BarkingTreefrogIf you would like a reprint, please email me, or check my researchgate profile.  Also, my complete publication list is available via my google scholar profile.






Escallón, C., L.K. Belden, and I.T. Moore.  2019. The cloacal microbiome changes with the breeding season in a wild bird. Integrative Organismal Biology 1: oby009.

Kueneman, J., M. Bletz, V. McKenzie, C.G. Becker, M. Joseph, J. Abarca, H. Archer, A. Arellano, A. Bataille, M. Becker, L. Belden, A. Crottini, R. Geffers, C. Haddad, R. Harris, W. Holden, M. Hughey, M. Jarek, P. Kearns, J. Kerby, J. Kielgast, A. Kurabayashi, A. Longo, A. Loudon, D. Medina, J. Núnez, R.G. Perl, A. Pinto-Tomás, F. Rabemananjara, E. Rebollar, A. Rodriguez, L. Rollins-Smith, R. Stevenson, C. Tebbe, G. Vargas Asensio, B. Waldman, J. Walke, S. Whitfield, K. Zamudio, I. Zúñiga Chaves, D. Woodhams, and M. Vences. 2019. Community richness of amphibian skin bacteria correlates with bioclimate at the global scale. Nature Ecology & Evolution 3: 381–389.

Rebollar, E.A., T. Bridges, M.C. Hughey, D. Medina, L.K. Belden, and R.N. Harris. 2019. Integrating the role of antifungal bacteria into skin symbiotic communities of three Neotropical frog species.  The ISME Journal: published online 13 March 2019

Medina, D., M.C. Hughey, J.B.  Walke, M.H. Becker, K. Pontarelli, S. Sun, B. Badgley, and L.K. Belden. 2019. Amphibian skin fungal communities vary across host species and regions, but do not correlate with infection by a pathogenic fungus. Environmental Microbiology, online

Estrada, A., M.C. Hughey, D. Medina, E.A. Rebollar, R.N. Harris, and L.K. Belden.  2019. Skin bacterial communities of neo-tropical treefrog populations vary with local environmental conditions at the time of sampling. PeerJ 7:e7044


Rebollar, E.A., A. Gutiérrez-Preciado, C. Noecker, A. Eng, M.C. Hughey, D. Medina, J.B. Walke, E. Borenstein, R.V. Jensen, L.K. Belden, and R.N. Harris. 2018. The skin microbiome of the Neotropical frog Craugastor fitzingeri: Inferring potential bacterial-host-pathogen interactions from metagenomic data. Frontiers in Microbiology 9: 466. doi: 10.3389/fmicb.2018.00466

Hopkins, S.R., C.M. McGregor, L.K. Belden, and J.M. Wojdak.  2018. Handling times and saturating transmission functions in a snail-worm symbiosis. Oecologia 188: 277-287.


Orlofske, S.A., L.K. Belden, and W.A. Hopkins. 2017. Effects of Echinostoma trivolvis metacercariae infection during development and metamorphosis of the wood frog (Lithobates sylvaticus) Comparative Biochemistry and Physiology, A 203: 40-48.

Hopkins, S.R., J.M. Wojdak, and L.K. Belden. 2017. Defensive symbionts mediate host-parasite interactions at multiple scales. Trends in Parasitology 33: 53-64. DOI:

Zemmer, S.A., J. Wyderko, I. Cedillos, L. Clay, J. Da Silva Neto, E.F. Benfield, and L.K. Belden. 2017.  Seasonal and annual variation in trematode infection of stream-dwelling snails, Elimia proxima, in the southern Appalachian mountains of Virginia. Journal of Parasitology 103: 213-220. DOI:

Thomason, C.A., A. Leon, L. Kirkpatrick, L.K. Belden and D.M. Hawley. 2017. Eye of the finch: characterization of the ocular microbiome of Haemorhous mexicanus in relation to mycoplasmal conjunctivitis. Environmental Microbiology, 14: 1439-1449.  DOI: 10.1111/1462-2920.13625

Medina, D., M.C. Hughey, M.H. Becker, J.B. Walke, T.P. Umile, E. Burzynski, A. Iannetta, K.P.C. Minbiole, and L.K. Belden.  2017. Variation in metabolite profiles of amphibian skin bacterial communities across elevations in the Neotropics. Microbial Ecology 74: 227-238.  doi:10.1007/s00248-017-0933-y

