Betsy Arnold

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Courses

Scholarly Contributions

Chapters

• Hamzazai, A., Lee, M., Garcia, K., Pat, E., Ibarra, A., Woytenko, E., & Arnold, A. E. (2019). Evaluating recruitment of soilborne microbes to seeds and their effects on seed germination of crop plants in agricultural and non-agricultural soils. In Handbook of Plant & Crop Physiology, 4th Edition. Taylor & Francis.
• Stump, S., Sarmiento, C., Zalamea, C., Dalling, J., Davis, A., Shaffer, J., & Arnold, A. E. (2019). Colonization of seeds by soilborne fungi: linking seed dormancy-defense syndromes, evolutionary constraints, and fungal traits.. In Seed Endophytes: Biotechnology and Biology. Springer.
• Baltrus, D. A., Spraker, J., & Arnold, A. E. (2018). Quantifying re-association of a facultative endohyphal bacterium with a filamentous fungus. In Plant Pathogenic Fungi and Oomycetes(pp 1-12). Humana.
• Stump, S. M., Sarmiento, C., Zalamea, P. C., Dalling, J. W., Davis, A. S., Shaffer, J. P., & Arnold, A. E. (2018). Colonization of seeds by soilborne fungi: linking seed dormancy-defense syndromes, evolutionary constraints, and fungal traits. In Seed Endophytes: Biotechnology and Biology(pp Pending). Springer.
• Van Bael, S., Estrada, C., & Arnold, A. (2017). Foliar endophyte communities and leaf traits in tropical trees. In The Fungal Community: its organization and role in the ecosystem(pp In press). Marcel-Dekker.

Journals/Publications

• Chen, K. H., Liao, H. L., Arnold, A. E., Korotkin, H. B., Wu, S. H., Matheny, P. B., & Lutzoni, F. (2021). Comparative transcriptomics of fungal endophytes in co-culture with their hosts reveals fungal trophic lability and neutral to slightly positive effects on moss growth rates. New Phytologist.
• Colon-Carrion, N., Lozada Troche, C., & Arnold, A. E. (2022). Responses of endophytic fungi in a Puerto Rican rainforest to damage from Hurricane Maria. Ecology and Evolution, See CV.
• Shaffer, J. P., Carter, M. E., Spraker, J. E., Clark, M., Smith, B., Hockett, K. C., Baltrus, D. A., & Arnold, A. E. (2021). Transcriptional profiles of a foliar fungal endophyte (Pestalotiopsis, Ascomycota) and its endohyphal bacterium (Luteibacter, Gammaproteobacteria) in co-culture support sulfur exchange and growth regulation.. mSystems.
• Tellez, P. H., Arnold, A. E., Leo, A. B., Kitajima, K., & VanBael, S. (2021). Traits along the leaf economics spectrum structure communities and traits of foliar endophytic symbionts. New Phytologist.
• U'Ren, J. M., Oita, S., Lutzoni, F., Miadlikowska, J., Ball, B., Carbone, I., Valles, D., Trouet, V., & Arnold, A. E. (2021). Climate drives endophyte diversity and hotspots at a trans-biome scale. Current Biology.
• Arnold, A. E. (2021). Endophytism in Pezizomycetes: the exception or the rule?. New Phytologist. doi:https://doi.org/10.1111/nph.17886
• Arnold, A. E., & Sandberg, D. C. (2021). Clohesyomyces symbioticus sp. nov., a fungal endophyte associated with roots of water smartweed (Persicaria amphibia). Plant and Fungal Systematics, 66, 201-210. doi:DOI: https://doi.org/10.35535/pfsyst-2021-0018
• Arnold, A. E., Harrington, A. H., Huang, Y., U'Ren, J. M., Massimo, N. C., Knight-Connoni, V., & Inderbitzin, P. (2021). Coniochaeta elegans sp. nov., C. montana sp. nov., and C. nivea sp. nov.: three new species of endophytes with distinctive morphology and functional traits.. International Journal of Systematic and Evolutionary Microbiology. doi:doi: 10.1099/ijsem.0.005003
• Arnold, A. E., Harrington, A. H., U'Ren, J. M., Oita, S., & Inderbitzin, P. (2021). Two new endophytic species enrich the Coniochaeta endophytica / C. prunicola clade: Coniochaeta palaoa sp. nov. and C. lutea sp. nov.. Plant and Fungal Systematics, 66, 66-78. doi:DOI: https://doi.org/10.35535/pfsyst-2021-0006
• Bowman, E., & Arnold, A. E. (2021). Drivers and implications of distance decay differ for ectomycorrhizal and foliar endophytic fungi across an anciently fragmented landscape.. The ISME Journal. doi:https://doi.org/10.1002/ecs2.3750
• Bowman, E., Hayden, D., & Arnold, A. E. (2021). Fire history and local factors shape ectomycorrhizal communities associated with Pinus ponderosa in mountains of the Madrean Sky Island Archipelago. Fungal Ecology, 49, e101013. doi:https://doi.org/10.1016/j.funeco.2020.101013
• Fehmi, J., Rasmussen, C., & Arnold, A. E. (2021). The pioneer effect advantage in plant invasions: site priming of native grasslands by invasive grasses. Ecosphere.
• Franco, M. E., Wisecaver, J. H., Arnold, A. E., Ju, Y. M., Slot, J. C., Ahrendt, S., Moore, L. P., Eastman, K. E., Scott, K., Konkel, Z., Mondo, S. J., Kuo, A., Hayes, R. D., Haridas, S., Andreopoulos, B., Riley, R., LaButti, K., Pangilinan, J., Lipzen, A., , Amirebrahimi, M., et al. (2021). Ecological generalism drives hyperdiversity of secondary metabolite gene clusters in xylarialean endophytes. New Phytologist, 233(3), 1317-1330.
• Geml, J., Arnold, A. E., Semenova-Nelsen, T. A., Nouhra, E. R., Drechsler-Santos, E. R., Góes-Neto, A., Morgado, L. N., Ódor, P., Hegyi, B., Grau, O., Ibáñez, A., Tedersoo, L., & Lutzoni, F. (2021). Community dynamics of soil-borne fungal communities along elevation gradients in neotropical and paleotropical forests. Molecular Ecology. doi:DOI: 10.1111/mec.16368
• Oita, S., Carey, J., Kline, I., Ibáñez, A., Yang, N., Hom, E. F., Carbone, I., U'Ren, J. M., & Arnold, A. E. (2021). Methodological approaches frame insights into endophyte richness and community composition. Microbial Ecology, 82(1), 21-34.
• Oita, S., Ibáñez, A., Lutzoni, F., Miadlikowska, J., Geml, J., Lewis, L. A., Hom, E. F., Carbone, I., U'Ren, J. M., & Arnold, A. E. (2021). Climate and seasonality drive the richness and composition of tropical fungal endophytes at a landscape scale. Communications Biology, 4(1), 313.
• Qu, W., Kithsiri Wijeratne, E. M., Bashyal, B. P., Xu, Y. M., Liu, M. X., Inacio, M. C., Arnold, A. E., U'Ren, J. M., & Gunatilaka, A. A. (2021). Strobiloscyphones A-F, 6-isopentylsphaeropsidones and other metabolites from Strobiloscypha sp. AZ0266, a leaf-associated fungus of Douglas fir. Journal of Natural Products, 84, 2575-2586. doi:DOI: 10.1021/acs.jnatprod.1c00662
• Zalamea, P., Sarmiento, C., Arnold, A. E., Davis, A., Ferrer, A., & Dalling, J. (2021). Closely related tree species support distinctive communities of seed-associated fungi in a lowland tropical forest.. Journal of Ecology, 109, 1858-1872. doi:https://doi.org/10.1111/1365-2745.13611
• Dalling, J. W., Davis, A. S., Arnold, A. E., Sarmiento, C., & Zalamea, P. (2020). Extending plant defense theory to seeds. Annual Reviews of Ecology, Evolution, and Systematics, 51, 123-141. doi:doi.org/10.1146/annurev-ecolsys-012120-115156
• Xu, Y. M., Arnold, A. E., U Ren, J. M., Xuan, L. J., Wang, W. Q., & Gunatilaka, A. A. (2020). Teratopyrones A-C, Dimeric Naphtho-γ-Pyrones and Other Metabolites from sp. AK1128, a Fungal Endophyte of. Molecules (Basel, Switzerland), 25(21).
  More info

