for kin discrimination. We performed a field experiment on spatial segregation and two laboratory experiments on agonistic behavior among non-soldiers and by soldiers of the cerataphidine aphid, Ceratovacuna japonica. For the test of spatial segregation, we introduce a new method of nearest-neighbor analysis, by constructing a minimum spanning tree from the map of individual locations and comparing the numbers of within-group and between-group connections. The results provide no evidence of kin recognition abilities in this species. Members of different clones showed no tendency to segregate spatially, nor to direct displacement attempts against non-kin when competing for feeding sites. Soldiers were indiscriminately aggressive toward early instar reproductives from their own and other colonies. We discuss the implications of these findings for several evolutionary hypotheses on the maintenance of the soldier morph in aphids.

%B Journal of the New York Entomological Society %V 102 %P 287-298 %8 Jul %@ 0028-7199 %G English %M WOS:A1994QP37800001 %! J New York Entomol SJ New York Entomol S %0 Book Section %B Molecular methods in ecology and evolution %D 1994 %T Diversity within diversity: molecular approaches to studying microbial interactions with insects. %A Kane, M %A Pierce, N. E. %B Molecular methods in ecology and evolution %I Birkhauser Verlag %P 509-524 %G eng %0 Book Section %B Molecular methods in ecology and evolution %D 1994 %T Diversity within diversity: molecular approaches to studying microbial interactions with insects. %A Kane, M %A Pierce, N. E. %B Molecular methods in ecology and evolution %I Birkhauser Verlag %P 509-524 %G eng %0 Journal Article %J Journal of the New York Entomological Society %D 1994 %T Absence of Kin Discrimination Behavior in a Soldier-Producing Aphid, Ceratovacuna-Japonica (Hemiptera, Pemphigidae, Cerataphidini) %A Carlin, N. F. %A Gladstein, D. S. %A Berry, A. J. %A Pierce, N. E. %X

Certain aphid species produce sterile soldiers, morphologically and behaviorally specialized individuals that defend fertile colony members, some or all of which are clonemates. If the soldier morph is maintained by inclusive fitness advantages, its altruism should preferentially benefit relatives, suggesting a potential role for kin discrimination. We performed a field experiment on spatial segregation and two laboratory experiments on agonistic behavior among non-soldiers and by soldiers of the cerataphidine aphid, Ceratovacuna japonica. For the test of spatial segregation, we introduce a new method of nearest-neighbor analysis, by constructing a minimum spanning tree from the map of individual locations and comparing the numbers of within-group and between-group connections. The results provide no evidence of kin recognition abilities in this species. Members of different clones showed no tendency to segregate spatially, nor to direct displacement attempts against non-kin when competing for feeding sites. Soldiers were indiscriminately aggressive toward early instar reproductives from their own and other colonies. We discuss the implications of these findings for several evolutionary hypotheses on the maintenance of the soldier morph in aphids.

%B Journal of the New York Entomological Society %V 102 %P 287-298 %8 Jul %@ 0028-7199 %G English %M WOS:A1994QP37800001 %! J New York Entomol SJ New York Entomol S %0 Book Section %B Caterpillars: Ecological and Evolutionary Constraints on Foraging %D 1993 %T The effects of ant mutualism on the foraging and diet of lycaenid caterpillars. %A Baylis, M %A Pierce, N. E. %B Caterpillars: Ecological and Evolutionary Constraints on Foraging %I Chapman and Hall %C New York %P 404-421 %G eng %0 Journal Article %J Molecular Biology and Evolution %D 1993 %T The Lepidopteran Mitochondrial Control Region - Structure and Evolution %A Taylor, M. F. J. %A Mckechnie, S. W. %A Pierce, N. %A Kreitman, M. %K control region %K DNA %K Evolution %K gene organization %K genome %K heteroplasmy %K jalmenus-evagoras %K lepidoptera %K mitochondrial %K model %K nucleotide-sequence %K ribosomal-subunit rna %K size %K tetrahymena %X