Hughey, M.C., J. Delia, and L.K. Belden. 2017. Diversity and stability of egg bacterial assemblages: The role of paternal care in the glassfrog, Hyalinobatrachium colymbiphyllum. Biotropica 49: 792-802. doi:10.1111/btp.12461

Walke, J.B., M.H. Becker, M.C. Hughey, M.C. Swartwout, R.V. Jensen, and L.K. Belden. 2017.  Dominance-function relationships in the amphibian skin microbiome. Environmental Microbiology 19: 3387-3397. doi: 10.1111/1462-2920.13850

Medina, D., J.B. Walke, Z. Gajewski, M.H. Becker, M.C. Swartwout, and L.K. Belden. 2017. Culture media and individual hosts affect the recovery of culturable bacterial diversity from amphibian skin. Frontiers in Microbiology 8: 1574. doi: 10.3389/fmicb.2017.01574

Hughey, M.C., J.A. Peña, R. Reyes, D. Medina, L.K. Belden, and P.A. Burrowes. 2017. Skin bacterial microbiome of a generalist Puerto Rican frog varies along elevation and land use gradients. PeerJ 5: e3688. DOI: 10.7717/peerj.3688

Thomason, C.A., N. Mullen, L.K. Belden, M. May, and D.M. Hawley. 2017. Resident microbiome disruption with antibiotics enhances virulence of a colonizing pathogen. Scientific Reports 7: 16177. doi:10.1038/s41598-017-16393-3


Rebollar, E.A., M.C. Hughey, D. Medina, R.N. Harris, R. Ibañez, and L.K. Belden.  2016. Skin bacterial diversity of Panamanian frogs is associated with host susceptibility and presence of Batrachochytrium dendrobatidis.  The ISME Journal 10: 1682-1695. doi:10.1038/ismej.2015.234

Rebollar, E.A., R.E. Antwis, L.K. Belden, M.H. Becker, M.C. Bletz, R. Brucker, X. Harrison, M.C. Hughey, J. Kueneman, A.H. Loudon, V. McKenzie, D. Medina, K.P.C. Minbiole, L. Rollins-Smith, J.B. Walke, S. Weiss, D.C. Woodhams and R.N. Harris.  2016. Using “omics” and multi-omics approaches to guide probiotic selection to mitigate chytridiomycosis and other emerging infectious diseases. Frontiers in Microbiology 7:68 doi:10.3389/fmicb.2016.00068

Hopkins, S.R., J. Ocampo, J.M. Wojdak, and L.K. Belden. 2016. Host community composition and defensive symbionts determine trematode parasite abundance in host communities. Ecosphere 7(3):e01278.

Hughey, M.C., J.B. Walke, M.H. Becker , T.P. Umile, E. Burzynski, K.P.C. Minbiole, A. Iannetta, C.N. Santiago, W.A. Hopkins, and L.K. Belden. 2016. Impact of short-term exposure to coal combustion waste on the skin microbiome of spring peepers, Pseudacris crucifer. Applied and Environmental Microbiology 82(12):3493-3502. doi:10.1128/AEM.00045-16  

Walke, J.B. and L.K. Belden. 2016. Harnessing the microbiome to prevent fungal infections: lessons from amphibians. PLoS Pathogens 12(9): e1005796. doi:10.1371/journal.ppat.1005796

Escallón, C., M.H. Becker, J.B. Walke, R.V. Jensen, G. Comier, L.K. Belden and I.T. Moore.  2016. Cloacal bacterial phylogenetic diversity, and relative abundance of Chlamydiae, increases with testosterone levels in a free-living tropical bird. Functional Ecology 31: 192-203. doi: 10.1111/1365-2435.12696


Woodhams D.C., R. Alford, R.E. Antwis, H. Archer, M.H. Becker, L.K. Belden, S.C. Bell, M. Bletz, J.H. Daskin, L.A. Davis, S.V. Flechas, A. Lauer, A.G. Peña, R.N. Harris, W.M. Holden, M.C. Hughey, R. Ibañez, R. Knight, J. Kueneman, F. Rabemanajara, L.K. Reinert, L.A. Rollins-Smith, F. Roman-Rodriguez, S.D. Shaw, J.B. Walke, and V. McKenzie. 2015. Antifungal isolates database to centralize data on amphibian skin-associated bacteria and their function against emerging fungal pathogens including Batrachochytrium dendrobatidis. Ecology 96: 595.