  Bioassay-guided fractionation of a cytotoxic extract derived from a solid potato dextrose agar (PDA) culture of sp. AK1128, a fungal endophyte of , afforded three new naphtho-γ-pyrone dimers, teratopyrones A-C (-), together with five known naphtho-γ-pyrones, aurasperone B (), aurasperone C (), aurasperone F (), nigerasperone A (), and fonsecin B (), and two known diketopiperazines, asperazine () and isorugulosuvine (). The structures of - were determined on the basis of their spectroscopic data. Cytotoxicity assay revealed that nigerasperone A () was moderately active against the cancer cell lines PC-3M (human metastatic prostate cancer), NCI-H460 (human non-small cell lung cancer), SF-268 (human CNS glioma), and MCF-7 (human breast cancer), with ICs ranging from 2.37 to 4.12 μM while other metabolites exhibited no cytotoxic activity up to a concentration of 5.0 μM.
• de Amorim, M. R., Wijeratne, E. M., Zhou, S., Arnold, A. E., Batista, A. N., Batista, J. M., Dos Santos, L. C., & Gunatilaka, A. A. (2020). An epigenetic modifier induces production of 3-(4-oxopyrano)-chromen-2-ones in sp. AST0006, an endophytic fungus of. Tetrahedron, 76(43).
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  Incorporation of the epigenetic modifier suberoylanilide hydroxamic acid (SAHA) into a potato dextrose broth culture of the endophytic fungus sp. AST0006 affected its polyketide biosynthetic pathway providing two new 3-(4-oxopyrano)-chromen-2-ones, aspyranochromenones A () and B (), and the isocoumarin, (-)-6,7-dihydroxymellein (). Eight additional metabolites (-) and two biotransformation products of SAHA (-) were also encountered. The planar structures and relative configurations of the new metabolites - were elucidated with the help of high-resolution mass, 1D and 2D NMR spectroscopic data and the absolute configurations of - were determined by comparison of experimental and calculated ECD data. Possible biosynthetic pathways to and are presented.
• Arnold, A. E., Davidowitz, G., Moore, A., & von Arx, M. (2019). Diversity and distribution of microbial communities in floral nectar of two night-blooming plants of the Sonoran Desert. PloS one, 14(12), e0225309.
• Arnold, A. E., von Arx, M., Moore, A. F., & Davidowitz, G. (2019). Diversity and composition of cultivable microbial communities in nectar of moth-pollinated plants in the Sonoran Desert. PLoS ONE, 14, e0225309.
• Baltrus, D., Clark, M., Inderbitzin, P., Knight-Connoni, V., Pignatta, D., & Arnold, A. E. (2019). A complete genome sequence for Luteibacter pinisoli MAH-14. Microbiobiology Resource Announcements, 8, e00774-19.
• Carbone, I., White, J., Miadlikowska, J., Arnold, A. E., Miller, M., U'Ren, J., & Lutzoni, F. (2019). T-BAS version 2.1: Tree-based alignment selector toolkit for evolutionary placement and viewing of alignments and metadata on curated and custom reference trees. Microbiobiology Resource Announcements, 8, e00328-19.
• Harrington, A. H., Del Olmo-Ruiz, M., Sandberg, D. C., Hoffman, M. T., Huang, Y. L., U'Ren, J. M., & Arnold, A. E. (2019). Coniochaeta endophytica sp. nov., a foliar endophyte that showcases challenges in defining species boundaries, even with multiple lines of evidence. Fungal Systematics and Evolution, In review.
• Jaspers, C., Consortium, C. R., Arnold, A. E., Miller, D., Bosch, T., & Voolstra, C. (2019). Resolving metaorganism identities through a holistic framework combining reductionist and integrative approaches.. Zoology. doi:https://doi.org/10.1016/j.zool.2019.02.007 (cover).
• Lutzoni, F., Miadlikowska, J., Zimmerman, N., Carbone, I., May, G., & Arnold, A. E. (2019). Host availability drives distributions of fungal endophytes in the imperiled boreal realm. Nature Ecology & Evolution, 3, 1430-1437.
• Lutzoni, F., Uren, J. M., Magain, N., Miller, M. A., Arnold, A. E., Miadlikowska, J., White, J. B., & Carbone, I. (2019). T-BAS Version 2.1: Tree-Based Alignment Selector toolkit for evolutionary placement of DNA sequences and viewing alignments and specimen metadata on curated and custom trees. Microbiology Resource Announcements. doi:10.1128/MRA.00328-19
• McDade, L. A., Steidl, R. J., Arnold, A. E., Boyd, A. E., & McIntosh, M. E. (2019). Demography of a declining, endangered cactus in the Sonoran Desert. Plant Species Biology, 2109, 1-10. doi:https://doi.org/10.1111/1442-1984.12251
• McIntosh, M. E., Boyd, A. E., Arnold, A. E., Steidl, R. J., & McDade, L. A. (2019). Demography of a declining, endangered cactus in the Sonoran Desert. Plant Species Biology. doi:10.1111/1442-1984.12251
• Taylor, M., Mannan, R., U'Ren, J., Garber, N., Gallery, R., & Arnold, A. E. (2019). Age-related variation in the oral microbiome of urban Cooper’s hawks (Accipiter cooperii). BMC Microbiology, In press.
• Arnold, A. E., & Epps, M. (2018). Interaction networks of macrofungi and mycophagous beetles reflect diurnal variation and the size and spatial arrangement of resources. Fungal Ecology, 37, 48-56.
• Arnold, A. E., Baltrus, D. A., Davis, A. S., Dalling, J. W., Gallery, R. E., Sarmiento, C., Zalamea, P., & Justin, S. P. (2018). Context-dependent and variable effects of endohyphal bacteria on interactions between fungi and seeds. Fungal Ecology, 36, 117-127. doi:https://doi.org/10.1016/j.funeco.2018.08.008
• Arnold, A. E., Sandberg, D. C., Uren, J. M., Garber, N. P., Massimo, N., Bowman, E. A., & Huang, Y. (2018). Using collections data to infer biogeographic, environmental, and host structure in communities of endophytic fungi. Mycologia. doi:doi: 10.1080/00275514.2018.1442078.
• Bowman, E., & Arnold, A. E. (2018). Distributions of ectomycorrhizal and foliar endophytic fungal communities associated with Pinus ponderosa along a spatially constrained elevation gradient. American Journal of Botany, 105, 687-699.
• Chen, K., Liao, H., Arnold, A. E., & Lutzoni, F. (2018). Metatranscriptomic analysis of active fungal communities across a senescence gradient of the moss Dicranum scoparium. New Phytologist, 218, 1597-1611.
• Daru, B., Bowman, E. A., Pfister, D., & Arnold, A. E. (2018). A novel proof of concept for capturing the diversity of endophytic fungi preserved in herbarium specimens. Proceedings of the Royal Society B, 374, e1753. doi:10.1098/rstb.2017.0395
• Epps, M. J., & Arnold, A. E. (2018). Quantifying beetle-macrofungal associations in a temperate biodiversity hotspot. Mycologia, 110, 269-285.
• Gunatilaka, L., Arnold, A. E., Uren, J. M., Espinosa-Artiles, P., Wijeratne, E. K., Xu, Y., & Padumadasa, C. (2018). Cytotoxic and Other Metabolites from Teratosphaeria sp. FL2137, a fungus associated with foliage of Pinus clausa. Journal of Natural Products. doi:10.1021/acs.jnatprod.7b00838
• Huang, Y. L., Bowman, E. A., Massimo, N. C., Garber, N. P., U'Ren, J. M., Sandberg, D. C., & Arnold, A. E. (2018). Using collections data to infer biogeographic, environmental, and host structure in communities of endophytic fungi. Mycologia, 110, 47-62.
• Huang, Y. L., Zimmerman, N. B., & Arnold, A. E. (2018). Observation on the early establishment of foliar endophytic fungi in leaf discs and living leaves of a model woody angiosperm, Populus trichocarpa (Salicaceae). Journal of Fungi, 4, e58.
• Lutzoni, F., Nowak, M., Alfaro, M., Miadlikowska, J., Swofford, D., Arnold, A. E., Hibbett, D., Hilu, K., James, T., Quandt, D., & Magallon, S. (2018). Synchronized radiations in plants and fungi linked to symbiosis. Nature Communications, 9, e5451. doi:https://doi.org/10.1038/s41467-018-07849
• Padumadasa, C., Xu, Y., Wijeratne, E. K., Espinosa, P., U'Ren, J. M., Arnold, A. E., & Gunatilaka, A. (2018). Cytotoxic and non-cytotoxic metabolites from Teratosphaeria sp. FL2137, a fungus associated with Pinus clausa. Journal of Natural Products, 81, 616-624.
• Shaffer, J. P., Zalamea, P., Sarmiento, C., Gallery, R. E., Dalling, J. W., Baltrus, D. A., & Arnold, A. E. (2018). Context-dependent and variable effects of endohyphal bacteria on interactions between fungi and seeds. Fungal Ecology, 36, 117-127.
• Zalamea, P., Dalling, J. W., Sarmiento, C., Arnold, A. E., Delevich, C., Berbow, M. A., Ndobegang, A., Gripenberg, S., & Davis, A. S. (2018). Dormancy-defense syndromes and trade-offs between physical and chemical defenses in seeds. Ecology, 99, 1988-1998. doi:DOI: 10.1002/ecy.2419.
• Araldi-Brondolo, S. J., Spraker, J. S., Shaffer, J. P., Woytenko, E. H., Baltrus, D. A., Gallery, R. E., & Arnold, A. E. (2017). Bacterial endosymbionts: master modulators of fungal phenotypes. Microbiology Spectrum, 5, e0056.
• Arnold, A. E., Andersen, E. M., Taylor, M. J., & Steidl, R. J. (2017). Using cytochrome b to identify nests and museum specimens of cryptic songbirds. Conservation Genetics Resources, 9, 451-458.
• Arnold, A., & Del Olmo, M. (2017). Community structure of fern-affiliated endophytes in three neotropical forests. Journal of Tropical Ecology, 33, 60-73.
• Baltrus, D. A., Dougherty, K., Arendt, K., Huntemann, M., Clum, A., Pillay, M., Palaniappan, K., Vargese, N., Mikhailova, N., Stamatis, D., Reddy, T., Ngan, C. Y., Daum, C., Shapiro, N., Markowitz, V., Ivanova, N., Kyrpides, N., Woyke, T., & Arnold, A. E. (2017). Absence of genome reduction in diverse, facultative endohyphal bacteria. Microbial Genomics, 3, e000101.
• Bashyal, B., Kithsiri Wijeratne, E., Tillotson, J., Arnold, A. E., Chapman, E., & Gunatilaka, L. (2017). Chlorinated dehydrocurvularins and alterperylenepoxide A from Alternaria sp. AST0039, a fungal endophyte of Astragalus lentiginosus. Journal of Natural Products, 80, 427-433.
• Grubiani, J., Shi, T., Araujo, A., Arnold, A. E., Chapman, E., & Gunatilaka, A. (2017). An epigenetic modifier induces production of (10S)-verruculide B, an inhibitor of protein tyrosine phosphatases by Phoma sp. nov. LG219, a fungal endophyte of Parkinsonia microphylla. Bioorganic and Medicinal Chemistry, 25, 1860-1866.
• Luo, J., Xu, Y., Sandberg, D., Arnold, A. E., & Gunatilaka, L. (2017). Montagnuphilones A-G, azaphilones from Montagnulaceae sp. DM0194, a fungal endophyte of submerged roots of Persicaria amphibia. Journal of Natural Products, 80, 76-81.
• Sarmiento, C., Zalamea, P., Dalling, J. W., Davis, A. S., Stump, S. M., U'Ren, J. M., & Arnold, A. E. (2017). Soilborne fungi have host affinity and host-specific effects on seed germination and survival in a lowland tropical forest. Proceedings of the National Academy of Sciences USA, 114, 11458-11464.
• Shaffer, J. P., U'Ren, J. M., Baltrus, D. A., Gallery, R. E., & Arnold, A. E. (2017). An endohyphal bacterium (Chitinophaga, Bacteroidetes) influences carbon source use by Fusarium keratoplasticum (F. solani species complex, Nectriaceae). Frontiers in Microbiology, 8, e350.
• Torres-Cruz, T., Tobias, T., Almatruk, M., Hesse, C., Kuske, C., Desiro, A., Benucci, G., Bonito, G., Stajich, J., Dunlap, C., Arnold, A. E., & Porras-Alfaro, A. (2017). Bifiguratus adelaidae, gen. nov. sp. nov., a new lineage of Mucoromycotina with endophytic and soil-dwelling strains. Mycologia, 109, 363-378.
• Arendt, K., Hockett, K., Araldi-Brondolo, S., Baltrus, D., & Arnold, A. (2016). Isolation of endohyphal bacteria from foliar fungi and in vitro establishment of their symbiotic associations. Applied and Environmental Microbiology, 82, 2943-2499.
• Carbone, I., White, J., Miadlikowska, J., Arnold, A. E., Miller, M., Kauff, F., Schoch, C., U'Ren, J., May, G., & Lutzoni, F. (2016). Enhancing fungal species discovery and description using T-BAS: Tree-Based Alignment Selector toolkit for phylogenetic-based placement, alignment downloads, and metadata visualization. Bioinformatics, doi: 10.1093/bioinformatics/btw808.
• Chagnon, P., U'Ren, J., Lutzoni, F., Miadlikowska, J., & Arnold, A. (2016). Interaction type influences ecological network structure more than local abiotic conditions: evidence from fungal symbionts at a continental scale. Oecologia, 180, 181-191.
• Huang, Y., Devan, M., Furr, S., U'Ren, J., & Arnold, A. (2016). Pervasive effects of wildfire on foliar endophyte communities in montane forest trees. Microbial Ecology, 71, 452-458.
• Kithsiri Wijeratne, E., Gunaherath, G., Chapla, V., Tillotson, J., de la Cruz, F., Kang, M., U'Ren, J., Araujo, A., Arnold, A., Chapman, E., & Gunatilaka, A. (2016). Oxaspirol B with p97 inhibitory activity and other oxaspirols from Lecythophora sp. FL1375 and FL1031, endolichenic fungal strains inhabiting Parmotrema tinctorum and Cladonia evansii. Journal of Natural Products, 79, 340-352.
• Ohkura, M., Hughe-Hallett, J., Worley, J., fisher, J., Love, B., Arnold, A., & Orbach, M. (2016). Ophidiomyces ophiodiicola on a captive black racer (Coluber constrictor) and a garter snake (Thamnophis sirtalis) in Pennsylvania. Journal of Zoo and Wildlife Medicine, 47, 341-346.
• Shaffer, J. P., Sarmiento, C., Zalamea, P., Gallery, R. E., Davis, A. S., Baltrus, D. A., & Arnold, A. E. (2016). Diversity, Specificity, and Phylogenetic Relationships of Endohyphal Bacteria in Fungi that Inhabit Tropical Seeds and Leaves. Frontiers Ecology and Evolution. doi:https://doi.org/10.3389/fevo.2016.00116
• Sousa, J., Aguilar-Perez, M., Arnold, A., Rios, N., Kursar, T., Coley, P., & Cubilla-Rios, L. (2016). Chemical constituents and their antibacterial activity from the tropical endophytic fungus Diaporthe sp. F2934. Journal of Applied Microbiology, 120, 1501-1508.
• U'Ren, J., Miadlikowska, J., Zimmerman, N., Lutzoni, F., Stajich, J., & Arnold, A. (2016). Contributions of North American endophytes to the phylogeny, ecology, and taxonomy of the Xylariaceae.. Molecular Phylogenetics and Evolution, 98, 210-232.
• Arnold, A., Xu, Y., Bashyal, B., Liu, M., Espinosa-Artiles, P., U'Ren, J., & Gunatilaka, A. (2015). Cytotoxic cytochalasins and other metabolites from Xylariaceae sp. FL0390, a fungal endophyte of Spanish moss. Natural Products Communications, In press.
• Chen, K., Miadlikowska, J., Molnar, K., Arnold, A. E., Gaya, E., Gueidan, C., & Lutzoni, F. (2015). Phylogenetic analyses of eurotiomycetous endophytes reveal their close affinities to Chaetothyriales, Eurotiales, and a new order – Phaeomoniellales. Molecular Phylogenetics and Evolution, 85, 117-130.
• Corrales, A., Arnold, A. E., Ferrer, A., Turner, B., & Dalling, J. (2015). Variation in ectomycorrhizal communities associated with Oreomunnea mexicana (Juglandaceae) in tropical montane forests. Mycorrhiza, 26, 1-16.
• Martinson, E., Hackett, J., Machado, C., & Arnold, A. E. (2015). Metatranscriptome analysis of fig flowers suggests mechanisms for mutualism stability and gall induction. PLoS One, 10, e0130745.
• Massimo, N., Devan, M. N., Arendt, K., Wilch, M., Riddle, J., Furr, S., Steen, C., U'Ren, J., Sandberg, D., & Arnold, A. E. (2015). Fungal endophytes of desert plants: infrequent in culture, but diverse and distinctive symbionts. Microbial Ecology, 70, 61-76.
• Wei, H., Xu, Y., Espinosa-Artiles, P., Liu, M. X., Luo, J., U'Ren, J. M., Arnold, A. E., & Gunatilaka, A. A. (2015). Sesquiterpenes and other constituents of Xylaria sp NC1214, a fungal endophyte of the moss Hypnum sp.. PHYTOCHEMISTRY, 118, 102-108.
• Wijeratne, E. M., Xu, Y., Arnold, A. E., & Gunatilaka, A. A. (2015). Pulvinulin A, Graminin C, and cis-Gregatin B - New Natural Furanones from Pulvinula sp 11120, a Fungal Endophyte of Cupressus arizonica. NATURAL PRODUCT COMMUNICATIONS, 10(1), 107-111.
• Xu, Y., Mazefoli, J., Oliveira, M., U'Ren, J., Arnold, A., & Gunatilaka, A. (2015). Anteaglonialides A–F, Spironaphtho-1,8-dioxincyclohexa--butyrolactones and Palmarumycins CE1–CE3 from Anteaglonium sp. FL0768, a fungal endophyte of sand spikemoss, Selaginella arenicola. Journal of Natural Products, In press.
• Zalamea, C., Sarmiento, C., Arnold, A. E., Davis, A., & Dalling, J. (2015). Do microbes and abrasion by soil particles influence seed persistence and loss of physical dormancy in tropical seedbanks?. Frontiers in Plant Science, 5, e799. doi:doi: 10.3389/fpls.2014.00799
• Almeida, C., Ortega, H., Higginbotham, S., Spadafora, C., Arnold, A. E., Coley, P., Kursar, T., Gerwick, W., & Cubilla-Rios, L. (2014). Chemical and bioactive natural products from Microthyriaceae sp., an endophytic fungus from a tropical grass.. Letters in Applied Microbiology, 58, 59-64.
• Del Olmo Ruiz, M., & Arnold, A. E. (2014). Interannual variation and host affiliations of endophytic fungi associated with ferns at La Selva, Costa Rica.. Mycologia, 106, 8-21.
• Higginbotham, S., Wong, W., Linington, R., Spadafora, C., Iturrado, L., & Arnold, A. E. (2014). Sloth fur as a novel source of fungi with potent anti-parasitic and anti-bacterial activity. PLOS One, 9, e84549.
• Higgins, L., Arnold, A. E., Coley, P., & Kursar, T. (2014). Communities of fungal endophytes in tropical forest grasses: highly diverse host- and habitat generalists characterized by strong spatial structure. Fungal Ecology, 8, 1-11.
• Luo, J., Wang, X., Zu, Y., U'Ren, J., Arnold, A. E., Kong, L., & Gunatilaka, L. (2014). Delitschiapyrone A, a pyrone-naphthalenone adduct bearing an unprecedented pentacyclic ring system from the leaf-associated fungus Delitschia sp. FL1581. Organic Letters, 16, 5944-5947.
• Miadlikowska, J., Coauthors, 2., Arnold, A. E., Lutzoni, F., & Stenroos, S. (2014). Multigene phylogenetic analysis for 1307 fungi representing 1139 infrageneric taxa, 312 genera, and 66 families of the class Lecanoromycetes (Ascomycota). Molecular Phylogenetics and Evolution, 79, 132-168.
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  (Editor's choice)
• Nilsson, H., Co-authors, 6., Arnold, A. E., & Co-authors, 4. (2014). A distributed effort to improve annotation of ITS sequences from plant pathogenic fungi. Fungal Diversity, 67, 11-19.
• Oono, R., Lutzoni, F., Arnold, A. E., Kaye, L., U'Ren, J., May, G., & Carbone, I. (2014). Genetic variation in horizontally transmitted symbionts of pine needles reveals population structure in cryptic species. American Journal of Botany, 101, 1362-1374.
• Sandberg, D. C., Battista, L. J., & Arnold, A. E. (2014). Fungal endophytes of aquatic macrophytes: diverse host-generalists characterized by tissue preferences and geographic structure. Microbial Ecology, 67, 735-747.
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  Abstract: Most studies of endophytic symbionts have focused on terrestrial plants, neglecting the ecologically and economically important plants present in aquatic ecosystems. We evaluated the diversity, composition, host and tissue affiliations, and geographic structure of fungal endophytes associated with common aquatic plants in lentic waters in northern Arizona, USA. Endophytes were isolated in culture from roots and photosynthetic tissues during two growing seasons. A total of 226 isolates representing 60 putative species was recovered from 9,600 plant tissue segments. Although isolation frequency was low, endophytes were phylogenetically diverse and species-rich. Comparisons among the most thoroughly sampled species and reservoirs revealed that isolation frequency and diversity did not differ significantly between collection periods, among species, among reservoirs, or as a function of depth. However, community structure differed significantly among reservoirs and tissue types. Phylogenetic analyses of a focal genus (Penicillium) corroborated estimates of species boundaries and informed community analyses, highlighting clade- and genotype-level affiliations of aquatic endophytes with both sediment- and waterborne fungi, and endophytes of proximate terrestrial plants. Together these analyses provide a first quantitative examination of endophytic associations in roots and foliage of aquatic plants and can be used to optimize survey strategies for efficiently capturing fungal biodiversity at local and regional scales. © 2014 Springer Science+Business Media New York.
• U'Ren, J., Riddle, J., Monacell, J. T., Carbone, I., Miadlikowska, J., & Arnold, A. E. (2014). Tissue storage and primer selection influence pyrosequencing-based inferences of diversity and community composition of endolichenic and endophytic fungi. Molecular Ecology Resources, 14, 1032-1048.
• Higginbotham, S., Arnold, A. E., Ibanez, A., Spadafora, C., Coley, P., & Kursar, T. (2013). Bioactivity of fungal endophytes as a function of their taxonomy and the taxonomy and distribution of host plants.. PLoS One.
• Higginbotham, S., Wong, W., Linington, R., Spadafora, C., Iturrado, L., & Arnold, A. E. (2014). Sloth fur as a novel source of fungi with potent anti-parasitic and anti-bacterial activity.. PLoS One.
• Hoffman, M., Gunatilaka, M., Wijeratne, E., Gunatilaka, A. L., & Arnold, A. E. (2013). IAA production by endophytic Pestalotiopsis is enhanced by an endohyphal bacterium. PLoS One, 8, e73132.
• Lau, M. K., Arnold, A. E., & Johnson, N. C. (2013). Factors influencing communities of foliar fungal endophytes in riparian woody plants. Fungal Ecology, 6(5), 365-378.
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  Abstract: Riparian areas within a given arid region frequently contain broadly similar plant communities despite substantive geographic separation. Whether they also harbor similar communities of fungal symbionts, or feature assemblages unique to each riparian zone, is unknown. We examined fungal endophytes in foliage of woody angiosperms in six riparian areas in Arizona. Abundance and diversity differed among host species according to leaf longevity and phytochemistry, and among sites as a function of rainfall. Community composition varied among sites and host species. Comparison with regional data revealed that riparian areas harbor different subsets of the regional mycota rather than a consistent group of riparian taxa. Overall a high species- and phylogenetic richness of endophytes was recovered, especially among Mycosphaerella and affiliated anamorphs. Variation in endophyte communities across sites despite the relative consistency of plant communities underscores the importance of riparian zones both singly and in combination for harboring fungal biodiversity. © 2013 Elsevier Ltd and The British Mycological Society.
• Lau, M., Collins-Johnson, N., & Arnold, A. E. (2013). Fungal endophytes in riparian areas of Arizona illustrate effects of microenvironment on symbiont assemblages. Fungal Ecology, 6, 365-378.
• Martinson, E. O., Jandér, K. C., Peng, Y., Chen, H., Machado, C. A., Arnold, A. E., & Herre, E. A. (2013). Relative investment in egg load and poison sac in fig wasps: Implications for physiological mechanisms underlying seed and wasp production in figs. Acta Oecologica.
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  Abstract: Fig pollinating wasps and most non-pollinator wasps apply secretions from their poison sacs into oviposited flowers that appear necessary to the formation of the galls that their developing offspring consume. Thus, both eggs and poison sac secretions appear to be essential for wasp reproduction, but the relative investment in each is unknown. We measured relative investment in poison sac and egg production in pollinating and non-pollinating wasps associated with seven species of monoecious Panamanian figs representing both active and passive pollination syndromes. We then collected similar data for four fig hosts in China, where some wasp species in the genus Eupristina have lost the ability to pollinate ("cheaters"). All wasps examined possessed large poison sacs, and we found a strong positive correlation between poison sac size and absolute egg production. In the Panamanian species, the relative poison sac to egg investment was highest in the externally ovipositing non-pollinator wasps, followed by active pollinators, then by passive pollinators. Further, pollinator wasps of fig species with demonstrated host sanctions against "cheating" wasps showed higher investment in the poison sac than wasps of species without sanctions. In the Chinese samples, relative investment in the poison sac was indistinguishable between pollinators and "cheaters" associated with the same fig species. We suggest that higher relative investment in poison sac across fig wasp species reflects higher relative difficulty in initiating formation of galls and subsequently obtaining resources from the fig. We discuss the implications for the stability of the fig-wasp mutualism, and for the ability of non-pollinators to exploit this mutualism.
• Ortega, H. E., Graupner, P. R., Asai, Y., Tendyke, K., Qiu, D., Shen, Y. Y., Rios, N., Arnold, A. E., Coley, P. D., Kursar, T. A., Gerwick, W. H., & Cubilla-Rios, L. (2013). Mycoleptodiscins A and B, cytotoxic alkaloids from the endophytic fungus Mycoleptodiscus sp. F0194. Journal of Natural Products, 76(4), 741-744.
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  PMID: 23560689;Abstract: Two novel reddish-orange alkaloids, mycoleptodiscin A (1) and mycoleptodiscin B (2), were isolated from liquid cultures of the endophytic fungus Mycoleptodiscus sp. that had been isolated from Desmotes incomparabilis in Panama. Elucidation of their structures was accomplished using 1D and 2D NMR spectroscopy in combination with IR spectroscopic and MS data. These compounds are indole-terpenes with a new skeleton uncommon in nature. Mycoleptodiscin B (2) was active in inhibiting the growth of cancer cell lines with IC 50 values in the range 0.60-0.78 μM. © 2013 The American Chemical Society and American Society of Pharmacognosy.
• Ortega, H., Graupner, P. R., Asai, Y., TenDyke, D., Qiu, D. Y., Shen, Y. Y., Rios, N., Arnold, A. E., Coley, P. D., Kursar, T. A., Gerwick, W. H., & Cubilla-Rios, L. (2013). Mycoleptodiscins A and B, cytotoxic alkaloids from the endophytic fungus Mycoleptodiscus sp. F0194. Journal of Natural Products, 76, 741-744.
• Tedersoo, L., Arnold, A. E., & Hansen, K. (2013). Novel aspects in the life cycle and biotrophic interactions in Pezizomycetes (Ascomycota, Fungi). Molecular Ecology, 22(6), 1488-1493.
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  PMID: 23599958;Abstract: The ascomycete class Pezizomycetes (single order Pezizales) is known for its cup-shaped fruit bodies and the evolution of edible truffles and morels, but little is known about the ontogeny and ecology of this large and ecologically diverse fungal group. In this issue of Molecular Ecology, Healy et al. (2013) make a great leap forward by describing and identifying asexual, anamorphic structures that produce mitotic spores in many ectomycorrhiza-forming truffle and nontruffle species on soil surfaces worldwide (Fig.). Although such anamorphic forms have been reported sporadically from certain ectomycorrhizal and saprotrophic Pezizomycetes (e.g. Warcup), Healy et al. (2013) demonstrate that these terricolous asexual forms are both taxonomically and geographically more widespread and, in fact, much more common than previously understood. We anticipate that deeper insight into other substrates, provided by molecular analyses of materials such as dead wood and seeds, is likely to reveal numerous anamorphs of saprotrophic and pathogenic Pezizomycetes as well (see Marek et al.). © 2013 Blackwell Publishing Ltd.
• Tedersoo, L., Arnold, A. E., & Hansen, K. (2013). Novel aspects in the life cycle and biotrophic interactions in the Pezizomycetes. Molecular Ecology.
• Wanigesekara, A., Wijeratne, E. M., Arnold, A. E., & Gunatilaka, A. L. (2013). 10;-Deoxy-10 alpha-hydroxyascochlorin, a new cell migration inhibitor and other metabolites from Acremonium sp., a fungal endophyte in Ephedra trifurca. Natural Products Communications, 8, 601-604.
• Xu, Y. M., Espinosa-Artiles, P., Liu, M. X., Arnold, A. E., & Gunatilaka, A. L. (2013). Secoemestrin D, a cytotoxic epitetrathiodioxopiperizine, and Emericellenes A-E, five sesterterpenoids from Emericella sp AST0036, a fungal endophyte of Astragalus lentiginosus.. Journal of Natural Products, 76, 2330-2336.
• Arnold, A. E., Hoffman, M., Gunatilaka, M., Wijeratne, E., Gunatilaka, A., & Arnold, A. E. (2012). IAA production by endophytic Pestalotiopsis is enhanced by an endohyphal bacterium. PLoS ONE.
• Bascom-Slack, C. A., Arnold, A. E., & Strobel, S. A. (2012). Student-directed discovery of the plant microbiome and its products. Science, 338(6106), 485-486.
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  PMID: 23112324;Abstract: Endophyte Discovery, an IBI Prize-winning module, allows novice scientists to engage in meaningful discovery through inquiry-based research experiences.
• Bascom-Slack, C., Arnold, A., & Strobel, S. (2012). Student-directed discovery of the plant microbiome and its products. Science, 338, 485-486.
• Devan, M., Furr, S., & Arnold, A. (2012). Persistent effects of wildfire on fungal endophyte communities in Quercus hypoleucoides and Juniperus deppeana. Microbial Ecology.
• Gazis, R., Miadlikowska, J., Lutzoni, F., Arnold, A., & Chaverri, P. (2012). Culture-based study of endophytes associated with rubber trees in Peru reveals a new class of Pezizomycotina: Xylonomycetes. Molecular Phylogenetics and Evolution, 65, 294-304.
• Martinson, E., Herre, E., Machado, C., & Arnold, A. (2012). Culture-free survey reveals diverse fungal communities associated with figs (Ficus spp.) in Panama. Microbial Ecology, 64, 1073-1084.
• Martińez-Luis, S., Cherigo, L., Arnold, E., Spadafora, C., Gerwick, W. H., & Cubilla-Rios, L. (2012). Antiparasitic and anticancer constituents of the endophytic fungus Aspergillus sp. strain F1544. Natural Product Communications, 7(2), 165-168.
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  PMID: 22474943;Abstract: With the combined goal of finding the best anti-parasitic and anti-cancer activities as well as isolating the bioactive agents and studying their structures and biological properties, we proceeded to perform a small-scale cultivation of Aspergillus sp. strain F1544 using Potato Dextrose, Malt Extract, Czapek Dox and Eight Vegetables media. From the more promising extracts (obtaining using potato dextrose and czapek dox media in large scale) of this fungus, we isolated the five compounds: pseurotin A (1), 14-norpseurotin A (2), FD-838 (3), and pseurotin D (4), and fumoquinone B (5). All compounds showed good antileishmanial and moderate anticancer activities.
• Milani, N. A., Lawrence, D. P., Arnold, A. E., & D., H. (2012). Origin of pisatin demethylase (PDA) in the genus Fusarium. Fungal Genetics and Biology, 49(11), 933-942.
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  PMID: 22985693;Abstract: Host specificity of plant pathogens can be dictated by genes that enable pathogens to circumvent host defenses. Upon recognition of a pathogen, plants initiate defense responses that can include the production of antimicrobial compounds such as phytoalexins. The pea pathogen Nectria haematococca mating population VI (MPVI) is a filamentous ascomycete that contains a cluster of genes known as the pea pathogenicity (PEP) cluster in which the pisatin demethylase (PDA) gene resides. The PDA gene product is responsible for the detoxification of the phytoalexin pisatin, which is produced by the pea plant (Pisum sativum L.). This detoxification activity allows the pathogen to evade the phytoalexin defense mechanism. It has been proposed that the evolution of PDA and the PEP cluster reflects horizontal gene transfer (HGT). Previous observations consistent with this hypothesis include the location of the PEP cluster and PDA gene on a dispensable portion of the genome (a supernumerary chromosome), a phylogenetically discontinuous distribution of the cluster among closely related species, and a bias in G. +. C content and codon usage compared to other regions of the genome. In this study we compared the phylogenetic history of PDA, beta-tubulin, and translation elongation factor 1-alpha in three closely related fungi (Nectria haematococca, Fusarium oxysporum, and Neocosmospora species) to formally evaluate hypotheses regarding the origin and evolution of PDA. Our results, coupled with previous work, robustly demonstrate discordance between the gene genealogy of PDA and the organismal phylogeny of these species, and illustrate how HGT of pathogenicity genes can contribute to the expansion of host specificity in plant-pathogenic fungi. © 2012 Elsevier Inc.
• Milani, N., Lawrence, D., Arnold, A., & H.D., v. H. (2012). Origin of pistatin demethylase (PDA) in the genus Fusarium. Fungal Genetics and Biology, 49, 933-942..
• Molinar, E., Rios, N., Spadafora, C., Arnold, A. E., Coley, P. D., Kursar, T. A., Gerwick, W. H., & Cubilla-Rios, L. (2012). Coibanoles, a new class of meroterpenoids produced by Pycnoporus sanguineus. Tetrahedron Letters, 53(8), 919-922.
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  Abstract: Three new terpenoids of mixed biosynthetic origin were isolated from the culture filtrate of the endophytic fungus Pycnoporus sanguineus. Their structures were determined by extensive spectroscopic analyses. We have named these tricyclic and tetracyclic metabolites 'coibanoles A-C' in reference to Coiba Island and Coiba National Park, Panamá, from which the plant and endophyte were collected. The extract was inactive to the human parasites Trypanosoma cruzi, Leishmania donovani, and Plasmodium falciparum at a test concentration of 10 μg/mL. © 2011 Elsevier Ltd. All rights reserved.
• Molinar, E., Rios, N., Spadafora, C., Arnold, A., Coley, P., Kursar, T., Gerwick, W., & Cubilla-Rios, . L. (2012). Coibanoles, a new class of meroterpenoids produced by Pycnoporus sanguineus. Tetrahedron Letters.
• U, R. J., Lutzoni, F., Miadlikowska, J., Laetsch, A., & Arnold, A. (2012). Host and geographic structure of endophytic and endolichenic fungi at a continental scale. American Journal of Botany.
• Varughese, T., Rios, N., Higginbotham, S., Arnold, A. E., Coley, P. D., Kursar, T. A., Gerwick, W. H., & Rios, L. C. (2012). Antifungal depsidone metabolites from Cordyceps dipterigena, an endophytic fungus antagonistic to the phytopathogen Gibberella fujikuroi. Tetrahedron Letters, 53(13), 1624-1626.
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  Abstract: Among thirty four endophytic fungal strains screened for in vitro antagonism, the endophytic fungus Cordyceps dipterigena was found to strongly inhibit mycelial growth of the plant pathogenic fungus Gibberella fujikuroi. Two new depsidone metabolites, cordycepsidone A (1) and cordycepsidone B (2), were isolated from the PDA culture extract of C. dipterigena and identified as being responsible for the antifungal activity. Elucidation of their chemical structures was carried out using 1D and 2D NMR spectroscopy in combination with IR and MS spectroscopic data. Cordycepsidone A displayed strong and dose-dependent antifungal activity against the plant pathogenic fungus G. fujikuroi. The isolates were inactive in bioassays for malaria (Plasmodium falciparum), leishmaniasis (Leishmania donovani), Chagas's disease (Trypanosoma cruzi), and cytotoxicity at 10 μg/mL. The compounds were also found to be inactive against several bacterial strains at 50 μg/mL. © 2011 Elsevier Ltd. All rights reserved.
• Wijeratne, E., Bashyal, B., Liu, M., Rocha, D., Kamal, G., Gunaherath, B., U, R. J., Gunatilaka, M., Arnold, A., Whitesell, L., & Gunatilaka, A. (2012). Geopyxins A-E, ent-kaurane diterpenoids from endolichenic fungal strains, Geopyxis aff. majalis and Geopyxis sp. AZ0066: structure-activity relationships of geopyxins and their analogues. Journal of Natural Products.
• Dalling, J. W., Davis, A. S., Schutte, B. J., & Arnold, A. E. (2011). Seed survival in soil: Interacting effects of predation, dormancy and the soil microbial community. Journal of Ecology, 99(1), 89-95.
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  Abstract: Plant defence theory provides a robust framework for understanding interactions between plants and antagonists, and for interpreting broad patterns in the functional-trait composition of plant communities. However, this framework has been built almost entirely on traits expressed by seedlings and mature plants. No equivalent seed defence theory exists that recognizes the distinct suite of natural enemies that seeds encounter, and the unique constraints to their response. Furthermore, most attention has been paid to insect and vertebrate seed predators active above ground, whereas microbes in soil also have large effects on seed survival, particularly for plants that recruit from soil seed banks. We suggest that concurrent selection on seed dormancy and resistance to microbial antagonists should result in distinct seed defence syndromes. We predict that species with physical seed dormancy will rely on physical defences to exclude predators and pathogens, and rapid seed germination to escape pathogens at the emergence stage. In contrast, species with physiological seed dormancy will deploy a continuum of physical and chemical defences, depending on soil pathogen pressure and duration of seed persistence. Finally, seeds of some species persist in the soil in a non-dormant, imbibed state, and lack obvious chemical and physical defences. These seeds may be especially dependent upon protection from beneficial seed-inhabiting microbes. Framing a general 'seed defence theory' may help to account for the distribution of seed dormancy types across ecosystems. We predict that physiological dormancy will be favoured in dry or well-drained environments where pathogen pressure is relatively low, germination cues are most unpredictable, and seedling recruitment success is most variable. In contrast, physical dormancy should be favoured in warm and moist environments where pathogen pressure is high, and where germination cues are a stronger predictor of recruitment success. Persistent, non-dormant seeds are restricted to relatively aseasonal environments where favourable conditions for recruitment can occur over most of the year. Synthesis. Integrating seed defence and dormancy traits can provide new insights into selection on dormancy types, and will help elucidate major trends in seed ecology and evolution. Understanding how seeds are defended also may improve our ability to predict plant regeneration and help develop innovative management strategies for weedy and invasive species. © 2010 The Authors. Journal of Ecology © 2010 British Ecological Society.
• Higgins, K. L., Coley, P. D., Kursar, T. A., & Arnold, A. E. (2011). Culturing and direct PCR suggest prevalent host generalism among diverse fungal endophytes of tropical forest grasses. Mycologia, 103(2), 247-260.
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  PMID: 20943570;Abstract: Most studies examining endophytic fungi associated with grasses (Poaceae) have focused on agronomically important species in managed ecosystems or on wild grasses in subtropical, temperate and boreal grasslands. However grasses first arose in tropical forests, where they remain a significant and diverse component of understory and forest-edge communities. To provide a broader context for understanding grass-endophyte associations we characterized fungal endophyte communities inhabiting foliage of 11 species of phylogenetically diverse C3 grasses in the understory of a lowland tropical forest at Barro Colorado Island, Panama. Our sample included members of early-arising subfamilies of Poaceae that are endemic to forests, as well as more recently arising subfamilies that transitioned to open environments. Isolation on culture media and direct PCR and cloning revealed that these grasses harbor species-rich and phylogenetically diverse communities that lack the endophytic Clavicipitaceae known from diverse woodland and pasture grasses in the temperate zone. Both the incidence and diversity of endophytes was consistent among grass species regardless of subfamily, clade affiliation or ancestral habitat use. Genotype and phylogenetic analyses suggest that these endophytic fungi are predominantly host generalists, shared not only among distinctive lineages of Poaceae but also with non-grass plants at the same site. © 2011 by The Mycological Society of America.
• Lawrence, D. P., Kroken, S., Pryor, B. M., & Arnold, A. E. (2011). Interkingdom gene transfer of a hybrid NPS/PKS from bacteria to filamentous ascomycota. PLoS ONE, 6(11).
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  PMID: 22140558;PMCID: PMC3226686;Abstract: Nonribosomal peptides (NRPs) and polyketides (PKs) are ecologically important secondary metabolites produced by bacteria and fungi using multidomain enzymes called nonribosomal peptide synthetases (NRPSs) and polyketide synthases (PKSs), respectively. Previous phylogenetic analyses of fungal NRPSs and PKSs have suggested that a few of these genes were acquired by fungi via horizontal gene transfer (HGT) from bacteria, including a hybrid NPS/PKS found in Cochliobolus heterostrophus (Dothideomycetes, Ascomycota). Here, we identify this hybrid gene in fungi representing two additional classes of Ascomycota (Aspergillus spp., Microsporum canis, Arthroderma spp., and Trichophyton spp., Eurotiomycetes; Chaetomium spp. and Metarhizium spp., Sordariomycetes) and use phylogenetic analyses of the most highly conserved domains from NRPSs (adenylation (A) domain) and PKSs (ketoacyl synthase (KS) domain) to examine the hypothesis that the hybrid NPS7/PKS24 was acquired by fungi from bacteria via HGT relatively early in the evolution of the Pezizomycotina. Our results reveal a unique ancestry of the A domain and KS domain in the hybrid gene relative to known fungal NRPSs and PKSs, provide strong evidence for HGT of the hybrid gene from a putative bacterial donor in the Burkholderiales, and suggest the HGT event occurred early in the evolution of the filamentous Ascomycota. © 2011 Lawrence et al.