For several species of lepidoptera, most of the similar to 350-bp mitochondrial control-region sequences were determined. Six of these species are in one genus, Jalmenus; are closely related; and are believed to have undergone recent rapid speciation. Recent speciation was supported by the observation of low interspecific sequence divergence. Thus, no useful phylogeny could be constructed for the genus. Despite a surprising conservation of control-region length, there was little conservation of primary sequences either among the three lepidopteran genera or between lepidoptera and Drosophila. Analysis of secondary structure indicated only one possible feature in common-inferred stem loops with higher-than-random folding energies-although the positions of the structures in different species were unrelated to regions of primary sequence similarity. We suggest that the conserved, short length of control regions is related to the observed lack of heteroplasmy in lepidopteran mitochondrial genomes. In addition, determination of flanking sequences for one Jalmenus species indicated (i) only weak support for the available model of insect 12S rRNA structure and (ii) that tRNA translocation is a frequent event in the evolution of insect mitochondrial genomes.

%B Molecular Biology and Evolution %V 10 %P 1259-1272 %8 Nov %@ 0737-4038 %G English %M WOS:A1993MH06200009 %! Mol Biol EvolMol Biol Evol %0 Book Section %B Caterpillars: Ecological and Evolutionary Constraints on Foraging %D 1993 %T The effects of ant mutualism on the foraging and diet of lycaenid caterpillars. %A Baylis, M %A Pierce, N. E. %B Caterpillars: Ecological and Evolutionary Constraints on Foraging %I Chapman and Hall %C New York %P 404-421 %G eng %0 Journal Article %J Molecular Biology and Evolution %D 1993 %T The Lepidopteran Mitochondrial Control Region - Structure and Evolution %A Taylor, M. F. J. %A Mckechnie, S. W. %A Pierce, N. %A Kreitman, M. %K control region %K DNA %K Evolution %K gene organization %K genome %K heteroplasmy %K jalmenus-evagoras %K lepidoptera %K mitochondrial %K model %K nucleotide-sequence %K ribosomal-subunit rna %K size %K tetrahymena %X

For several species of lepidoptera, most of the similar to 350-bp mitochondrial control-region sequences were determined. Six of these species are in one genus, Jalmenus; are closely related; and are believed to have undergone recent rapid speciation. Recent speciation was supported by the observation of low interspecific sequence divergence. Thus, no useful phylogeny could be constructed for the genus. Despite a surprising conservation of control-region length, there was little conservation of primary sequences either among the three lepidopteran genera or between lepidoptera and Drosophila. Analysis of secondary structure indicated only one possible feature in common-inferred stem loops with higher-than-random folding energies-although the positions of the structures in different species were unrelated to regions of primary sequence similarity. We suggest that the conserved, short length of control regions is related to the observed lack of heteroplasmy in lepidopteran mitochondrial genomes. In addition, determination of flanking sequences for one Jalmenus species indicated (i) only weak support for the available model of insect 12S rRNA structure and (ii) that tRNA translocation is a frequent event in the evolution of insect mitochondrial genomes.

%B Molecular Biology and Evolution %V 10 %P 1259-1272 %8 Nov %@ 0737-4038 %G English %M WOS:A1993MH06200009 %! Mol Biol EvolMol Biol Evol %0 Journal Article %J Physiological Entomology %D 1992 %T Lack of Compensation by Final Instar Larvae of the Myrmecophilous Lycaenid Butterfly, Jalmenus-Evagoras, for the Loss of Nutrients to Ants %A Baylis, M. %A Pierce, N. E. %K ant attendance %K artificial diet %K compensation %K efficiency %K jalmenus-evagoras %K lepidoptera %K lycaenidae %K mutualism %K myrmecophily %K nutrient loss %K water-content %X

Larvae and pupae of the Australian lycaenid butterfly, Jalmenus evagoras Donovan (Lepidoptera; Lycaenidae), are protected from parasites and predators by attendant ants. In return, the juveniles of J.evagoras secrete to the ants a solution containing substantial amounts of sugars and amino acids. Larvae of J.evagoras were reared from hatching until adult eclosion either with or without ants. Experiments were performed to examine whether fifth (final) instar larvae attempt to compensate for the nutrient loss to ants, by consuming more food, digesting food more efficiently, or extending development time. The presence or absence of ants had no effect on the feeding rate, efficiency of digestion or development time of fifth instar larvae. Larvae with ants converted a smaller proportion of ingested food into biomass, and consequently grew less than their counterparts without ants. Thus fifth instar larvae of J.evagoras do not appear to compensate for the nutrient loss to ants. Possible reasons for the failure to compensate are discussed.