Becker, M.H, J.B. Walke, S. Cikanek, A.E. Savage, N. Mattheus, K.P.C. Minbiole, T. Umile, R.N. Harris, L.K. Belden, and B. Gratwicke. 2015. Composition of symbiotic bacteria as a predictor of survival in Panamanian golden frogs infected with Batrachochytrium dendrobatidis. Proceedings of the Royal Society, B 282: 20142881.

Becker, M.H., J.B. Walke, L. Murrill*, D.C. Woodhams, L.K. Reinert, L.A. Rollins-Smith, T.P. Umile, K.P.C. Minbiole, and L.K. Belden. 2015. The phylogenetic distribution of symbiotic bacteria from Panamanian amphibians that inhibit growth of the lethal fungal pathogen Batrachochytrium dendrobatidis. Molecular Ecology 24: 1628-1641. doi: 10.1111/mec.13135.

Hopkins, S.R., L.J. Boyle*, L.K. Belden and J.M. Wojdak. 2015. Dispersal of a defensive symbiont depends on contact between hosts, host size and resource availability Oecologia 179: 307-318.

Wyderko, J.A, E.F. Benfield, J. Maerz, K. Cecala, and L.K. Belden. 2015. Variable infection of stream salamanders in the southern Appalachians by the trematode Metagonimoides oregonensis (family: Heterophyidae). Parasitology Research 114(8): 3159-3165.

Walke, J.B., M.H. Becker , M.C. Hughey, M.C. Swartwout*, R.V. Jensen, and L.K. Belden. 2015. Most of the dominant members of amphibian skin bacterial communities can be readily cultured. Applied and Environmental Microbiology 81: 6589-6600.

Walke, J.B., M.H. Becker, S.C. Loftus, L.L. House, T. Teotonio, K.P.C. Minbiole, and L.K. Belden. 2015. Community structure and function of amphibian skin microbes: an experiment with bullfrogs exposed to a chytrid fungus. PLoS ONE 10(10): e0139848. doi:10.1371/journal.pone.0139848.

Belden, L.K., M.C. Hughey, E.A. Rebollar, T.P. Umile, S.C. Loftus, E.A. Burzynski, K.P.C. Minbiole, L.L. House, R.V. Jensen, M.H. Becker, J.B. Walke, D. Medina, R. Ibáñez, and R.N. Harris. 2015. Panamanian frog species host unique skin bacterial communities. Frontiers in Microbiology 6:1171 doi: 10.3389/fmicb.2015.01171


C.L. Searle, L.K. Belden, P. Du, and A.R. Blaustein. 2014. Stress and chytridiomycosis: Exogenous exposure to corticosterone does not alter amphibian susceptibility to a fungal pathogen. Journal of Experimental Zoology Part A: Ecological Genetics and Physiology 321A: 243-253.

E.A. Rebollar, M.C. Hughey, R.N. Harris, R.J. Domangue, D. Medina, R. Ibáñez, and L.K. Belden. 2014. The lethal fungus Batrachochytrium dendrobatidis is present in lowland tropical forests of far Eastern Panamá. PLoS ONE 9(4): e95484. doi: 10.1371/journal.pone.0095484.

Walke, J.B., M.H. Becker, S.C. Loftus, L.L. House, G. Cormier, R.V. Jensen and L.K. Belden. 2014. Amphibian skin may select for rare environmental microbes. The ISME Journal: doi:10.1038/ismej.2014.77.

Becker, M.H., C.L. Richards-Zawacki, B. Gratwicke and L.K. Belden. 2014. The effect of captivity on the cutaneous bacterial community of the critically endangered Panamanian golden frog, Atelopus zeteki. Biological Conservation 176: 199-206.

Wojdak, J.M., R.M. Edman*, J.A.Wyderko*, S.A. Zemmer, and L.K. Belden. 2014. Host density and competency determine host diversity effects on infection by trematode parasites PLoS ONE: 9(8): e105059. doi:10.1371/journal.pone.0105059.

Hughey, M.C., M.H. Becker, J.B. Walke, M.C. Swartwout*, and L.K. Belden. 2014. Batrachochytrium dendrobatidis in Virginia amphibians: within and among site variation in infection. Herpetological Review 45(1): 1-5.