• Martínez-Luis, S., Cherigo, L., Higginbotham, S., Arnold, E., Spadafora, C., Ibañez, A., Gerwick, W. H., & Cubilla-Rios, L. (2011). Screening and evaluation of antiparasitic and in vitro anticancer activities of Panamanian endophytic fungi. International Microbiology, 14(2), 95-102.
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  PMID: 22069153;PMCID: PMC3375117;Abstract: Many compounds produced by fungi have relevant pharmaceutical applications. The purpose of this study was to collect and isolate entophytic fungi from different regions of Panama and then to test their potential therapeutic activities against Leishmania donovani, Plasmodium falciparum, and Trypanosoma cruzi as well as their anticancer activities in MCF-7 cells. Of the 25 fungal isolates obtained, ten of them had good anti-parasitic potential, showing selective activity against L. donovani; four had significant anti-malarial activity; and three inhibited the growth of T. cruzi. Anticancer activity was demonstrated in four isolates. Of the active isolates, Edenia sp. strain F0755, Xylaria sp. strain F1220, Aspergillus sp. strain F1544, Mycoleptodiscus sp. strain F0194, Phomopsis sp. strain F1566, Pycnoporus sp. strain F0305, and Diaporthe sp. strain F1647 showed the most promise based on their selective bioactivity and lack of toxicity in the assays.
• Moreno, E., Varughese, T., Spadafora, C., Arnold, A., Coley, P., Kursar, T., Gerwick, W., & Cubilla-Rios, L. (2011). Chemical constituents of the new endophytic fungus Mycosphaerella sp. nov. and their anti-parasitic activity. Natural Product Communications, 6, 1-6.
• Wang, X., Bashyal, B., Wijeratne, E., U, R. J., Gunatilaka, M., Arnold, A., & Gunatilaka, A. (2011). Smardaesidins A-G, new isopimarane and 20-Nor-isopimarane diterpenoids isolated from Smardaea sp., endophyte of the moss Ceratodon purpureus. Journal of Natural Products.
• Arnold, A. E., Lamit, L. J., Gehring, C. A., Bidartondo, M. I., & Callahan, H. (2010). Interwoven branches of the plant and fungal trees of life. New Phytologist, 185(4), 874-878.
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  PMID: 20356341;
• Caballero-George, C., Bolaños, J., Ochoa, E., Carballo, J. L., Cruz, J. A., & Arnold, A. E. (2010). Protocol to isolate sponge-associated fungi from tropical waters and an examination of their cardioprotective potential. Current Trends in Biotechnology and Pharmacy, 4(4), 881-899.
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  Abstract: Fungi represent an essential component of biodiversity not only because of their high species richness at multiple scales, but also because of their ecological, evolutionary and socio-economic significance. Despite poorly understood natural history and uncertain estimates of diversity, marine fungi have been identified as a major source of new natural products with pharmacological applications. The aims of this study were (1) to characterize fungi associated with marine sponges in protected areas of the Pacific and Caribbean coasts of Panama, and (2) to examine their effects through radioligand binding assays on endothelin ETA (ETA) and neuropeptide Y Y1 (Y1) receptors, which provide an indication of cardioprotective potential. A total of 369 marine sponges were collected in areas of high biodiversity along the Panamanian coasts, including 156 from the western Caribbean and 213 from the eastern Pacific. From these, 2,747 and 2,263 fungal isolates were recovered, respectively, with variable isolation frequencies when sponge fragments were cultivated on five media. After determining the seasonality, geographic stucture, and taxonomic diversity of these fungal assemblages, we identified five strains that inhibited by > 50% the binding of [3H] BQ-123 and one the binding of [3H] neuropeptide Y to the ETA and Y1 receptors, respectively, at 100 μg/ml. Further studies are required to determine whether these interactions are agonistic or antagonistic. Drawing from our methods for isolating and screening these fungi we propose a general protocol for capturing, cataloguing, and assessing the pharmacological potential of previously undiscovered fungi associated with marine sponges.
• Epps, M. J., & Arnold, A. E. (2010). Diversity, abundance and community network structure in sporocarp-associated beetle communities of the central Appalachian Mountains. Mycologia, 102(4), 785-802.
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  PMID: 20648747;Abstract: Although arthropods are abundant and diverse in and on macrofungal sporocarps, their associations with fungi seldom have been described at a community level. We examined sporocarpassociated beetle communities in two primary sites in the Appalachian Mountains and foothills, assessing beetle diversity and abundance in relation to study site, sampling season (early vs. late summer), and sporocarp characteristics such as taxonomic position, dry mass and age. From 758 sporocarps representing.tdthomgt;180 species we recovered 15 404 adult beetles representing 72 species and 15 families, primarily Staphylinidae (tdthomgt;98% of individuals and of 64% morphospecies). The probability of sporocarp colonization by beetles, beetle abundance and diversity differed among fungal species and were positively associated with sporocarp dry mass. Sporocarp age was positively correlated with beetle diversity and abundance (as measured in a focal species, Megacollybia platyphylla, Tricholomataceae), and its effects were independent of dry mass. Many beetle species were generalists, visiting a wide breadth of fungi in both the Agaricales and Polyporales; however, several beetle taxa showed evidence of specialization on particular fungal hosts. Host association data were used to examine the structure underlying sporocarpbeetle associations. Here we present the first evidence of nested community structure in the sporocarpbeetle interaction network. © 2010 by The Mycological Society of America.
• Hoffman, M. T., & Arnold, A. E. (2010). Diverse bacteria inhabit living hyphae of phylogenetically diverse fungal endophytes. Applied and Environmental Microbiology, 76(12), 4063-4075.
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  PMID: 20435775;PMCID: PMC2893488;Abstract: Both the establishment and outcomes of plant-fungus symbioses can be influenced by abiotic factors, the interplay of fungal and plant genotypes, and additional microbes associated with fungal mycelia. Recently bacterial endosymbionts were documented in soilborne Glomeromycota and Mucoromycotina and in at least one species each of mycorrhizal Basidiomycota and Ascomycota. Here we show for the first time that phylogenetically diverse endohyphal bacteria occur in living hyphae of diverse foliar endophytes, including representatives of four classes of Ascomycota. We examined 414 isolates of endophytic fungi, isolated from photosynthetic tissues of six species of cupressaceous trees in five biogeographic provinces, for endohyphal bacteria using microscopy and molecular techniques. Viable bacteria were observed within living hyphae of endophytic Pezizomycetes, Dothideomycetes, Eurotiomycetes, and Sordariomycetes from all tree species and biotic regions surveyed. A focus on 29 fungus/bacterium associations revealed that bacterial and fungal phylogenies were incongruent with each other and with taxonomie relationships of host plants. Overall, eight families and 15 distinct genotypes of endohyphal bacteria were recovered; most were members of the Proteobacteria, but a small number of Bacillaceae also were found, including one that appears to occur as an endophyte of plants. Frequent loss of bacteria following subculturing suggests a facultative association. Our study recovered distinct lineages of endohyphal bacteria relative to previous studies, is the first to document their occurrence in foliar endophytes representing four of the most species-rich classes of fungi, and highlights for the first time their diversity and phylogenetic relationships with regard both to the endophytes they inhabit and the plants in which these endophyte-bacterium symbiota occur. Copyright © 2010, American Society for Microbiology. All Rights Reserved.
• Parrent, J. L., Peay, K., Arnold, A. E., Comas, L. H., Avis, P., & Tuininga, A. (2010). Moving from pattern to process in fungal symbioses: Linking functional traits, community ecology and phylogenetics. New Phytologist, 185(4), 882-886.
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  PMID: 20356343;
• Peay, K. G., Bidartondo, M. I., & Arnold, A. E. (2010). Not every fungus is everywhere: Scaling to the biogeography of fungal-plant interactions across roots, shoots and ecosystems. New Phytologist, 185(4), 878-882.
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  PMID: 20356342;
• U'Ren, J. M., Lutzoni, F., Miadlikowska, J., & Arnold, A. E. (2010). Community Analysis Reveals Close Affinities Between Endophytic and Endolichenic Fungi in Mosses and Lichens. Microbial Ecology, 60(2), 340-353.
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  PMID: 20625714;Abstract: Endolichenic fungi live in close association with algal photobionts inside asymptomatic lichen thalli and resemble fungal endophytes of plants in terms of taxonomy, diversity, transmission mode, and evolutionary history. This similarity has led to uncertainty regarding the distinctiveness of endolichenic fungi compared with endophytes. Here, we evaluate whether these fungi represent distinct ecological guilds or a single guild of flexible symbiotrophs capable of colonizing plants or lichens indiscriminately. Culturable fungi were sampled exhaustively from replicate sets of phylogenetically diverse plants and lichens in three microsites in a montane forest in southeastern Arizona (USA). Intensive sampling combined with a small spatial scale permitted us to decouple spatial heterogeneity from host association and to sample communities from living leaves, dead leaves, and lichen thalli to statistical completion. Characterization using data from the nuclear ribosomal internal transcribed spacer and partial large subunit (ITS-LSU rDNA) provided a first estimation of host and substrate use for 960 isolates representing five classes and approximately 16 orders, 32 families, and 65 genera of Pezizomycotina. We found that fungal communities differ at a broad taxonomic level as a function of the phylogenetic placement of their plant or lichen hosts. Endolichenic fungal assemblages differed as a function of lichen taxonomy, rather than substrate, growth form, or photobiont. In plants, fungal communities were structured more by plant lineage than by the living vs. senescent status of the leaf. We found no evidence that endolichenic fungi are saprotrophic fungi that have been "entrapped" by lichen thalli. Instead, our study reveals the distinctiveness of endolichenic communities relative to those in living and dead plant tissues, with one notable exception: we identify, for the first time, an ecologically flexible group of symbionts that occurs both as endolichenic fungi and as endophytes of mosses. © 2010 Springer Science+Business Media, LLC.
• Vega, F. E., Simpkins, A., Aime, M. C., Posada, F., Peterson, S. W., Rehner, S. A., Infante, F., Castillo, A., & Arnold, A. E. (2010). Fungal endophyte diversity in coffee plants from Colombia, Hawai'i, Mexico and Puerto Rico. Fungal Ecology, 3(3), 122-138.
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  Abstract: Coffee (Coffea arabica) plant tissues were surface-sterilized and fungal endophytes isolated using standard techniques, followed by DNA extraction and sequencing of the internal transcribed spacer region (ITS). A total of 843 fungal isolates were recovered and sequenced (Colombia, 267; Hawai'i, 393; Mexico, 109; Puerto Rico, 74) yielding 257 unique ITS genotypes (Colombia, 113; Hawai'i, 126; Mexico, 32; Puerto Rico, 40). The most abundant taxa were Colletotrichum, Fusarium, Penicillium, and Xylariaceae. Overall, 220 genotypes were detected in only one of the countries sampled; only two genotypes were found in all four countries. Endophytes were also isolated from Coffea canephora, Coffea congensis, Coffea liberica, Coffea macrocarpa, Coffea racemosa, and Coffea stenophylla in Hawai'i. The high biodiversity of fungal endophytes in coffee plants may indicate that most of these are " accidental tourists" with no role in the plant, in contrast to endophytes that could be defined as " influential passengers" and whose role in the plant has been elucidated. This study, the most comprehensive analysis of fungal endophytes associated with a single host species, demonstrates that coffee plants serve as a reservoir for a wide variety of fungal endophytes that can be isolated from various plant tissues, including the seed, and illustrates the different fungal communities encountered by C. arabica in different coffee-growing regions of the world. © 2009.
• Arnold, A. E., Miadlikowska, J., Higgins, K. L., Sarvate, S. D., Gugger, P., Way, A., Hofstetter, V., Kauff, F., & Lutzoni, F. (2009). A phylogenetic estimation of trophic transition networks for ascomycetous Fungi: Are lichens cradles of symbiotrophic Fungal diversification?. Systematic Biology, 58(3), 283-297.
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  PMID: 20525584;Abstract: Fungi associated with photosynthetic organisms are major determinants of terrestrial biomass, nutrient cycling, and ecosystem productivity from the poles to the equator. Whereas most fungi are known because of their fruit bodies (e.g., saprotrophs), symptoms (e.g., pathogens), or emergent properties as symbionts (e.g., lichens), the majority of fungal diversity is thought to occur among species that rarely manifest their presence with visual cues on their substrate (e.g., the apparently hyperdiverse fungal endophytes associated with foliage of plants). Fungal endophytes are ubiquitous among all lineages of land plants and live within overtly healthy tissues without causing disease, but the evolutionary origins of these highly diverse symbionts have not been explored. Here, we show that a key to understanding both the evolution of endophytism and the diversification of the most species-rich phylum of Fungi (Ascomycota) lies in endophyte-like fungi that can be isolated from the interior of apparently healthy lichens. These "endolichenic" fungi are distinct from lichen mycobionts or any other previously recognized fungal associates of lichens, represent the same major lineages of Ascomycota as do endophytes, largely parallel the high diversity of endophytes from the arctic to the tropics, and preferentially associate with green algal photobionts in lichen thalli. Using phylogenetic analyses that incorporate these newly recovered fungi and ancestral state reconstructions that take into account phylogenetic uncertainty, we show that endolichenism is an incubator for the evolution of endophytism. In turn, endophytism is evolutionarily transient, with endophytic lineages frequently transitioning to and from pathogenicity. Although symbiotrophic lineages frequently give rise to free-living saprotrophs, reversions to symbiosis are rare. Together, these results provide the basis for estimating trophic transition networks in the Ascomycota and provide a first set of hypotheses regarding the evolution of symbiotrophy and saprotrophy in the most species-rich fungal phylum. Copyright © Society of Systematic Biologists 2009.
• Horton, T. R., Arnold, A. E., & Bruns, T. D. (2009). FESIN workshops at ESA - The mycelial network grows. Mycorrhiza, 19(4), 283-285.
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  PMID: 19002507;
• Maddison, D. R., & Arnold, A. E. (2009). A review of the Bembidion (Odontium) aenulum subgroup (Coleoptera: Carabidae), with description of a new species. Zootaxa, 45-61.
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  Abstract: The Bembidion aenulum subgroup of the subgenus Odontium is reviewed. This subgroup, widespread in the United States east of the Rocky Mountains, previously has been considered to consist of only one species, Bembidion aenulum Hayward. However, morphological studies coupled with analyses of 28S ribosomal RNA, cytochrome oxidase I, CAD, and wingless genes reveal that eastern members of this group belong to a new species herein named Bembidion paraenulum new species. This species ranges from New Hampshire in the northeast, through Virginia and North Carolina, south to Florida and west to Mississippi. Both species are described and illustrated. Copyright © 2009 Magnolia Press.
• Rodriguez, R. J., White Jr, J. F., Arnold, A. E., & Redman, R. S. (2009). Fungal endophytes: Diversity and functional roles: Tansley review. New Phytologist, 182(2), 314-330.
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  PMID: 19236579;Abstract: All plants in natural ecosystems appear to be symbiotic with fungal endophytes. This highly diverse group of fungi can have profound impacts on plant communities through increasing fitness by conferring abiotic and biotic stress tolerance, increasing biomass and decreasing water consumption, or decreasing fitness by altering resource allocation. Despite more than 100 yr of research resulting in thousands of journal articles, the ecological significance of these fungi remains poorly characterized. Historically, two endophytic groups (clavicipitaceous (C) and nonclavicipitaceous (NC)) have been discriminated based on phylogeny and life history traits. Here, we show that NC-endophytes represent three distinct functional groups based on host colonization and transmission, in planta biodiversity and fitness benefits conferred to hosts. Using this framework, we contrast the life histories, interactions with hosts and potential roles in plant ecophysiology of C- and NC-endophytes, and highlight several key questions for future work in endophyte biology. © 2009 New Phytologist.
• U'Ren, J. M., Dalling, J. W., Gallery, R. E., Maddison, D. R., Davis, E. C., Gibson, C. M., & Arnold, A. E. (2009). Diversity and evolutionary origins of fungi associated with seeds of a neotropical pioneer tree: a case study for analysing fungal environmental samples. Mycological Research, 113(4), 432-449.
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  PMID: 19103288;Abstract: Fungi associated with seeds of tropical trees pervasively affect seed survival and germination, and thus are an important, but understudied, component of forest ecology. Here, we examine the diversity and evolutionary origins of fungi isolated from seeds of an important pioneer tree (Cecropia insignis, Cecropiaceae) following burial in soil for five months in a tropical moist forest in Panama. Our approach, which relied on molecular sequence data because most isolates did not sporulate in culture, provides an opportunity to evaluate several methods currently used to analyse environmental samples of fungi. First, intra- and interspecific divergence were estimated for the nu-rITS and 5.8S gene for four genera of Ascomycota that are commonly recovered from seeds. Using these values we estimated species boundaries for 527 isolates, showing that seed-associated fungi are highly diverse, horizontally transmitted, and genotypically congruent with some foliar endophytes from the same site. We then examined methods for inferring the taxonomic placement and phylogenetic relationships of these fungi, evaluating the effects of manual versus automated alignment, model selection, and inference methods, as well as the quality of BLAST-based identification using GenBank. We found that common methods such as neighbor-joining and Bayesian inference differ in their sensitivity to alignment methods; analyses of particular fungal genera differ in their sensitivity to alignments; and numerous and sometimes intricate disparities exist between BLAST-based versus phylogeny-based identification methods. Lastly, we used our most robust methods to infer phylogenetic relationships of seed-associated fungi in four focal genera, and reconstructed ancestral states to generate preliminary hypotheses regarding the evolutionary origins of this guild. Our results illustrate the dynamic evolutionary relationships among endophytic fungi, pathogens, and seed-associated fungi, and the apparent evolutionary distinctiveness of saprotrophs. Our study also elucidates the diversity, taxonomy, and ecology of an important group of plant-associated fungi and highlights some of the advantages and challenges inherent in the use of ITS data for environmental sampling of fungi. © 2008 The British Mycological Society.
• Bruns, T. D., Arnold, A. E., & Hughes, K. W. (2008). Fungal networks made of humans: UNITE, FESIN, and frontiers in fungal ecology. New Phytologist, 177(3), 586-588.
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  PMID: 18211473;
• Bruns, T. D., Blackwell, M., Edwards, I., F., A., Horton, T., Zhang, N., Kõljalg, U., May, G., Kuyper, T. W., Bever, J. D., Gilbert, G., Taylor, J. W., DeSantis, T. Z., Pringle, A., Borneman, J., Thorn, G., Berbee, M., Mueller, G. M., Andersen, G. L., , Vellinga, E. C., et al. (2008). Preserving accuracy in GenBank. Science, 319(5870), 1616-.
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  PMID: 18356505;
• Feldman, T. S., O'Brien, H. E., & Arnold, A. E. (2008). Moths that vector a plant pathogen also transport endophytic fungi and mycoparasitic antagonists. Microbial Ecology, 56(4), 742-750.
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  PMID: 18491176;Abstract: Claviceps paspali, a common fungal pathogen of Paspalum grasses, attracts moth vectors by producing sugary exudates in the grass florets it infects. These exudates also support mycoparasitic Fusarium species that may negatively influence C. paspali fitness. We examined the potential for moths on which C. paspali depends to also transmit mycoparasitic Fusarium and fungal endophytes, which inhabit asymptomatic plant tissue and may influence host susceptibility to pathogens. We quantified infections by C. paspali, Fusarium spp., and endophytic fungi associated with Paspalum spp. at focal sites in the southeastern USA and used data from the nuclear internal transcribed spacer (ITS rDNA) to compare communities of plant-associated and moth-borne fungi. ITS sequences of moth-borne fungi were identical to reference sequences of mycoparasitic Fusarium heterosporum and to three distinct endophytic fungi isolated from Paspalum species. Our results demonstrate an unexpected overlap of fungal communities between disparate locations and among plant species and plant tissues, and suggest an unexpected role of moths, which vector a plant pathogen, to transmit other guilds of fungi. In turn, the potential for insects to transmit plant pathogens as well as mycoparasites and endophytic fungi suggests complex interactions underlying a commonly observed grass-pathogen system. © 2008 Springer Science+Business Media, LLC.
• Hoffman, M. T., & Arnold, A. E. (2008). Geographic locality and host identity shape fungal endophyte communities in cupressaceous trees. Mycological Research, 112(3), 331-344.
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  PMID: 18308531;Abstract: Understanding how fungal endophyte communities differ in abundance, diversity, taxonomic composition, and host affinity over the geographic ranges of their hosts is key to understanding the ecology and evolutionary context of endophyte-plant associations. We examined endophytes associated with healthy photosynthetic tissues of three closely related tree species in the Cupressaceae (Coniferales): two native species within their natural ranges [Juniperus virginiana in a mesic semideciduous forest, North Carolina (NC); Cupressus arizonica, under xeric conditions, Arizona (AZ)], and a non-native species planted in each site (Platycladus orientalis). Endophytes were recovered from 229 of 960 tissue segments and represented at least 35 species of Ascomycota. Isolation frequency was more than threefold greater for plants in NC than in AZ, and was 2.5 (AZ) to four (NC) times greater for non-native Platycladus than for Cupressus or Juniperus. Analyses of ITS rDNA for 109 representative isolates showed that endophyte diversity was more than twofold greater in NC than in AZ, and that endophytes recovered in AZ were more likely to be host-generalists relative to those in NC. Different endophyte genera dominated the assemblages of each host species/locality combination, but in both localities, Platycladus harboured less diverse and more cosmopolitan endophytes than did either native host. Parsimony and Bayesian analyses for four classes of Ascomycota (Dothideomycetes, Sordariomycetes, Pezizomycetes, Eurotiomycetes) based on LSU rDNA data (ca 1.2 kb) showed that well-supported clades of endophytes frequently contained representatives of a single locality or host species, underscoring the importance of both geography and host identity in shaping a given plant's endophyte community. Together, our data show that not only do the abundance, diversity, and taxonomic composition of endophyte communities differ as a function of host identity and locality, but that host affinities of those communities are variable as well. © 2007 The British Mycological Society.
• Jiménez-Romero, C., Ortega-Barría, E., Arnold, A. E., & Cubilla-Rios, L. (2008). Activity against Plasmodium falciparum of lactones isolated from the endophytic fungus Xylaria sp.. Pharmaceutical Biology, 46(10-11), 700-703.
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  Abstract: Three lactones were isolated from the culture medium of the endophytic fungus Xylaria sp. Grev. (Xylariaceae). The major compound, which showed weak activity (13 μg/mL) against a chloroquine-resistant strain of Plasmodium falciparum, was identified as (+)-phomalactone (1). The others were 6-(1-propenyl)-3,4,5,6-tetrahydro-5-hydroxy-4H-pyran-2-one (2) and 5-hydroxymellein (3). Compounds 1 and 2 are reported for the first time as constituents of Xylaria. Also, this is the first report of the activity of the compounds 1-3 against a chloroquine-resistant Plasmodium falciparum strain. © 2008 Informa UK Ltd.
• Kluger, C. G., Dalling, J. W., Gallery, R. E., Sanchez, E., Weeks-Galindo, C., & Arnold, A. E. (2008). Host generalists dominate fungal communities associated with seeds of four neotropical pioneer species. Journal of Tropical Ecology, 24(3), 351-354.
• Mejía, L. C., Rojas, E. I., Maynard, Z., Bael, S. V., Arnold, A. E., Hebbar, P., Samuels, G. J., Robbins, N., & Herre, E. A. (2008). Endophytic fungi as biocontrol agents of Theobroma cacao pathogens. Biological Control, 46(1), 4-14.
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  Abstract: Fungal endophytes isolated from healthy Theobroma cacao tissues were screened in vitro for antagonism against major pathogens of cacao. Of tested endophytic morphospecies, 40% (21/52), 65% (28/43) and 27% percent (4/15) showed in vitro antagonism against Moniliophthora roreri (frosty pod rot), Phytophthora palmivora (black pod rot) and Moniliophthora perniciosa (witches broom), respectively. The most common antagonistic mechanism was simple competition for substrate. Nonetheless, 13%, 21%, and 0% of tested morphospecies showed clear antibiosis against M. roreri, P. palmivora, and M. perniciosa, respectively. One isolate of Trichoderma was observed to be parasitic on M. roreri. Endophyte species that were common in the host plants under natural conditions often are good colonizers and grow fast in vitro whereas antibiosis producers usually appear to be relatively rare in nature, tend to grow slowly in vitro, and often are not good colonizers. We suggest that there is an inherent general trade-off between fast growth (high colonization) and production of chemicals that produce antibiosis reactions. Finally, field trials assessing the effects of three endophytic fungi (Colletotrichum gloeosporioides, Clonostachys rosea and Botryosphaeria ribis) on pod loss due to M. roreri and Phytophthora spp. were conducted at four farms in Panama. Although the overall incidence of black pod rot was very low during the tests, treatment with C. gloeosporioides significantly decreased pod loss due to that disease. We observed no decrease in pod loss due to frosty pod rot, but treatment with C. rosea reduced the incidence of cacao pods with sporulating lesions of M. roreri by 10%. The observed reduction in pod loss due to Phytophthora spp., and sporulation by M. roreri, supports the potential of fungal endophytes as biological control agents. Further, these studies suggest that combined information from field censuses of endophytic fungi, in vitro studies, and greenhouse experiments can provide useful a priori criteria for identifying desirable attributes for potential biocontrol agents. © 2008 Elsevier Inc. All rights reserved.
• Rojas, E. I., Herre, E. A., Mej́ia, L. C., Arnold, A. E., Chaverri, P., & Samuels, G. J. (2008). Endomelanconiopsis, a new anamorph genus in the Botryosphaeriaceae. Mycologia, 100(5), 760-775.
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  PMID: 18959162;Abstract: A new lineage is discovered within the Botryosphaeriaceae (Ascomycetes, Dothideomycetes, incertae sedis). Consistent with current practice of providing generic names for independent lineages, this lineage is described as Endomelanconiopsis gen. nov., with the anamorphic species E. endophytica sp. nov. and E. microspora comb. nov. (5 Endomelanco-nium microsporum). Endomelanconiopsis is characterized by eustromatic conidiomata and holoblastically produced, brown, nonapiculate, unicellular conidia, each with a longitudinal germ slit. Phylogenetic analysis of partial sequences of LSU, ITS and translation elongation factor 1 alpha (tef1) indicate that E. endophytica is sister of E. microspora and that they are nested within the Botryosphaeriaceae. However because there is no support for the ''backbone'' of the Botryosphaeriacae we are not able to see the interrelationships among the many genera in the family. Neither species is known to have a teleomorph. Endomelanconiopsis differs from En-domelanconium because conidia of the type species of Endomelanconium, E. pini, are papillate at the base, conidiogenous cells proliferate sympodially and the pycnidial wall is thinner; we postulate that the teleomorph of E. pini as yet unknown is an inoperculate discomycete. Endomelanconiopsis endo-phytica was isolated as an endophyte from healthy leaves of Theobroma cacao (cacao, Malvaceae) and Heisteria concinna (Erythroplaceae) in Panama. En-domelanconiopsis microspora was isolated from soil in Europe. © 2008 by The Mycological Society of America.
• Agrawal, A. A., Ackerly, D. D., Adler, F., Arnold, A. E., Cáceres, C., Doak, D. F., Post, E., Hudson, P. J., Maron, J., Mooney, K. A., Power, M., Schemske, D., Stachowicz, J., Strauss, S., Turner, M. G., & Werner, E. (2007). Filling key gaps in population and community ecology. Frontiers in Ecology and the Environment, 5(3), 145-152.
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  Abstract: We propose research to fill key gaps in the areas of population and community ecology, based on a National Science Foundation workshop identifying funding priorities for the next 5-10 years. Our vision for the near future of ecology focuses on three core areas: predicting the strength and context-dependence of species interactions across multiple scales; identifying the importance of feedbacks from individual interactions to ecosystem dynamics; and linking pattern with process to understand species coexistence. We outline a combination of theory development and explicit, realistic tests of hypotheses needed to advance population and community ecology. © The Ecological Society of America.
• Agrawal, A. A., Ackerly, D. D., Adler, F., Arnold, A. E., Cáceres, C., Doak, D. F., Post, E., Hudson, P. J., Maron, J., Mooney, K. A., Power, M., Stachowicz, J., Strauss, S., Turner, M. G., & Werner, E. (2007). The authors reply [2]. Frontiers in Ecology and the Environment, 5(6), 294-295.
• Arnold, A. E. (2007). Understanding the diversity of foliar endophytic fungi: progress, challenges, and frontiers. Fungal Biology Reviews, 21(2-3), 51-66.
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  Abstract: Fungal endophytes, a diverse group of primarily ascomycetous fungi defined functionally by their occurrence within asymptomatic photosynthetic tissues of plants, occur in all major lineages of land plants and in natural and anthropogenic communities ranging from the arctic to the tropics. Because of the tremendous diversity they encompass, ecological questions regarding the interactions of endophytes with the plants in which they live - and with other organisms that in turn interact with endophyte-plants symbiota - are difficalt to address. The goals of this review are to highlight progress, challenges, and frontiers in the study of foliar endophyte diversity, with the ultimate goal of encouraging research that both bridges the gap between, and advances, research in alpha taxonomy and ecology. I focus on four themes that are reflected in the recent and rapidly expanding literature regarding endophyte biology: (1) the taxonomic and ecological distinctiveness of endophytes relative to other nonpathogenic plant-associated fungi; (2) the insights that can be gained from studies that consider genotypes as the relevant unit of biological organization, especially in the context of traditional species-level taxonomy and robust phylogenetic methods that tie these genotypes and species together in an explicit evolutionary context; (3) the context-dependency of endophyte communities, highlighting the importance of both the identify of host plants and the geographic location in which plants occur; and (4) the complexity of the endophyte-pathogen-saprotroph continuum, and the challeges and exciting frontiers that lie in understanding the evolutionary relationships and ecological lability of fungi that exhibit these ecological modes. I argue that never before has the study of endophytic fungi been more exciting or more tractable, and that the potential for endophyte researchers to inform diverse areas of biology has never been greater. © 2007.
• Arnold, A. E., & Lutzoni, F. (2007). Diversity and host range of foliar fungal endophytes: Are tropical leaves biodiversity hotspots?. Ecology, 88(3), 541-549.
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  PMID: 17503580;Abstract: Fungal endophytes are found in asymptomatic photosynthetic tissues of all major lineages of land plants. The ubiquity of these cryptic symbionts is clear, but the scale of their diversity, host range, and geographic distributions are unknown. To explore the putative hyperdiversity of tropical leaf endophytes, we compared endophyte communities along a broad latitudinal gradient from the Canadian arctic to the lowland tropical forest of central Panama. Here, we use molecular sequence data from 1403 endophyte strains to show that endophytes increase in incidence, diversity, and host breadth from arctic to tropical sites. Endophyte communities from higher latitudes are characterized by relatively few species from many different classes of Ascomycota, whereas tropical endophyte assemblages are dominated by a small number of classes with a very large number of endophytic species. The most easily cultivated endophytes from tropical plants have wide host ranges, but communities are dominated by a large number of rare species whose host range is unclear. Even when only the most easily cultured species are considered, leaves of tropical trees represent hotspots of fungal species diversity, containing numerous species not yet recovered from other biomes. The challenge remains to recover and identify those elusive and rarely cultured taxa with narrower host ranges, and to elucidate the ecological roles of these little-known symbionts in tropical forests. © 2007 by the Ecological Society of America.
• Arnold, A. E., & M., B. (2007). Fungal endophytes nearly double minimum leaf conductance in seedlings of a neotropical tree species. Journal of Tropical Ecology, 23(3), 369-372.
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  Abstract: Drought strongly influences plant phenology, growth and mortality in tropical forests, thereby shaping plant performance, population dynamics and community structure (Bunker & Carson 2005, Condit et al. 1995). Microbial symbionts of plants profoundly influence host water relations (Lösch & Gansert 2002), but are rarely considered in studies of tropical plant physiology. In particular, plant-fungus associations, which are ubiquitous in plant communities and especially common in tropical forests, play important and varied roles in plant water status. Fungal pathogens associated with roots, vascular tissue and foliage may interfere with water uptake and transport, increase rates of foliar transpiration, and induce xylem embolism and tissue death (Agrios 1997). In contrast, rhizosphere mutualists such as ecto- and arbuscular mycorrhizal fungi may benefit hosts by increasing surface area for water uptake, enhancing stomatal regulation of water loss, and increasing root hydraulic conductivity (Auge 2001, Lösch & Gansert 2002). Copyright © 2007 Cambridge University Press.
• Arnold, A. E., Henk, D. A., Eells, R. L., Lutzoni, F., & Vilgalys, R. (2007). Diversity and phylogenetic affinities of foliar fungal endophytes in loblolly pine inferred by culturing and environmental PCR. Mycologia, 99(2), 185-206.
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  PMID: 17682771;Abstract: We examined endophytic fungi in asymptomatic foliage of loblolly pine (Pinus taeda) in North Carolina, USA, with four goals: (i) to evaluate morphotaxa, BLAST matches and groups based on sequence similarity as functional taxonomic units; (ii) to explore methods to maximize phylogenetic signal for environmental datasets, which typically contain many taxa but few characters; (iii) to compare culturing vs. culture-free methods (environmental PCR of surface sterilized foliage) for estimating endophyte diversity and species composition; and (iv) to investigate the relationships between traditional ecological indices (e.g. Shannon index) and phylogenetic diversity (PD) in estimating endophyte diversity and spatial heterogeneity. Endophytes were recovered in culture from 87 of 90 P. taeda leaves sampled, yielding 439 isolates that represented 24 morphotaxa. Sequence data from the nuclear ribosomal internal transcribed spacer (ITS) for 150 isolates revealed 59 distinct ITS genotypes that represented 24 and 37 unique groups based on 90% and 95% sequence similarity, respectively. By recoding ambiguously aligned regions to extract phylogenetic signal and implementing a conservative phylogenetic backbone constraint, we recovered well supported phylogenies based on ca. 600 bp of the nuclear ribosomal large subunit (LSUrDNA) for 72 Ascomycota and Basidiomycota, 145 cultured endophytes and 33 environmental PCR samples. Comparisons with LSUrDNA-delimited species showed that morphotaxa adequately estimated total species richness but rarely corresponded to biologically meaningful groups. ITS BLAST results were variable in their utility, but ITS genotype groups based on 90% sequence similarity were concordant with LSUrDNA-delimited species. Environmental PCR yielded more genotypes per sampling effort and recovered several distinct clades relative to culturing, but some commonly cultured clades were never found (Sordariomycetes) or were rare relative to their high frequency among cultures (Leotiomycetes). In contrast to traditional indices, PD demonstrated spatial heterogeneity in endophyte assemblages among P. taeda trees and study plots. Our results highlight the need for caution in designating taxonomic units based on gross cultural morphology or ITS BLAST matches, the utility of phylogenetic tools for extracting robust phylogenies from environmental samples, the complementarity of culturing and environmental PCR, the utility of PD relative to traditional ecological indices, and the remarkably high diversity of foliar fungal endophytes in this simplified temperate ecosystem. © 2007 by The Mycological Society of America.
• Gallery, R. E., Dalling, J. W., & Arnold, A. E. (2007). Diversity, host affinity, and distribution of seed-infecting fungi: A case study with Cecropia. Ecology, 88(3), 582-588.