%B Physiological Entomology %V 17 %P 107-114 %8 Jun %@ 0307-6962 %G English %M WOS:A1992HZ99700001 %! Physiol EntomolPhysiol Entomol %0 Journal Article %J Physiological Entomology %D 1992 %T Lack of Compensation by Final Instar Larvae of the Myrmecophilous Lycaenid Butterfly, Jalmenus-Evagoras, for the Loss of Nutrients to Ants %A Baylis, M. %A Pierce, N. E. %K ant attendance %K artificial diet %K compensation %K efficiency %K jalmenus-evagoras %K lepidoptera %K lycaenidae %K mutualism %K myrmecophily %K nutrient loss %K water-content %X

Larvae and pupae of the Australian lycaenid butterfly, Jalmenus evagoras Donovan (Lepidoptera; Lycaenidae), are protected from parasites and predators by attendant ants. In return, the juveniles of J.evagoras secrete to the ants a solution containing substantial amounts of sugars and amino acids. Larvae of J.evagoras were reared from hatching until adult eclosion either with or without ants. Experiments were performed to examine whether fifth (final) instar larvae attempt to compensate for the nutrient loss to ants, by consuming more food, digesting food more efficiently, or extending development time. The presence or absence of ants had no effect on the feeding rate, efficiency of digestion or development time of fifth instar larvae. Larvae with ants converted a smaller proportion of ingested food into biomass, and consequently grew less than their counterparts without ants. Thus fifth instar larvae of J.evagoras do not appear to compensate for the nutrient loss to ants. Possible reasons for the failure to compensate are discussed.

%B Physiological Entomology %V 17 %P 107-114 %8 Jun %@ 0307-6962 %G English %M WOS:A1992HZ99700001 %! Physiol EntomolPhysiol Entomol %0 Journal Article %J Ecological Entomology %D 1991 %T The Effect of Host-Plant Quality on the Survival of Larvae and Oviposition by Adults of an Ant-Tended Lycaenid Butterfly, Jalmenus-Evagoras %A Baylis, M. %A Pierce, N. E. %K ant attendance %K Behavior %K cinnabar moth %K coevolution %K euphydryas-chalcedona %K growth %K host-plant quality %K jalmenus-evagoras %K lepidoptera %K lycaenidae %K nitrogen %K nitrogenous fertilizer %K oviposition %K pieris-rapae %K protein %K salvinia moth %K selection %X

1. Juveniles of the Australian lycaenid butterfly, Jalmenus evagoras (Donovan), secrete to ants a solution of sugars and amino acids, primarily serine. The attendant ants protect the larvae and pupae from parasites and predators.2. The effect of caterpillar nutrition on the defence provided by ants was investigated. Potted food plants of Acacia decurrens were either given water containing nitrogenous fertilizer or were given water alone. Fertilized plants had a higher nitrogen content than unfertilized plants.3. Fifth instar larvae of J.evagoras feeding on fertilized plants attracted a larger ant guard than those feeding on unfertilized plants. In the absence of caterpillars, ants were not differentially attracted to fertilized and unfertilized plants.4. In the presence of ants, over a 10-day period, larvae on fertilized plants survived better than larvae on unfertilized plants. In the absence of ants larvae survived equally on fertilized and unfertilized plants. It is concluded that larvae on fertilized plants attracted a larger ant guard, and thereby survived better, than larvae on unfertilized plants.5. Adult females of J.evagoras preferred to lay egg batches on fertiizedd rather than unfertilized plants, but they did not lay larger egg batches.