Ellison, A.R., T. Tunstall, M.C. Hughey, E.A. Rebollar, L.K. Belden, R.N. Harris, K.R. Lips and K.R. Zamudio. 2014. More than skin deep: functional genomic basis for resistance to amphibian chytridiomycosis. Genome Biology and Evolution doi:10.1093/gbe/evu285.


Orlofske, S.A., L.K. Belden and W.A. Hopkins. 2013. Larval wood frog (Rana [=Lithobates] sylvatica) development and physiology following gradual exposure to the trematode parasite, Echinostoma trivolvis. Comparative Biochemistry and Physiology, Part A: Molecular and Integrative Physiology 164: 529-536.

Chambers, D.L., J.M. Wojdak, P. Du and L.K. Belden. 2013. Pond acidification may explain differences in corticosterone among salamander populations. Physiological and Biochemical Zoology 86: 224-232.

Wojdak, J.M., L. Clay*, S. Moore*, T. Williams*, and L.K. Belden. 2013. Echinostoma trivolvis (Digenea; Echinostomatidae) second intermediate host preference matches host-suitability. Parasitology Research 112: 799-805.

Hopkins, S.R.*, J.A. Wyderko*, R. Sheehy, L.K. Belden, and J.M. Wojdak. 2013. Parasite predators exhibit a rapid numerical response to increased parasite abundance and reduce transmission to hosts. Ecology and Evolution 3: 4427-4438.


Rubbo, M.J., L.K. Belden, S.I. Storrs, J.J. Cole, and J.M. Kiesecker. 2012. Species loss in the brown world: are heterotrophic systems inherently stable? Aquatic Sciences 74: 397-404.

Blaustein, A.R., S.S. Gervasi, P.T.J. Johnson, J.T. Hoverman, L.K. Belden, P.W. Bradley and G.Y. Xie. 2012. Ecophysiology meets conservation: understanding the role of disease in amphibian population declines. Philosophical Transactions of the Royal Society, B 367: 1688-1707.

Belden, L.K., W.E. Peterman, S.A. Smith, L.R. Brooks*, E. F. Benfield, W.P. Black, Z. Yang, and J.M. Wojdak. 2012. Metagonimoides oregonensis (Digenea, Heterophyidae) infection in pleurocerid snails and Desmognathus quadramaculatus salamander larvae in southern Appalachian streams. Journal of Parasitology 98: 760-767.

Detwiler, J.T., A.M. Zajac, D.J. Minchella, and L.K. Belden. 2012. Revealing cryptic diversity in a definitive host: Echinostomes in muskrats. Journal of Parasitology 98: 1148-1155.


Sokol, E.R., E.F. Benfield, L.K. Belden, and H.M. Valett. 2011. The assembly of ecological communities inferred from taxonomic and functional composition. American Naturalist 177: 630-644.

Belden, L.K. and J.M. Wojdak. 2011. The combined influence of trematode parasites and predatory salamanders on wood frog (Rana sylvatica) tadpoles. Oecologia 166: 1077-1086.

Chambers, D.L., J.M. Wojdak, P. Du and L.K. Belden. 2011. Corticosterone levels during larval development in the amphibians Rana sylvatica and Ambystoma jeffersonianum reared under laboratory, mesocosm, or free-living conditions. Copeia: 530-538.


Searle, C.L., L.K. Belden, B.A. Bancroft, B.A. Han, L.M. Biga and A.R. Blaustein. 2010. Experimental examination of the effects of ultraviolet-B radiation in combination with other stressors on frog larvae. Oecologia 162: 237-245.

Belden, L.K., J.C. Wingfield and J.M. Kiesecker. 2010. Variation in the hormonal stress response among larvae of three amphibian species. Journal of Experimental Zoology Part A: Ecological Genetics and Physiology 313A: 524-531.

Keesing, F., L.K. Belden, P. Daszak, A. Dobson, D. Harvell, R.D. Holt, P. Hudson, A. Jolles, K. Jones, C. Mitchell, T. Bogich and R.S. Ostfeld. 2010. Impacts of biodiversity on the emergence and transmission of infectious diseases. Nature 468: 647-652.


Budischak, S.A., L. K. Belden, and W.A. Hopkins. 2009. The relative toxicity of malathion to trematode-infected and non-infected Rana palustris tadpoles. Archives of Environmental Contamination and Toxicology 56: 123-128.

Orlofske, S.A., L.K. Belden and W.A. