Journal publications and other peer-reviewed works


Published or in press as of September 2013 • Asterisk indicates that lead author was a student


Hoffman*, M.T., M. Gunatilaka, E.M.K. Wijeratne, A.A.L. Gunatilaka, and A.E. Arnold. 2013. IAA

production by endophytic Pestalotiopsis is enhanced by an endohyphal bacterium. PLoS One, in press.


Higgins*, K.L., A.E. Arnold, P. Coley, T. Kursar. 2013. Communities of fungal endophytes in tropical forest grasses: highly diverse host- and habitat generalists characterized by strong spatial structure. Fungal Ecology, in press.


Sandberg*, D.C., L.J. Battista, A.E. Arnold. Host affiliations and geographic distributions of fungal endophytes inhabiting aquatic plants in northern Arizona. Microbial Ecology, in press.


Del Olmo Ruiz*, M. A.E. Arnold. 2013. Interannual variation and host affiliations of endophytic fungi

associated with ferns at La Selva, Costa Rica. Mycologia, in press.


Higginbotham, S.J., A.E. Arnold, A. Ibañez, C. Spadafora, P.D. Coley, T.A. Kursar. 2013. Bioactivity of fungal endophytes as a function of their taxonomy and the taxonomy and distribution of host plants. PLoS One, 8: e73192.


Lau*, M., N. Johnson, A.E. Arnold. 2013. Fungal endophytes in riparian areas of Arizona illustrate effects of microenvironment on symbiont assemblages. Fungal Ecology, in press.


Martinson*, E.O., K.C. Jander, Y.Q. Peng, H.H. Chen, C.A. Machado, A.E. Arnold, E.A. Herre. 2013. Relative investment in egg loads and poison sacs in fig wasps: implications for physiological mechanisms underlying seed and wasp production in figs. Acta Oecologica, in press.


Ortega, H. et al. 2013. Mycoleptodiscin A and B, cytotoxic alkaloids from the endophytic fungus Mycoleptodiscus sp. F0194. Journal of Natural Products, 76: 741-744.


Wanigesekara, A., E.M. Kithsiri Wijeratne, A.E. Arnold, and A.A. Leslie Gunatilaka. 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, in press.


Tedersoo, L., A.E. Arnold, K. Hansen. 2013. Novel aspects in the life cycle and biotrophic interactions in the Pezizomycetes. Molecular Ecology 22: 1488-1493.


Bascom-Slack, C., A.E. Arnold, S.A. Strobel. 2012. Student-directed discovery of the plant microbiome and its products. Science 338: 485-486.


Milani*, N.A., D.P. Lawrence, A.E. Arnold, and H.D. vanEtten. 2012. Origin of pistatin demethylase (PDA) in the genus Fusarium. Fungal Genetics and Biology 49: 933-942.


Martinson*, E.O., E.A. Herre, C.A. Machado, A.E. Arnold. 2012. Culture-free survey reveals diverse fungal communities associated with figs (Ficus spp.) in Panama. Microbial Ecology 64: 1073-1084.


Gazis*, R., J. Miadlikowska, F. Lutzoni, A.E. Arnold, P. Chaverri. 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.


U’Ren*, J.M., F. Lutzoni, J. Miadlikowska, A. Laetsch, A.E. Arnold. 2012. Host and geographic structure of endophytic and endolichenic fungi at a continental scale. American Journal of Botany 99: 898-914.


Varughese, T., et al. 2012 Antifungal depsidone metabolites from Cordyceps dipterigena, an endophytic fungus antagonistic to the phytopathogen Giberella fujikoroi. Tetrahedron Letters 28: 1624-1626.


Molinar, E., et al. 2012. Coibanoles, a new class of meroterpenoids produced by Pycnoporus sanguineus. Tetrahedron Letters 53: 919-922.


Lawrence*, D.P., B.M. Pryor, S.B. Kroken, A.E. Arnold. 2011. Interkingdom horizontal gene transfer of a hybrid NRPS/PKS from bacteria to filamentous Ascomycota. PLoS ONE 6: e28231.


Wijeratne, E.M.K., et al. 2012. Geopyxins A-E, ent-kaurane diterpenoids from endolichenic fungi, Geopyxis aff. majalis and Geopyxis sp. AZ0066: structure-activity relationships of geopyxins and their analogues. Journal of Natural Products 75: 361-369.


Wang, X.N., B.P. Bashyal, E.M.K. Wijeratne, J.M. U’Ren, M. Gunatilaka, A.E. Arnold, A.A.L. Gunatilaka. 2011. Smardaesidins A-G, new isopimarane and 20-Nor-isopimarane diterpenoids isolated from Smardaea sp., endophyte of  Ceratodon purpureus. Journal of Natural Products 74: 2052-2061.


Moreno, E., et al. 2011. Chemical constituents of the new endophytic fungus Mycosphaerella sp. nov. and their antiparasitic activity. Natural Products Communications 6: 835-840.


Dalling, J.W., A.S. Davis, B.J. Schutte, A.E. Arnold. 2011. Seed survival in soil: integrating effects of predation, dormancy, and the soil microbial community. Journal of Ecology 99: 89-95.


Higgins* K.L., P.D. Coley, T.A. Kursar, A.E. Arnold. 2011. Culturing and direct PCR suggest prevalent host-generalism among fungal endophytes of tropical grasses. Mycologia 103: 247-260.


U’Ren*, J.M., F. Lutzoni, J. Miadlikowska, A.E. Arnold. 2010. Intensive sampling reveals ecological distinctiveness and continua among culturable symbiotrophic and saprotrophic Ascomycota in a montane forest. Microbial Ecology 60: 340-353.


Epps*, M.J. and A.E. Arnold. 2010. Diversity, abundance, and community network structure in sporocarp-associated beetle communities in the central Appalachian Mountains. Mycologia 102: 785-802.


Hoffman*, M.T. and A.E. Arnold. 2010. Diverse bacteria inhabit living hyphae of phylogenetically diverse foliar endophytes. Applied and Environmental Microbiology 76: 4063-4075.


Arnold, A.E., L.J. Lamit, M. Bidartondo, C. Gehring, H.S. Callahan. 2010. Interwoven branches of the plant and fungal trees of life. New Phytologist 185: 874-878.


Peay, K., M. Bidartondo, A.E. Arnold. 2010. Not every fungus is everywhere: scaling to the biogeo-graphy of fungal-plant interactions across roots, shoots, and ecosystems. New Phytologist 185: 878-882.


Parrent*, J.L., K. Peay, A.E. Arnold, L. Comas, P. Avis, A. Tuininga. 2010. Moving from pattern to process in fungal symbioses: linking functional traits, community ecology, and phylogenetics. New Phytologist 185: 882-886.


Maddison, D.R. and A.E. Arnold. 2009. A review of the Bembidion (Odontium) aenulum subgroup (Coleoptera: Carabidae), with description of a new species. Zootaxa 2214: 45-61.


Arnold, A.E., J. Miadlikowska, K.L. Higgins, S.D. Sarvate, P. Gugger, A. Way, V. Hofstetter, F. Kauff, F. Lutzoni. 2009. Hyperdiverse fungal endophytes and endolichenic fungi elucidate the evolution of major ecological modes in the Ascomycota. Systematic Biology 58: 283-297.


U'Ren*, J.M., J.W. Dalling, R.E. Gallery, D.R. Maddison, E.C. Davis, C.M. Gibson, A.E. Arnold. 2009. Diversity and evolutionary origins of fungi associated with seeds of a neotropical pioneer tree: a case study for analyzing fungal environmental samples. Mycological Research 113: 432-449.


Rodriguez, R., J. White, A.E. Arnold, R. Redman. 2009. Fungal endophytes: diversity and ecological roles. New Phytologist 182: 314-330.


Arnold, A.E. 2008. Endophytic fungi: hidden components of tropical community ecology. Tropical Forest Community Ecology. (S. Schnitzer and W. Carson, eds.) Blackwell Scientific, Inc. pp. 254-271.


Hoffman*, M. and A.E. Arnold. 2008. Geography and host identity interact to shape communities of endophytic fungi in cupressaceous trees. Mycological Research 112: 331-344.


Kluger*, C., J.W. Dalling, R. E. Gallery, E. Sanchez, C. Weeks-Galindo, A.E. Arnold. 2008. Prevalent host-generalism among fungi associated with seeds of four neotropical pioneer species. Journal of Tropical Ecology 24: 351-354.


Feldman*, T.S., H. O’Brien, A.E. Arnold. 2008. Moth dispersal of mycoparasites and endophytes associated with Claviceps paspali and the grass Paspalum (Poaceae). Microbial Ecology 56: 742-750.


Bruns, T., A.E. Arnold, K. Hughes. 2008. Fungal networks made of humans: UNITE, FESIN, and frontiers in fungal ecology. New Phytologist 177: 586-588.


Mejía*, L.C., E.I. Rojas, Z. Maynard, A.E. Arnold, S. Van Bael, G.J. Samuels, N. Robbins, E.A. Herre. 2008. Endophytic fungi as biocontrol agents of Theobroma cacao pathogens. Biological Control 46: 4-14.


Jiménez-Romero, C., E. Ortega-Barria, A.E. Arnold, L. Cubilla-Rios. 2008. Activity against Plasmodium falciparum of lactones isolated from the endophytic fungus Xylaria sp. Pharmaceutical Biology 10-11: 1-4.