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  PMID: 17503585;Abstract: Recruitment limitation has been proposed as an important mechanism contributing to the maintenance of tropical tree diversity. For pioneer species, infection by fungi significantly reduces seed survival in soil, potentially influencing both recruitment success and adult distributions. We examined fresh seeds of four sympatric Cecropia species for evidence of fungal infection, buried seeds for five months in common gardens below four C. insignis crowns in central Panama, and measured seed survival and fungal infection of inviable seeds. Seed survival varied significantly among species and burial sites, and with regard to local (Panama) vs. foreign (Costa Rica) maternal seed sources. Fresh seeds contained few cultivable fungi, but >80% of soil-incubated seeds were infected by diverse Ascomycota, including putative pathogens, saprophytes, and endophytes. From 220 isolates sequenced for the nuclear internal transcribed spacer region (ITS), 26 of 73 unique genotypes were encountered more than once. Based on the most common genotypes, fungal communities demonstrate host affinity and are structured at the scale of individual crowns. Similarity among fungal communities beneath a given crown was significantly greater than similarity among isolates found under different crowns. However, the frequency of rare species suggests high fungal diversity and fine-scale spatial heterogeneity. These results reveal complex plant-fungal interactions in soil and provide a first indication of how seed survival in tropical forests may be affected by fungal community composition. © 2007 by the Ecological Society of America.
• Gallery, R. E., Dalling, J. W., Wolfe, B. T., & Arnold, A. E. (2007). The influence of seed source, Habitat and fungi on cecropia seed survival in two neotropical forests. Seed Dispersal: Theory and its Application in a Changing World, 479-498.
• Higgins, K. L., Arnold, A. E., Miadlikowska, J., Sarvate, S. D., & Lutzoni, F. (2007). Phylogenetic relationships, host affinity, and geographic structure of boreal and arctic endophytes from three major plant lineages. Molecular Phylogenetics and Evolution, 42(2), 543-555.
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  PMID: 17005421;Abstract: Although associated with all plants, fungal endophytes (microfungi that live within healthy plant tissues) represent an unknown proportion of fungal diversity. While there is a growing appreciation of their ecological importance and human uses, little is known about their host specificity, geographic structure, or phylogenetic relationships. We surveyed endophytic Ascomycota from healthy photosynthetic tissues of three plant species (Huperzia selago, Picea mariana, and Dryas integrifolia, representing lycophytes, conifers, and angiosperms, respectively) in northern and southern boreal forest (Québec, Canada) and arctic tundra (Nunavut, Canada). Endophytes were recovered from all plant species surveyed, and were present in
• Tsai, Y. J., Maloney, K., & Arnold, A. E. (2007). Biotic and abiotic factors influencing the distribution of the Huachuca springsnail (Pyrgulopsis thompsoni). Journal of Freshwater Ecology, 22(2), 213-218.
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  Abstract: The Huachuca springsnail (Pyrgulopsis thompsoni) is a species of concern inhabiting springs of the Huachuca Mountains in southeastern Arizona, USA. We documented springsnail distribution and examined the major abiotic and biotic factors influencing springsnail abundance at eight spring channels before and after the onset of seasonal summer rains in 2003. Of the abiotic factors examined (total dissolved solids, pH, distance from spring source, spring channel, water temperature, and dissolved oxygen), only spring channel and total dissolved solids were strongly associated with springsnail abundance. However, correlation of total dissolved solids with pH, temperature, dissolved O 2, and distance downstream necessitated further exploration of these explanatory variables. We found that sampling locations with P. thompsoni were characterized by cooler, more oxygenated, and less-occluded water conditions relative to sampling locations without springsnails. Springsnail abundance was positively associated with abundance of caddisfly larvae (Helicopsyche sp.) but not significantly associated with abundance of a co-occurring snail (Physa sp.). The most important outcomes of this study were the documentation of major factors associated with springsnail abundance, and recovery of springsnails at greater distances downstream than previously documented.
• James, T. Y., Kauff, F., Schoch, C. L., Matheny, P. B., Hofstetter, V., Cox, C. J., Celio, G., Gueidan, C., Fraker, E., Miadlikowska, J., Lumbsch, H. T., Rauhut, A., Reeb, V., Arnold, A. E., Amtoft, A., Stajich, J. E., Hosaka, K., Sung, G., Johnson, D., , O'Rourke, B., et al. (2006). Reconstructing the early evolution of Fungi using a six-gene phylogeny. Nature, 443(7113), 818-822.
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  PMID: 17051209;Abstract: The ancestors of fungi are believed to be simple aquatic forms with flagellated spores, similar to members of the extant phylum Chytridiomycota (chytrids). Current classifications assume that chytrids form an early-diverging clade within the kingdom Fungi and imply a single loss of the spore flagellum, leading to the diversification of terrestrial fungi. Here we develop phylogenetic hypotheses for Fungi using data from six gene regions and nearly 200 species. Our results indicate that there may have been at least four independent losses of the flagellum in the kingdom Fungi. These losses of swimming spores coincided with the evolution of new mechanisms of spore dispersal, such as aerial dispersal in mycelial groups and polar tube eversion in the microsporidia (unicellular forms that lack mitochondria). The enigmatic microsporidia seem to be derived from an endoparasitic chytrid ancestor similar to Rozella allomycis, on the earliest diverging branch of the fungal phylogenetic tree. ©2006 Nature Publishing Group.
• Miadlikowska, J., Kauff, F., Hofstetter, V., Fraker, E., Grube, M., Hafellner, J., Reeb, V., Hodkinson, B. P., Kukwa, M., Lücking, R., Hestmark, G., Otalora, M. G., Rauhut, A., Büdel, B., Scheidegger, C., Timdal, E., Stenroos, S., Brodo, I., Perlmutter, G. B., , Ertz, D., et al. (2006). New insights into classification and evolution of the Lecanoromycetes (Pezizomycotina, Ascomycota) from phylogenetic analyses of three ribosomal RNA- and two protein-coding genes. Mycologia, 98(6), 1088-1103.
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  PMID: 17486983;Abstract: The Lecanoromycetes includes most of the lichen-forming fungal species (>13 500) and is therefore one of the most diverse class of all Fungi in terms of phenotypic complexity. We report phylogenetic relationships within the Lecanoromycetes resulting from Bayesian and maximum likelihood analyses with complementary posterior probabilities and bootstrap support values based on three combined multilocus datasets using a supermatrix approach. Nine of 10 orders and 43 of 64 families currently recognized in Eriksson's classification of the Lecanoromycetes (Outline of Ascomycota - 2006 Myconet 12:1-82) were represented in this sampling. Our analyses strongly support the Acarosporomycetidae and Ostropomycetidae as monophyletic, whereas the delimitation of the largest subclass, the Lecanoromycetidae, remains uncertain. Independent of future delimitation of the Lecanoromycetidae, the Rhizocarpaceae and Umbilicariaceae should be elevated to the ordinal level. This study shows that recent classifications include several nonmonophyletic taxa at different ranks that need to be recircumscribed. Our phylogenies confirm that ascus morphology cannot be applied consistently to shape the classification of lichen-forming fungi. The increasing amount of missing data associated with the progressive addition of taxa resulted in some cases in the expected loss of support, but we also observed an improvement in statistical support for many internodes. We conclude that a phylogenetic synthesis for a chosen taxonomic group should include a comprehensive assessment of phylogenetic confidence based on multiple estimates using different methods and on a progressive taxon sampling with an increasing number of taxa, even if it involves an increasing amount of missing data. © 2006 by The Mycological Society of America.
• Spatafora, J. W., Sung, G., Johnson, D., Hesse, C., O'Rourke, B., Serdani, M., Spotts, R., Lutzoni, F., Hofstetter, V., Miadlikowska, J., Reeb, V., Gueidan, C., Fraker, E., Lumbsch, T., Lücking, R., Schmitt, I., Hosaka, K., Aptroot, A., Roux, C., , Miller, A. N., et al. (2006). A five-gene phylogeny of Pezizomycotina. Mycologia, 98(6), 1018-1028.
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  PMID: 17486977;Abstract: Pezizomycotina is the largest subphylum of Ascomycota and includes the vast majority of filamentous, ascoma-producing species. Here we report the results fromweighted parsimony, maximum likelihood and Bayesian phylogenetic analyses of five nuclear loci (SSU rDNA, LSU rDNA, RPB1, RPB2 and EF-1α) from 191 taxa. Nine of the 10 Pezizomycotina classes currently recognized were represented in the sampling. These data strongly supported the monophyly of Pezizomycotina, Arthoniomycetes, Eurotiomycetes, Orbiliomycetes and Sordariomycetes. Pezizomycetes and Dothideomycetes also were resolved as monophyletic but not strongly supported by the data. Lecanoromycetes was resolved as paraphyletic in parsimony analyses but monophyletic in maximum likelihood and Bayesian analyses. Leotiomycetes was polyphyletic due to exclusion of Geoglossaceae. The two most basal classes of Pezizomycotina were Orbiliomycetes and Pezizomycetes, both of which comprise species that produce apothecial ascomata. The seven remaining classes formed a monophyletic group that corresponds to Leotiomyceta. Within Leotiomyceta, the supraclass clades of Leotiomycetes s.s. plus Sordariomycetes and Arthoniomycetes plus Dothideomycetes were resolved with moderate support. © 2006 by The Mycological Society of America.
• Lutzoni, F., Kauff, F., Cox, C. J., McLaughlin, D., Celio, G., Dentinger, C., Padamsee, M., Hibbett, D., James, T. Y., Baloch, E., Grube, M., Reeb, V., Hofstetter, V., Schoch, C., Arnold, A. E., Miadlikowska, J., Spatafora, J., Johnson, D., Hambleton, S., , Crockett, M., et al. (2004). Assembling the fungal tree of life: Progress, classification, and evolution of subcellular traits. American Journal of Botany, 91(10), 1446-1480.
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  PMID: 21652303;Abstract: Based on an overview of progress in molecular systematics of the true fungi (Fungi/Eumycota) since 1990, little overlap was found among single-locus data matrices, which explains why no large-scale multilocus phylogenetic analysis had been undertaken to reveal deep relationships among fungi. As part of the project "Assembling the Fungal Tree of Life" (AFTOL), results of four Bayesian analyses are reported with complementary bootstrap assessment of phylogenetic confidence based on (1) a combined two-locus data set (nucSSU and nucLSU rDNA) with 558 species representing all traditionally recognized fungal phyla (Ascomycota, Basidiomycota, Chytridiomycota, Zygomycota) and the Glomeromycota, (2) a combined three-locus data set (nucSSU, nucLSU, and mitSSU rDNA) with 236 species, (3) a combined three-locus data set (nucSSU, nucLSU rDNA, and RPB2) with 157 species, and (4) a combined four-locus data set (nucSSU, nucLSU, mitSSU rDNA, and RPB2) with 103 species. Because of the lack of complementarity among single-locus data sets, the last three analyses included only members of the Ascomycota and Basidiomycota. The four-locus analysis resolved multiple deep relationships within the Ascomycota and Basidiomycota that were not revealed previously or that received only weak support in previous studies. The impact of this newly discovered phylogenetic structure on supraordinal classifications is discussed. Based on these results and reanalysis of subcellular data, current knowledge of the evolution of septal features of fungal hyphae is synthesized, and a preliminary reassessment of ascomal evolution is presented. Based on previously unpublished data and sequences from GenBank, this study provides a phylogenetic synthesis for the Fungi and a framework for future phylogenetic studies on fungi.
• Arnold, A. E., & Herre, E. A. (2003). Canopy cover and leaf age affect colonization by tropical fungal endophytes: Ecological pattern and process in Theobroma cacao (Malvaceae). Mycologia, 95(3), 388-398.
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  PMID: 21156627;Abstract: Fungal endophytes inhabit healthy tissues of all terrestrial plant taxa studied to date and are diverse and abundant in leaves of tropical woody angiosperms. Studies have demonstrated that plant location and leaf age influence density of endophyte infection in leaves of tropical forest trees. However, ecological factors underlying these observations have not been explored in detail. Here, we establish that foliar endophytes of a tropical tree (Theobroma cacao, Malvaceae) are transmitted horizontally and that endophyte-free seedlings can be produced for experimental manipulation by protecting aerial tissues from surface wetting. At Barro Colorado Island, Panama, we used transects of endophyte-free seedlings to determine the importance of several factors (canopy cover, abundance of aerial and epiphytic propagules, leaf age, leaf chemistry, leaf toughness and duration of exposure to viable air spora) in shaping colonization by endophytic fungi. Endophytes colonized leaves of T. cacao more rapidly beneath the forest canopy than in cleared sites, reflecting local abundance of aerial and epiphytic propagules. The duration of exposure, rather than absolute leaf age, influenced endophyte infection, whereas leaf toughness and chemistry had no observed effect. Endophytes isolated from mature T. cacao grew more rapidly on media containing leaf extracts of T. cacao than on media containing extracts from other co-occurring tree species, suggesting that interspecific differences in leaf chemistry influence endophyte assemblages. Together, these data allow us to identify factors underlying patterns of endophyte colonization within healthy leaves of this tropical tree.
• Arnold, A. E., Mejía, L. C., Kyllo, D., Rojas, E. I., Maynard, Z., Robbins, N., & Herre, E. A. (2003). Fungal endophytes limit pathogen damage in a tropical tree. Proceedings of the National Academy of Sciences of the United States of America, 100(26), 15649-15654.
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  PMID: 14671327;PMCID: PMC307622;Abstract: Every plant species examined to date harbors endophytic fungi within its asymptomatic aerial tissues, such that endophytes represent a ubiquitous, yet cryptic, component of terrestrial plant communities. Fungal endophytes associated with leaves of woody angiosperms are especially diverse; yet, fundamental aspects of their interactions with hosts are unknown. In contrast to the relatively species-poor endophytes that are vertically transmitted and act as defensive mutualists of some temperate grasses, the diverse, horizontally transmitted endophytes of woody angiosperms are thought to contribute little to host defense. Here, we document high diversity, spatial structure, and host affinity among foliar endophytes associated with a tropical tree (Theobroma cacao, Malvaceae) across lowland Panama. We then show that inoculation of endophyte-free leaves with endophytes isolated frequently from naturally infected, asymptomatic hosts significantly decreases both leaf necrosis and leaf mortality when T. cacao seedlings are challenged with a major pathogen (Phytophthora sp.). In contrast to reports of fungal inoculation inducing systemic defense, we found that protection was primarily localized to endophyte-infected tissues. Further, endophyte-mediated protection was greater in mature leaves, which bear less intrinsic defense against fungal pathogens than do young leaves. In vitro studies suggest that host affinity is mediated by leaf chemistry, and that protection may be mediated by direct interactions of endophytes with foliar pathogens. Together, these data demonstrate the capacity of diverse, horizontally transmitted endophytes of woody angiosperms to play an important but previously unappreciated role in host defense.
• Arnold, A. E., & Asquith, N. M. (2002). Herbivory in a fragmented tropical forest: Patterns from islands at Lago Gatún, Panama. Biodiversity and Conservation, 11(9), 1663-1680.
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  Abstract: By imposing density-dependent mortality upon their hosts, specialist insect herbivores are thought to contribute to the maintenance of tree diversity in tropical forests. Forest fragmentation may alter patterns of herbivory, however, which may have important implications for tree species diversity in forest remnants. To explore effects of fragmentation on patterns of herbivory, we assessed folivory by Lepidopteran larvae on saplings of four focal tree species on eight artificial, forested islands at Lago Gatún, Panama. We explored the importance of island area, distance to larger land, exposure to dry season winds, tree species, and season in determining proportions of new leaves damaged by caterpillars, and proportions of leaf area lost to caterpillars, during two dry and wet seasons. We found that both measures of herbivory increased markedly with island area, that island isolation had no apparent effect on herbivory, and that interactions between season and exposure, and between tree species and season, were important determinants of herbivory rates. In addition, we observed species-specific differences in herbivory among host plants under various conditions imposed by fragmentation. We conclude that patterns of herbivory by Lepidopteran larvae are sensitive to fragmentation in this tropical forest. Differential herbivory among the four tree species considered here may have important implications for tree species dynamics on the islands of Lago Gatún.
• Arnold, A. E., Maynard, Z., & Gilbert, G. S. (2001). Fungal endophytes in dicotyledonous neotropical trees: Patterns of abundance and diversity. Mycological Research, 105(12), 1502-1507.
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  Abstract: Fungal endophytes have been found in every plant species examined to date and appear to be important, but largely unquantified, components of fungal biodiversity. Endophytes are especially little known in tropical forest trees, where their abundance and diversity are thought to be greatest. Here, we explore the occurrence of endophytes in a broad diversity of woody, angiospermous taxa in a lowland, moist tropical forest in central Panamá. We use similarity indices to assess host preference and spatial heterogeneity of endophytes associated with two co-occurring, but distantly related, understorey tree species in two sites of that forest, and assess the utility of indices based on frequencies of morphospecies occurrence (Morisita-Horn index) and on presence-absence data (Sørensen's index). We suggest that our understanding of fungal diversity will be enhanced by exploring ecological patterns underlying endophyte occurrence in host species, and discuss methods for assessing the proportion of fungal biodiversity represented by tropical endophytes.
• Arnold, A. E., Maynard, Z., Gilbert, G. S., Coley, P. D., & Kursar, T. A. (2000). Are tropical fungal endophytes hyperdiverse?. Ecology Letters, 3(4), 267-274.
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  Abstract: Fungal endophytes are ubiquitous fungi that inhabit healthy plant tissues without causing disease. Endophytes have been found in every plant species examined to date and may be important, but often overlooked, components of fungal biodiversity. In two sites in a lowland, moist tropical forest of central Panama, we quantified endophyte colonization patterns, richness, host preference, and spatial variation in healthy leaves of two co-occurring, understory tree species [Heisteria concinna (Olacaceae) and Ouratea lucens (Ochnaceae)]. From 83 leaves, all of which were colonized by endophytes, we isolated 418 endophyte morphospecies (estimated 347 genetically distinct taxa), most of which were represented by only a single isolate (59%). Among morphospecies encountered in more than one leaf (nonsingletons), we found evidence of host preference and spatial heterogeneity using both morphospecies frequencies and presence/absence records. Based on these data, we postulate that tropical endophytes themselves may be hyperdiverse and suggest that extrapolative estimates that exclude them will markedly underestimate fungal species diversity.
• Asquith, N. M., Terborgh, J., Arnold, A. E., & Riveros, C. M. (1999). The fruits the agouti ate: Hymenaea courbaril seed fate when its disperser is absent. Journal of Tropical Ecology, 15(2), 229-235.