%B Ecological Entomology %V 16 %P 1-9 %8 Feb %@ 0307-6946 %G English %M WOS:A1991ET42600001 %! Ecol EntomolEcol Entomol %0 Book Section %B Ant - Plant Interactions %D 1991 %T Variation in the Attractiveness of Lycaenid Butterfly Larvae to Ants %A Pierce, N. E. %A Nash, D. R. %A Baylis, M. %A Carper, E. R. %B Ant - Plant Interactions %I Oxford University Press %C Oxford %P 131-142 %G English %M WOS:A1991BV72N00011 %0 Journal Article %J Ecological Entomology %D 1991 %T The Effect of Host-Plant Quality on the Survival of Larvae and Oviposition by Adults of an Ant-Tended Lycaenid Butterfly, Jalmenus-Evagoras %A Baylis, M. %A Pierce, N. E. %K ant attendance %K Behavior %K cinnabar moth %K coevolution %K euphydryas-chalcedona %K growth %K host-plant quality %K jalmenus-evagoras %K lepidoptera %K lycaenidae %K nitrogen %K nitrogenous fertilizer %K oviposition %K pieris-rapae %K protein %K salvinia moth %K selection %X

1. Juveniles of the Australian lycaenid butterfly, Jalmenus evagoras (Donovan), secrete to ants a solution of sugars and amino acids, primarily serine. The attendant ants protect the larvae and pupae from parasites and predators.2. The effect of caterpillar nutrition on the defence provided by ants was investigated. Potted food plants of Acacia decurrens were either given water containing nitrogenous fertilizer or were given water alone. Fertilized plants had a higher nitrogen content than unfertilized plants.3. Fifth instar larvae of J.evagoras feeding on fertilized plants attracted a larger ant guard than those feeding on unfertilized plants. In the absence of caterpillars, ants were not differentially attracted to fertilized and unfertilized plants.4. In the presence of ants, over a 10-day period, larvae on fertilized plants survived better than larvae on unfertilized plants. In the absence of ants larvae survived equally on fertilized and unfertilized plants. It is concluded that larvae on fertilized plants attracted a larger ant guard, and thereby survived better, than larvae on unfertilized plants.5. Adult females of J.evagoras preferred to lay egg batches on fertiizedd rather than unfertilized plants, but they did not lay larger egg batches.