Rojas, E.I., E.A. Herre, L.C. Mejía, A.E. Arnold, P. Chaverri, G.J. Samuels. 2008. Endomelanconiopsis, a new anamorph genus in the Botryosphaeriaceae. Mycologia 100: 760-775.


Horton, T.R., A.E. Arnold, T.D. Bruns. 2008. FESIN workshops at ESA: the mycelial network grows. Mycorrhiza, online at DOI 10.1007/s00572-008-0210-5


Wijeratne, E.M.K., P.A. Paranagama, M.T. Marron, M.K. Gunatilaka, A.E. Arnold, A.A.L. Gunatilaka. 2008. Sesquiterpene quinines and related metabolites from Phyllosticta spinarum, a fungal strain endophytic in Platycladus orientalis in the Sonoran Desert. Journal of Natural Products 71:218-222.


Arnold, A.E. 2007. Understanding the diversity of foliar fungal endophytes: progress, challenges, and frontiers. Fungal Biology Reviews 21: 51-66.


Arnold, A.E. and F. Lutzoni. 2007. Diversity and host range of foliar fungal endophytes: Are tropical leaves biodiversity hotspots? Ecology 88: 541-549.


Arnold, A.E., D.A. Henk, R.L. Eells, F. Lutzoni, R. Vilgalys. 2007. Diversity and phylogenetic affinities of fungal endophytes in loblolly pine inferred by culturing and environmental PCR. Mycologia 99: 185-206.


Arnold, A.E. and B.M.J. Engelbrecht. 2007. Fungal endophytes nearly double minimum leaf conductance in seedlings  of a tropical tree. Journal of Tropical Ecology 23: 369-372.


Gallery*, R., J.W. Dalling, A.E. Arnold. 2007. Diversity, host affinity, and distribution of seed-infecting fungi:  A case study with neotropical Cecropia. Ecology 88: 582-588.


Gallery*, R.E., J.W. Dalling, B. Wolfe, A.E. Arnold. 2007. Role of seed-infecting fungi in the recruitment limitation of neotropical pioneer species. Seed dispersal: Theory and its application in a changing world. Eds. A. Dennis, R. Green, E. Schupp and D. Westcott. CABI Press. pp. 479-498.


Tsai*, Y.J, K. Maloney, A.E. Arnold. 2007. Abiotic and biotic factors influencing the distribution of Pyrgulopsis thompsoni, the Huachuca springsnail. Journal of Aquatic Ecology 22: 213-218.


Agrawal, A.A., D.A. Ackerly, F. Adler, A.E. Arnold, and 16 others. 2007. Filling key gaps in population and community ecology. Frontiers in Ecology and the Environment 5: 145-152.


Higgins*, K.L., A.E. Arnold, J. Miadlikowska, S.D. Sarvate, F. Lutzoni. 2007. Phylogenetic relationships, host affinity, and geographic structure of boreal and arctic endophytes from three major plant lineages. Molecular Phylogenetics and Evolution 42: 543-555.


Paranagama, P.A., E.M.K. Wijeratne, M.K. Gunatilaka, A.E. Arnold, A.A.L. Gunatilaka. 2007. Bioactive and other naphthopyrans from Corynespora sp. occurring in Usnea cavernosa: first report of metabolites of an endolichenic fungus. Journal of Natural Products 70: 1700-1705


Miadlikowska, J.M., et al. 2006. New insights from nuclear ribosomal and protein coding genes into classification and evolution of the Lecanoromycetes (Pezizomycotina, Ascomycota). Mycologia 98: 1088-1103.


James, T.Y., et al. 2006. Reconstructing the early evolution of the fungi using a six-gene phylogeny. Nature 443: 818-822.


Spatafora, J.W., et al. 2006. A five-gene phylogenetic analysis of the Pezizomycotina. Mycologia 98: 1018-1028.


Arnold, A.E. and L.C. Lewis. 2005.  Evolution of fungal endophytes, and their roles against insects. Ecological and Evolutionary Advances in Insect-Fungus Associations (F. Vega and M. Blackwell, eds.).  Oxford University Press, Oxford. pp. 74-96.


Lutzoni, F., et al. 2004. Assembling the fungal tree of life: progress, classification, and the evolution of subcellular traits. American Journal of Botany 91: 1446-1480.


Arnold*, A.E., L. Mejía, D. Kyllo, E. Rojas, Z. Maynard, N. Robbins, E.A. Herre. 2003. Fungal endophytes limit pathogen damage in a tropical tree. Proceedings of the National Academy of Sciences USA 100: 15649-15654.


Arnold*, A.E. and E.A. Herre. 2003.  Canopy cover and leaf age affect colonization by tropical fungal endophytes: Ecological pattern and process in Theobroma cacao (Malvaceae). Mycologia 95: 388-398.


Arnold*, A.E. and N.M. Asquith.  2002.  Herbivory in a fragmented tropical forest:  patterns from islands at Lago Gatún, Panamá.  Biodiversity and Conservation 11: 1663-1680.