Presentations

• Arnold, A. E. (2021). 4 oral presentations, contributed: senior author or second author on all; all co-authored/led by students. Ecological Society of America (3 talks), American Phytopathological Society, Southern Division (1 talk).
• Arnold, A. E. (2021). 6 invited presentations at national/international venues; sole or first author on 5, senior author on 1. Harvard University (1 talk), National Science Foundation (1 talk, 1 poster), Smithsonian Tropical Research Institute (1 talk, and panelist), University of Wisconsin (1 talk), International Association of Lichenology Symposium (1 talk).
• Arnold, A. E. (2020). 2 oral presentations, contributed: senior author or second author on both, both co-authored/led by students. Ecological Society of America (1 talk), American Association for the Advancement of Science (1 talk).
• Arnold, A. E. (2019, Various). 8 invited scholarly presentations: seminar/keynote/symposium/plenary. Harvard University, UA Ecosystem Genomics, Northern Arizona University, Mycological Society of America, Botanical Society of America (x2), UA UBRP program, RISE symposium. Various: Various.
• Arnold, A. E., & Various, V. (2019, Various). 8 contributed presentations at national and international meetings (* = poster, ** = student award). American Society for Horticultural Science*,**, Instituti Gulbenkian de Ciencia (Portugal)*, Botanical Society of America, Mycological Society of America, Fungal Genetics Conference*, British Ecological Society, West Coast Biological Sciences Undergraduate Meeting (x2)*. Various.
• Arnold, A. E. (2018, Summer/fall). 11 intramural presentations by undergraduates and graduate students for which I was senior author. Summer Research Experiences for Undergraduates, including Native American and international programs (6 presentations), graduate presentations in departmental seminars (4), undergraduate presentation in ASEMS symposium. Tucson: University of Arizona.
• Arnold, A. E. (2018, Summer/fall). 8 invited symposium/seminar presentations for which I was sole author. International Symbiosis Society Congress (Plenary); Gordon Research Conference; Coral Reef Biology Congress; University of Massachusetts, Amherst; University of Minnesota; University of Arizona Tech Launch; University of Arizona SWES. Oregon, New Hampshire, Queensland Australia, Amherst, Minneapolis St. Paul, and Tucson, respectively..
• Spraker, J., Shaffer, J. P., Baltrus, D. A., Traxler, M., & Arnold, A. E. (2018, Summer). 1 invited symposium presentation for which I was senior author. International Mycological Congress. San Juan, Puerto Rico.
• Arnold, A. E. (2017, Fall). Perspectives from leaves and lichens on the scale and distribution of the global endobiome. Invited seminar, University of Michigan (Herb Wagner Memorial Lecture). Ann Arbor, MI.
• Arnold, A. E. (2017, Summer). Interactions across boundaries promote symbiotic modulation of plant phenotypes.. Invited Symposium, Botanical Society of America. Fort Worth, TX.
• Bowman, E. A., & Arnold, A. E. (2017, Summer). Ectomycorrhizal and foliar endophytic fungal communities differ in sensitivity to climate-related factors along a spatially constrained elevation gradient. Yosemite Symbiosis Workshop. Yosemite National Park.
• Tellez, P., Arnold, A. E., & vanBael, S. (2017, Summer 2017). Tropical plants and fungal symbionts: leaf functional traits as drivers of plant-fungal endophyte interactions. Association for Tropical Biology and Conservation. Merida, Mexico.
• Tellez, P., Arnold, A. E., & vanBael, S. (2017, Summer). Tropical plants and fungal symbionts: leaf functional traits as drivers of plant-fungal endophyte interactions. Ecological Society of America. Portland, OR.
• U'Ren, J. M., Lutzoni, F., Miadlikowska, J., Leo, A. B., May, G., Carbone, I., & Arnold, A. E. (2017, Summer). Environmental drivers and spatial structuring of boreal endolichenic fungi and local, regional, and global scales. Future Arctic: Global Initiative on bryophyte and lichen Arctic research from species to ecosystems. Quebec, Canada.
• Zalamea, P., Sarmiento, C., Davis, A. S., Arnold, A. E., & Dalling, J. W. (2017, Summer). Seed-associated fungi in neotropical pioneers influence seed persistence, germination, and survival. Association for Tropical Biology and Conservation. Merida, Mexico.
• Zimmerman, N., U'Ren, J. M., & Arnold, A. E. (2017, Summer). High resolution genotyping reveals extensive diversification of trichome-associated fungi at high elevation sites in Hawai'i. Ecological Society of America. Portland, OR.
• Arnold, A. E. (2016, Fall). Discovering new agricultural resources from the plant microbiome. Invited scholarly presentation, Symbiota LLC.
• Arnold, A. E. (2016, Spring). Fungal endophytes from the tropics to the tundra: clues to the evolution of fungal symbioses.. Invited seminar, Stanford University.
• Arnold, A. E. (2016, Spring). Fungal endophytes from the tropics to the tundra: clues to the evolution of fungal symbioses.. Keynote presentation, Yosemite Symbiosis Conference.
• Arnold, A. E. (2016, Spring). Perspectives from leaves and lichens on the scale and distribution of the global endobiome. Invited seminar, Ecology and Evolutionary Biology, University of Arizona.
• Arnold, A. E., U'Ren, J., Miadlikowska, J., Carbone, I., Bowman, E., Huang, Y., May, G., & Lutzoni, F. (2016, Summer). Perspectives from leaves and lichens on the scale and distribution of the global endobiome. Invited symposium presentation, Mycological Society of America.
• Bowman, E., & Arnold, A. E. (2016, Summer). Fungal symbionts of ponderosa pine across a spatially constrained elevation gradient. Annual meeting, Mycological Society of America.
• Carbone, I., White, J., Miadlikowska, J., Arnold, A. E., Miller, M., Kauff, F., Schoch, C., U'Ren, J., May, G., & Lutzoni, F. (2016, Summer). Enhancing fungal species discovery and description using T-BAS: Tree-based alignment selector toolkit. Invited seminar, National Science Foundation.
• Chen, K., Liao, H., Arnold, A. E., & Lutzoni, F. (2016, Summer). Metatranscriptomic analysis of the moss Dicranum scoparium reveals active fungal communities and functionalities across a senescence gradient. Annual meeting, Mycological Society of America.
• Chen, K., Liao, H., Arnold, A. E., & Lutzoni, F. (2016, Summer). Metatranscriptomic analysis of the moss Dicranum scoparium reveals active fungal communities and functionalities across a senescence gradient. Invited symposium presentation, Botanical Society of America.
• Shaffer, J., & Arnold, A. E. (2016, Summer). Endohyphal bacterium (Chitinophaga sp.) influences broad-spectrum substrate use by its host fungus (Fusarium keratoplasticum). Annual meeting, Mycological Society of America.
• U'Ren, J., Miadlikowska, J., Lutzoni, F., Leo, A., May, G., Carbone, I., & Arnold, A. E. (2016, Summer). Environmental drivers and spatial structuring of boreal endolichenic fungi and local, regional, and global scales. Invited seminar, International Lichenology Symposium.
• U'Ren, J., Miadlikowska, J., Lutzoni, F., Leo, A., May, G., Carbone, I., Oita, S., & Arnold, A. E. (2016, Summer). Geographic and temporal structure of endophytic and endolichenic fungal communities of the boreal biome. Invited seminar, National Science Foundation.
• Zimmerman, N., Huang, Y., Johnson, J., Moore, D., & Arnold, A. E. (2016, Summer). The effects of foliar fungal endophytes on plant physiological performance. Annual meeting, Ecological Society of America.
• Arnold, A. E. (2015, Fall). Discovering new agricultural resources from the plant microbiome. Invited scholarly presentation, Symbiota LLC.
• Arnold, A. E. (2015, Fall). Fungal endophytes from the tropics to the tundra. Invited scholarly presentation, Swedish University of Agricultural Sciences.
• Arnold, A. E. (2015, Fall). Symbiosis, fungal endophytes, and biodiversity. Invited scholarly presentation, Swedish University of Agricultural Sciences, Uppsala, Sweden.
• Arnold, A. E. (2015, Spring). Discovering new resources among the fungi that live within healthy plants. Invited scholarly presentation, University of Arizona School of Natural Resources and the Environment.
• Arnold, A. E. (2015, Spring). Fungal endophytes from the tropics to the tundra: clues to the evolution of fungal symbioses. Invited scholarly presentation, University of Oregon.
• Arnold, A. E. (2015, Spring). Fungal endophytes from the tropics to the tundra: clues to the evolution of fungal symbioses. Invited scholarly presentation, University of Pittsburgh.
• Arnold, A., U'Ren, J., Miadlikowska, J., Carbone, I., & Lutzoni, F. (2015, Spring). Progress toward capturing the biodiversity of fungal endophytes. Invited symposium presentation, International Workshop on Ascomycete Systematics, Amsterdam, The Netherlands.
• Carbone, I., White, J., Miadlikowska, J., U'Ren, J., Arnold, A., & Lutzoni, F. (2015, Spring). New online tools for species delimitation and classification of unknown fungal endophytes. Invited symposium presentation, International Workshop on Ascomycete Systematics, Amsterdam, The Netherlands.
• Lutzoni, F., Carbone, I., White, J., Miadlikowska, J., U'Ren, J., & Arnold, A. (2015, Spring). Challenges to speeding up the name of new fungal species. Invited symposium presentation, International Workshop on Ascomycete Systematics, Amsterdam, The Netherlands.
• Lutzoni, F., U'Ren, J., Miadlikowska, J., Carbone, I., & Arnold, A. (2015, Spring). Filling in the gaps on the fungal tree of life through endophyte discovery. Invited symposium presentation, International Workshop on Ascomycete Systematics, Amsterdam, The Netherlands.
• Sarmiento, C., Zalamea, C., Dalling, J., Davis, A., & Arnold, A. (2015, Summer). Seed-associated fungi: effects on seed survival and germination of tropical pioneer species. Invited symposium presentation, Association for Tropical Biology and Conservation.
• Shaffer, J., Gallery, R., Baltrus, D., & Arnold, A. (2015, Summer). Endohyphal bacteria of tropical Sordariomycetes: community structure and relationships with other functional groups of bacteria in a lowland tropical rainforest. Contributed oral presentation, Mycological Society of America.
• U'Ren, J., Lutzoni, F., Miadlikowska, J., Leo, A., Gleason, T., Chen, K., May, G., Carbone, I., & Arnold, A. (2015, Spring). Geographic and temporal structure of endophytic and endolichenic fungal communities of the boreal biome. Invited symposium presentation, Fungal Genetics Conference, Asilomar, CA.
• Zimmerman, N., Arnold, A., & Vitousek, P. (2015, Spring). A highly diverse clade of melanized fungi associated with leaves and trichomes of the endemic tree Metrosideros polymorpha at high elevation sites in Hawai'i. Invited symposium presentation, Fungal Genetics Conference, Asilomar, CA.
• Zimmerman, N., Arnold, A., & Vitousek, P. (2015, Summer). A highly diverse clade of melanized fungi associated with leaves and trichomes of the endemic tree Metrosideros polymorpha at high elevation sites in Hawai'i. Contributed oral presentation, Ecological Society of America.
• Arnold, A. E. (2014, Fall). Using epigenetic modifiers and new substrates to enhance the pace and impact of ecologically-guided discovery of bioactive metabolites from tropical fungi. Invited symposium presentation, National Institutes of Health - International Cooperative Biodiversity Group, Panama.
• Arnold, A. E. (2014, Fall). What can >50,000 cultures tell us about the ecological specificity of endophytes and related fungi?. Invited symposium presentation, International Mycological Congress.
• Arnold, A. E. (2014, Spring). Fungal endophytes from the tropics to the tundra: clues to the evolution of plant-fungal symbioses. Invited seminar, Tulane University.
• Arnold, A. E. (2014, Spring). Fungal endophytes from the tropics to the tundra: clues to the evolution of plant-fungal symbioses. Invited seminar, University of Virginia.
• Baltrus, D., Arendt, K., & Arnold, A. E. (2014, Summer). Facultative endohyphal bacterial symbionts alter phenotypes of fungal endophyte hosts. Contributed presentation, Society for the Study of Evolution.
• Corrales, A., Arnold, A. E., Ferrer, A., & Dalling, J. (2014, Summer). Variation in ectomycorrhizal communities associated with stands of Oreomunnea mexicana (Juglandaceae) in montane tropical forests of western Panama. Contributed presentation, Latin American Mycological Congress.
• Arendt, K., Baltrus, D., & Arnold, A. E. (2013, Spring). Diversity and specificity of phenotypic effects of endohyphal bacteria on foliar fungal endophytes. Contributed presentation, NSF Dimensions of Biodiversity Boreal Endophyte Meeting.
• Arnold, A. E. (2013, Fall). Fungal endophytes from the tropics to the tundra: clues to the evolution of plant-fungal symbioses. Invited seminar, Northern Arizona University.
• Arnold, A. E. (2013, Fall). Fungal endophytes from the tropics to the tundra: clues to the evolution of plant-fungal symbioses. Invited seminar, University of Georgia.
• Arnold, A. E. (2013, Fall). Fungal endophytes from the tropics to the tundra: clues to the evolution of plant-fungal symbioses. Invited speaker, British Mycological Society.
• Arnold, A. E. (2013, Spring). Fungal endophytes from the tropics to the tundra: clues to the evolution of plant-fungal symbioses. Invited seminar, Michigan State University.
• Arnold, A. E. (2013, Winter). Fungal endophytes from the tropics to the tundra: clues to the evolution of plant-fungal symbioses. Invited seminar, Cornell University.
• Arnold, A. E., Dalling, J., Corrales Osorio, A., & McGuire, K. (2013, Summer). Variation in ectomycorrhizal community composition in montane forest along a soil nutrient gradient in a montane forest of western Panama. Invited symposium presentation, co-author, Association for Tropical Biology and Conservation.
• Chen, K., Miadlikowska, J., Molnar, K., Arnold, A. E., U'Ren, J., Gaya, E., & Lutzoni, F. (2013, Summer). Phylogenetic relationships of endophytic and endolichenic fungi reveal a new order within the class Eurotiomycetes. Contributed presentation, Mycological Society of America.
• Corrales Osorio, A., Dalling, J., Arnold, A. E., & McGuire, K. (2013, Summer). Variation in ectomycorrhizal community composition along a soil nutrient gradient in montane forest in western Panama. Contributed presentation, Mycological Society of America.
• Del Olmo Ruiz, M., Arnold, A. E., Del Olmo Ruiz, M., & Arnold, A. E. (2013, Summer). Diversity, distributions, and host affiliations of endophytes associated with tropical ferns. Invited symposium presentation, co-author, Association for Tropical Biology and Conservation.
• Sarmiento, C., Dalling, J., Zalamea, P. C., Davis, A., & Arnold, A. E. (2013, Summer). Seed - fungal interactions in tropical trees: Exploring fungal diversity in pioneer seeds. Contributed presentation, Association for Tropical Biology and Conservation.
• Sarmiento, C., Roche, D., Zalamea, P. C., Arnold, A. E., Davis, A., & Dalling, J. (2013, Summer). Physical defenses, persistence in the soil, and fungal associations of tropical pioneer tree seeds. Invited symposium presentation, co-author, Association for Tropical Biology and Conservation.
• Shaffer, J., Baltrus, D., & Arnold, A. E. (2013, Spring). Phylogenetic relationships and diversity of endohyphal bacteria of plant-associated Pezizomycotina. Contributed presentation, NSF Dimensions of Biodiversity Boreal Endophyte Meeting.
• U'Ren, J., & Arnold, A. E. (2013, Spring). A culture-based and culture-free assessment of the geographic and temporal variation of boreal endophytic and endolichenic fungal communities. Contributed presentation, NSF Dimensions of Biodiversity Boreal Endophyte Meeting.
• U'Ren, J., Massimo, N., Riddle, J., Steen, C., Arendt, K., Huang, Y., Miadlikowska, J., LeFevre, E., Ball, B., Wong, V., Monacell, J. T., Carbone, I., Lutzoni, F., May, G., & Arnold, A. E. (2013, Summer). A culture-based and culture-free assessment of the geographic and temporal variation of boreal endophytic and endolichenic fungal communities. Contributed presentation, Mycological Society of America.
• Wong, V., U'Ren, J., Miadlikowska, J., Monacell, J. T., Arendt, K., Shaffer, J., Arnold, A. E., Carbone, I., & May, G. (2013, Summer). Genomic comparison of closely related boreal endophytes. Contributed presentation, Mycological Society of America.

Poster Presentations

• Arnold, A. E. (2021). 6 poster presentations, contributed: senior author on all; all co-authored/led by students. Mycological Society of America (2), Research Insights in Semi-arid Ecosystems (2), Annual Biomedical Conference for Minority Students.
• Arnold, A. E. (2020). 6 poster presentations, contributed: senior author on all; all co-authored/led by students. Mycological Society of America (6 posters).
• Arnold, A. E., Wisecaver, J., & Uren, J. M. (2019, March). Comparative and population genomics of endophytic Xylariaceae. Department of Energy Joint Genome Institute Genomics of Energy and the Environment. San Francisco, CA: Department of Energy Joint Genome Institute.
• Arnold, A. E. (2018, Summer/fall). 5 senior-authored posters at regional, national and international conferences, led by students in my group. International Symbiosis Society Congress (2), American Indian Science and Engineering Society (2), Research in Semi-Arid Ecosystems Symposium (2). Oregon, Oklahoma, and Tucson, respectively.
• Arnold, A. E., Uren, J. M., Trouet, V. M., & Oita, S. (2018, August). Relationships of foliar endophyte communities in Picea mariana to tree age, biomass, and latitude. International Symbiosis Society.
• Sarmiento, C., Zalamea, P., Kuo, V., Delevich, C., Davis, A. S., Brown, T. A., Arnold, A. E., & Dalling, J. W. (2017, Summer). Impact of decadal persistence of tropical pioneer seeds on seed-associated fungal communities. Association for Tropical Biology and Conservation. Merida, Mexico.
• Chen, K., Liao, H., Arnold, A., & Lutzoni, F. (2015, Spring). Using metatranscriptomics to characterize functional shifts in endophytic fungi at plant senescence: Are endophytic fungi latent saprotrophs?. Contributed presentation, Fungal Genetics Conference, Asilomar, CA.
• Chen, K., Liao, H., Arnold, A., & Lutzoni, F. (2015, Summer). Using metatranscriptomics to characterize functional shifts in endophytic fungi at plant senescence: Are endophytic fungi latent saprotrophs?. Contributed presentation, Mycological Society of America.
• Huang, Y., Devan, M., Furr, S., U'Ren, J., & Arnold, A. (2015, Summer). Pervasive effects of wildfire on foliar endophytes in montane forest trees. Contributed presentation, Mycological Society of America.
• Shaffer, J., Gallery, R., Baltrus, D., & Arnold, A. (2015, Spring). Endohyphal bacteria of tropical Sordariomycetes: community structure and relationships with other functional groups of bacteria in a lowland tropical rainforest. Contributed presentation, Fungal Genetics Conference, Asilomar, CA.
• Arendt, K., Baltrus, D., & Arnold, A. E. (2014, Summer). Bacterial symbionts of endophytic fungi mediate functional shifts in plant-fungal interactions. Contributed presentation, Ecological Society of America.
• Chen, K., & Arnold, A. E. (2014, Summer). Phylogenetic relationships of endophytic and endolichenic fungi reveal a new order within the class Eurotiomycetes. Contributed presentation, International Mycological Congress.
• Nilsson, H., & Arnold, A. E. (2014, Summer). Improving ITS sequence data for identification of plant-pathogenic fungi. Contributed presentation, International Mycological Congress.
• von Arx, M., Moore, A. F., Davidowitz, G., & Arnold, A. E. (2014, Aug). Diversity and distribution of microbial communities in floral nectar in two typical plants of the Sonoran Desert. International Symposium on Insect-Plant Relationships, Neuchatel, Switzerland.
• Arendt, K., Baltrus, D., & Arnold, A. E. (2013, Summer). Diversity and specificity of phenotypic effects of endohyphal bacteria on foliar fungal endophytes. Contributed presentation, Mycological Society of America.
• Chen, K., Miadlikowska, J., Molnar, K., Arnold, A. E., Gaya, E., & Lutzoni, F. (2013, Summer). Phylogenetic relationships of endophytic and endolichenic fungi reveal a new order in Eurotiomycetes. Contributed presentation, Society for the Study of Evolution.
• Garcia, K., Schaffer, J., Sarmiento, C., Zalamea, C., Dalling, J., Davis, A., Baltrus, D. A., Gallery, R. E., & Arnold, A. E. (2013, August). Diversity and evolutionary relationships of bacteria affiliated with tropical seeds and seed-associated fungi. Mycological Society of America (MSA). Austin, TX: Mycological Society of America (MSA).
• Huang, Y., Devan, M. N., Furr, S., & Arnold, A. E. (2013, Summer). Persistent effects of wildfire on foliar endophytes of Quercus hypoleucoides and Juniperus deppeana in southeastern Arizona. Contributed presentation, Mycological Society of America.
• LeFevre, E., Arendt, K., Ball, B., Miadlikowska, J., Picard, K., U'Ren, J., Arnold, A. E., & Lutzoni, F. (2013, Summer). Understanding the spatial scaling of boreal endophytic fungal communities using environmental cloning and ion semiconductor amplicon sequencing. Contributed presentation, Mycological Society of America.
• Oono, R., Kaye, L., Arnold, A. E., May, G., Lutzoni, F., & Carbone, I. (2013, Spring). Population structures of horizontally-transmitted fungal endophytes associated with southeastern US pine hosts. Contributed presentation, Fungal Genetics Conference.
• Oono, R., Kaye, L., Arnold, A. E., May, G., Lutzoni, F., & Carbone, I. (2013, Summer). Population structures of horizontally-transmitted fungal endophytes associated with southeastern US pine hosts. Contributed presentation, Society for the Study of Evolution.
• Sandberg, D., Battista, L., & Arnold, A. E. (2013, Summer). Host affiliations and geographic distributions of fungal symbionts of aquatic plants. Contributed presentation, Mycological Society of America.
• Shaffer, J., Gallery, R. E., Baltrus, D. A., & Arnold, A. E. (2013, August). Phylogenetic relationships and diversity of endohyphal bacteria of plant-associated Pezizomycotina. Mycological Society of America (MSA). Austin, TX: Mycological Society of America (MSA).
• Shaffer, J., Gallery, R., Baltrus, D., & Arnold, A. E. (2013, Summer). Phylogenetic relationships and diversity of endohyphal bacteria of plant- associated Pezizomycotina. Contributed presentation, American Society for Microbiology.
• Shaffer, J., Gallery, R., Baltrus, D., & Arnold, A. E. (2013, Summer). Phylogenetic relationships and diversity of endohyphal bacteria of plant-associated Pezizomycotina. Contributed presentation, Mycological Society of America.
• Oono, R., Arnold, A., May, G., Lutzoni, F., & Carbone, I. (2012, July). Population structure in Lophodermium sp., a dominant fungal endophyte of loblolly pine. International Symbiosis Society. Krakow, Poland.
• Wong, V., Schwebach, C., Arnold, A., & et, a. l. (2012, June). Effects of climate warming on the occurrence of endophytic fungi in boreal Picea and Populus. Not Provided in APROL. Portland, OR: Ecological Society of America.
• Moy, J., Arnold, A., Predick, K., Levi, E., & Archer, S. (2011, November). Effects of microbial communities on in vitro and in situ degradation of plant material in an arid ecosystem. Research in Semi-Arid Ecosystems Symposium. Tucson, AZ.
• Oono, R., Arnold, A., May, G., Lutzoni, F., & Carbone, I. (2011, August). Population structure in Lophodermium sp., a dominant fungal endophyte of loblolly pine. Mycological Society of America annual meeting. Fairbanks, AK: Mycological Society of America.

Creative Productions

• Arnold, A. E. (2015. Website for lab, for outreach/recruitment/research. Internet: www.arnoldlab.net.
• Arnold, A., & Baltrus, D. (2015. Website for NSF-funded project on endohyphal bacteria. Internet: http://www.endohyphalbacteria.com.

Others

• Arnold, A. E. (2013, Winter). Updates to research websites: www.arnoldlab.net, www.endobiodiversity.org, http://publish.illinois.edu/tropicalseeds/.
• Arnold, A. E. (2010, Fall). NSF project websites. www.endobiodiversity.org
• Arnold, A. E. (2010, Fall). Provided all core content and most photographs for NSF project websites (www.endobiodiversity.org; www.tropicalseeds.org). www.endobiodiversity.org; www.tropicalseeds.org
• Arnold, A. E. (2010, Fall). Provided all core content and most photographs for lab website (www.arnoldlab.net). www.arnoldlab.net