%B Ecological Entomology %V 16 %P 1-9 %8 Feb %@ 0307-6946 %G English %M WOS:A1991ET42600001 %! Ecol EntomolEcol Entomol %0 Book Section %B Ant - Plant Interactions %D 1991 %T Variation in the Attractiveness of Lycaenid Butterfly Larvae to Ants %A Pierce, N. E. %A Nash, D. R. %A Baylis, M. %A Carper, E. R. %B Ant - Plant Interactions %I Oxford University Press %C Oxford %P 131-142 %G English %M WOS:A1991BV72N00011 %0 Book Section %B Towards a more exact ecology %D 1989 %T Butterfly-ant mutualisms %A Pierce, N. E. %B Towards a more exact ecology %I Blackwell %C Oxford %P 299-324 %G eng %0 Journal Article %J Oecologia %D 1989 %T The Effect of Adult Diet on the Biology of Butterflies .1. The Common Imperial Blue, Jalmenus-Evagoras %A Hill, C. J. %A Pierce, N. E. %B Oecologia %V 81 %P 249-257 %@ 0029-8549 %G English %M WOS:A1989AY34400016 %! OecologiaOecologia %0 Book Section %B Towards a more exact ecology %D 1989 %T Butterfly-ant mutualisms %A Pierce, N. E. %B Towards a more exact ecology %I Blackwell %C Oxford %P 299-324 %G eng %0 Journal Article %J Oecologia %D 1989 %T The Effect of Adult Diet on the Biology of Butterflies .1. The Common Imperial Blue, Jalmenus-Evagoras %A Hill, C. J. %A Pierce, N. E. %B Oecologia %V 81 %P 249-257 %@ 0029-8549 %G English %M WOS:A1989AY34400016 %! OecologiaOecologia %0 Journal Article %J Oecologia %D 1988 %T Local-Distribution of the Lycaenid Butterfly, Jalmenus-Evagoras, in Response to Host Ants and Plants %A Smiley, J. T. %A Atsatt, P. R. %A Pierce, N. E. %B Oecologia %V 76 %P 416-422 %@ 0029-8549 %G English %M WOS:A1988P639200014 %! OecologiaOecologia %0 Book Section %B Reproductive success: studies of selection and adaptation in contrasting breeding systems. %D 1988 %T Mating success and fecundity in an ant-tended lycaenid butterfly %A Elgar, MA %A Pierce, N. E. %B Reproductive success: studies of selection and adaptation in contrasting breeding systems. %I Chicago University Press %C Chicago %P 59-75 %G eng %0 Book Section %B Reproductive success: studies of selection and adaptation in contrasting breeding systems. %D 1988 %T Mating success and fecundity in an ant-tended lycaenid butterfly %A Elgar, MA %A Pierce, N. E. %B Reproductive success: studies of selection and adaptation in contrasting breeding systems. %I Chicago University Press %C Chicago %P 59-75 %G eng %0 Journal Article %J Oecologia %D 1988 %T Local-Distribution of the Lycaenid Butterfly, Jalmenus-Evagoras, in Response to Host Ants and Plants %A Smiley, J. T. %A Atsatt, P. R. %A Pierce, N. E. %B Oecologia %V 76 %P 416-422 %@ 0029-8549 %G English %M WOS:A1988P639200014 %! OecologiaOecologia %0 Journal Article %J Behavioral Ecology and Sociobiology %D 1987 %T The Costs and Benefits of Cooperation between the Australian Lycaenid Butterfly, Jalmenus-Evagoras, and Its Attendant Ants %A Pierce, N. E. %A Kitching, R. L. %A Buckley, R. C. %A Taylor, M. F. J. %A Benbow, K. F. %B Behavioral Ecology and Sociobiology %V 21 %P 237-248 %@ 0340-5443 %G English %M WOS:A1987K428000006 %! Behav Ecol SociobiolBehav Ecol Sociobiol %0 Book Section %B Oxford Surveys in Evolutionary Biology %D 1987 %T The evolution and biogeography of associations between lycaenid butterflies and ants. %A Pierce, N. E. %X

Many organisms make a living from seratching each other's backs. and many survive at the expense of othcrs. Once a complex interaction has arisen between two organisms. what elfect can such a relationship have on their subsequcnt cvolution·! This pllpcr will consider thc evolutionary conscquences of ussocintions UIllOlIg Iycllcnid buttcrflics, their host plants, unts. purusitoids. lind prcdators. Thc Lycacllidac llfC cspecially intcrc.'1ting from an ccologicul and evolutionary pcrspcctivc because they exhibit dramatic variety in their life histories. The larvae of many species associate with ants, and these relationships can be parasitic. commensal, or mutualistic. larvae cun be carnivorous or hcrbivorous: and some species interact with many species of ants, whereas others are species-specific. It is partly because of this complexity and diversity that the lycaenidae have not been studied as intensively as other buUerny families, and I will discuss at least three problems that have hampered our understanding of their ecology and evolution. In particular. more must be learned about the nature of the exocrine secretions of lycaenid larvae, and whether they function to reward. appease, and/or deceive their associated ants. The association between lycaenids and ants has had several important evolutionary consequences, and I will show how these relate to the question of why there are so many species of Iycaenid buuernies. Finally. I will discuss an unresolved pattern in the biogeography oflycaenid buuernies: association with ants in general, and species-specific interactions in particular, are far more common among Iycaenids found in Ethiopian, Oriental. and Australasian regions than among those from the Holarctic. 

%B Oxford Surveys in Evolutionary Biology %I Oxford University Press %V IV %P 89-116 %G eng %0 Journal Article %J Nature %D 1987 %T No room for loners. Review of Animal Societies: Theories and Facts, Ito Y, JL Brown and J Kikkawa (eds.), Japan Scientific Societies Press. %A May, RM %A Pierce, N. E. %B Nature %V 330 %P 32 %G eng %0 Journal Article %J Insectarium %D 1987 %T The remarkable life histories of lycaenid butterflies. %A Pierce, N. E. %B Insectarium %V 24 %P 4-11 %G eng %0 Book Section %B Oxford Surveys in Evolutionary Biology %D 1987 %T The evolution and biogeography of associations between lycaenid butterflies and ants. %A Pierce, N. E. %X

Many organisms make a living from seratching each other's backs. and many survive at the expense of othcrs. Once a complex interaction has arisen between two organisms. what elfect can such a relationship have on their subsequcnt cvolution·! This pllpcr will consider thc evolutionary conscquences of ussocintions UIllOlIg Iycllcnid buttcrflics, their host plants, unts. purusitoids. lind prcdators. Thc Lycacllidac llfC cspecially intcrc.'1ting from an ccologicul and evolutionary pcrspcctivc because they exhibit dramatic variety in their life histories. The larvae of many species associate with ants, and these relationships can be parasitic. commensal, or mutualistic. larvae cun be carnivorous or hcrbivorous: and some species interact with many species of ants, whereas others are species-specific. It is partly because of this complexity and diversity that the lycaenidae have not been studied as intensively as other buUerny families, and I will discuss at least three problems that have hampered our understanding of their ecology and evolution. In particular. more must be learned about the nature of the exocrine secretions of lycaenid larvae, and whether they function to reward. appease, and/or deceive their associated ants. The association between lycaenids and ants has had several important evolutionary consequences, and I will show how these relate to the question of why there are so many species of Iycaenid buuernies. Finally. I will discuss an unresolved pattern in the biogeography oflycaenid buuernies: association with ants in general, and species-specific interactions in particular, are far more common among Iycaenids found in Ethiopian, Oriental. and Australasian regions than among those from the Holarctic. 

%B Oxford Surveys in Evolutionary Biology %I Oxford University Press %V IV %P 89-116 %G eng %0 Journal Article %J Nature %D 1987 %T No room for loners. Review of Animal Societies: Theories and Facts, Ito Y, JL Brown and J Kikkawa (eds.), Japan Scientific Societies Press. %A May, RM %A Pierce, N. E. %B Nature %V 330 %P 32 %G eng %0 Journal Article %J Insectarium %D 1987 %T The remarkable life histories of lycaenid butterflies. %A Pierce, N. E. %B Insectarium %V 24 %P 4-11 %G eng %0 Journal Article %J Behavioral Ecology and Sociobiology %D 1987 %T The Costs and Benefits of Cooperation between the Australian Lycaenid Butterfly, Jalmenus-Evagoras, and Its Attendant Ants %A Pierce, N. E. %A Kitching, R. L. %A Buckley, R. C. %A Taylor, M. F. J. %A Benbow, K. F. %B Behavioral Ecology and Sociobiology %V 21 %P 237-248 %@ 0340-5443 %G English %M WOS:A1987K428000006 %! Behav Ecol SociobiolBehav Ecol Sociobiol %0 Journal Article %J American Naturalist %D 1986 %T Lycaenid Butterflies and Ants - 2-Species Stable Equilibria in Mutualistic, Commensal, and Parasitic Interactions %A Pierce, N. E. %A Young, W. R. %B American Naturalist %V 128 %P 216-227 %8 Aug %@ 0003-0147 %G English %M WOS:A1986D610300005 %! Am NatAm Nat %0 Journal Article %J Journal of Animal Ecology %D 1986 %T The Selective Advantage of Attendant Ants for the Larvae of a Lycaenid Butterfly, Glaucopsyche-Lygdamus %A Pierce, N. E. %A Easteal, S. %B Journal of Animal Ecology %V 55 %P 451-462 %8 Jun %@ 0021-8790 %G English %M WOS:A1986C733400004 %! J Anim EcolJ Anim Ecol %0 Journal Article %J American Naturalist %D 1986 %T Lycaenid Butterflies and Ants - 2-Species Stable Equilibria in Mutualistic, Commensal, and Parasitic Interactions %A Pierce, N. E. %A Young, W. R. %B American Naturalist %V 128 %P 216-227 %8 Aug %@ 0003-0147 %G English %M WOS:A1986D610300005 %! Am NatAm Nat %0 Journal Article %J Journal of Animal Ecology %D 1986 %T The Selective Advantage of Attendant Ants for the Larvae of a Lycaenid Butterfly, Glaucopsyche-Lygdamus %A Pierce, N. E. %A Easteal, S. %B Journal of Animal Ecology %V 55 %P 451-462 %8 Jun %@ 0021-8790 %G English %M WOS:A1986C733400004 %! J Anim EcolJ Anim Ecol %0 Journal Article %J Behavioral Ecology and Sociobiology %D 1985 %T The Influence of Ants on Host Plant-Selection by Jalmenus-Evagoras, a Myrmecophilous Lycaenid Butterfly %A Pierce, N. E. %A Elgar, M. A. %B Behavioral Ecology and Sociobiology %V 16 %P 209-222 %@ 0340-5443 %G English %M WOS:A1985AHJ8900003 %! Behav Ecol SociobiolBehav Ecol Sociobiol %0 Journal Article %J American Naturalist %D 1985 %T Lycaenid Butterflies and Ants - Selection for Nitrogen-Fixing and Other Protein-Rich Food Plants %A Pierce, N. E. %B American Naturalist %V 125 %P 888-895 %@ 0003-0147 %G English %M WOS:A1985ASK0500010 %! Am NatAm Nat %0 Journal Article %J Behavioral Ecology and Sociobiology %D 1985 %T The Influence of Ants on Host Plant-Selection by Jalmenus-Evagoras, a Myrmecophilous Lycaenid Butterfly %A Pierce, N. E. %A Elgar, M. A. %B Behavioral Ecology and Sociobiology %V 16 %P 209-222 %@ 0340-5443 %G English %M WOS:A1985AHJ8900003 %! Behav Ecol SociobiolBehav Ecol Sociobiol %0 Journal Article %J American Naturalist %D 1985 %T Lycaenid Butterflies and Ants - Selection for Nitrogen-Fixing and Other Protein-Rich Food Plants %A Pierce, N. E. %B American Naturalist %V 125 %P 888-895 %@ 0003-0147 %G English %M WOS:A1985ASK0500010 %! Am NatAm Nat %0 Book Section %B Biology of butterflies. XI Symp R Entomol Soc (Lond). %D 1984 %T Amplified species diversity: a case study of an Australian lycaenid butterfly and its attendant ants. %A Pierce, N. E. %B Biology of butterflies. XI Symp R Entomol Soc (Lond). %I Academic Press %C London %P 197-200 %G eng %0 Book Section %B Biology of butterflies. XI Symp R Entomol Soc (Lond). %D 1984 %T Amplified species diversity: a case study of an Australian lycaenid butterfly and its attendant ants. %A Pierce, N. E. %B Biology of butterflies. XI Symp R Entomol Soc (Lond). %I Academic Press %C London %P 197-200 %G eng %0 Journal Article %J Science %D 1981 %T Parasitoids as Selective Agents in the Symbiosis between Lycaenid Butterfly Larvae and Ants %A Pierce, N. E. %A Mead, P. S. %B Science %V 211 %P 1185-1187 %@ 0036-8075 %G English %M WOS:A1981LE42500028 %! ScienceScience %0 Journal Article %J Science %D 1981 %T Parasitoids as Selective Agents in the Symbiosis between Lycaenid Butterfly Larvae and Ants %A Pierce, N. E. %A Mead, P. S. %B Science %V 211 %P 1185-1187 %@ 0036-8075 %G English %M WOS:A1981LE42500028 %! ScienceScience