Arnold*, A.E., Z. Maynard, G. Gilbert.  2001.  Fungal endophytes in dicotyledonous neotropical trees: patterns of abundance and diversity.  Mycological Research 105: 1502-1507.


Arnold*, A.E., Z. Maynard, G. Gilbert, P.D. Coley, T.A. Kursar.  2000.  Are tropical fungal endophytes hyperdiverse?  Ecology Letters 3: 267-274.


Asquith*, N.M., J. Terborgh, A.E. Arnold, M. Riveros-C. 1999. The fruits the agouti ate:  Hymenaea courbaril seed fate when its disperser is absent: evidence from Lago Guri, Venezuela. Journal of Tropical Ecology 15: 229-235.


Book chapters, technical reports, and popular articles (not peer-reviewed)

Arnold, A.E. 2008. Hidden within our botanical richness, a treasure trove of fungal endophytes. 2008. The Plant Press 32: 13-15. pdf

Bashyal, B.P., A.M. Burns, M.X. Liu, P.A. Paranagama, C.J. Seliga, T.J. Turbyville, E.M.K. Wijeratne, J. Zhan, M.K. Gunatilaka, A.E. Arnold, S.H. Faeth, L. Whitesell, and A.A.L. Gunatilaka. 2007. Discovery of small molecule bioactive agents from endophytic fungi of the Sonoran Desert. Proceedings of the 6th International Symposium on Fungal Endophytes of Grasses. Grassland Research and Practice Series No. 13, New Zealand Grassland Association, Christchurch. pdf

Arnold, A.E. 2005. Diversity and Ecology of fungal endophytes in tropical forests. Current Trends in Mycological Research (S. Deshmukh, ed.). Oxford & IBH Publishing Co. Pvt. Ltd., New Delhi:  pp. 49-68. pdf

Van Bael, S.A., Z. Maynard, N. Robbins, J.F. Bischoff, A.E. Arnold, E. Rojas, L.C. Mejía, D.A. Kyllo, and E.A. Herre. 2005.  Emerging perspectives on the ecological roles of endophytic fungi in tropical plants. The Fungal Community: its organization and role in the ecosystem (J.F. White, Jr., J. Dighton, & P. Oudemans, eds). 3rd Edition. Marcel-Dekker, New York, NY.  Pp. 181-192. Reprint available on request.

Herre, E.A., S. A. Van Bael, Z. Maynard, N. Robbins, J.  Bischoff, A. E. Arnold, E. Rojas, L. C. Mejia, R. A. Cordero, C. Woodward, and D.A. Kyllo. 2005. Tropical plants as chimera: some implications of foliar endophytic fungi for the study of host plant defense, physiology, and genetics. Biotic Interactions in the Tropics (D.F.R.P. Burslem, M.A. Pinard, & S.E. Hartley, eds.) Cambridge University Press. pp. 226-237. Reprint available on request.

Mejia, L. C., E. Rojas, Z. Maynard, A.E. Arnold, D. Kyllo, N. Robbins, and E.A. Herre. 2003. Inoculation of beneficial endophytic fungi into Theobroma cacao tissues. In Proceedings of the 14th International Cocoa Research Conference, Accra-Ghana. Reprint available on request.

Arnold, A.E.  2001.  Fungal endophytes in neotropical trees:  Abundance, diversity, and ecological interactions.  In Tropical Ecosystems: Structure, Diversity and Human Welfare (K. N. Ganeshaiah, R. Uma Shaanker, & K. S. Bawa, eds.).  Oxford & IBH Publishing Co. Pvt. Ltd., New Delhi:  pp. 739-743. pdf

Arnold, A. E., L. Mejía, E. Rojas, Z. Maynard, N. Robbins and E. A. Herre.  2001.  Organismos endofíticos:  Microorganismos en plantas.  (Endophytic organisms:  microorganisms in plants.) El Uso de Microorganismos Benéficos en la Agricultura Moderna.  Universidad E.A.R.T.H., Guácimo, Limón, Costa Rica. Reprint (in Spanish) available on request.

Arnold, A.E.  1999.  Fungal endophytes of tropical trees: methods and potential for biological control of fungal  pathogens of cocoa.  Proceedings of the Research Methodology of Biocontrol of Plant Diseases Workshop, San Jose: CATIE. Reprint available on request.

Arnold, A.E.  1999.  Sustainable cocoa: the fungal community component. American Cocoa Research Institute Features in Integrated Pest Management for Cocoa:  Online:

Arnold, A.E.  1998.  Various entries, Encyclopaedia Britannica Online Rainforest Resources CD.

Arnold, A.E.  1993.  Flower color variation and morph-specific outcrossing in morning glories polymorphic for corolla color. Vertices (Duke University).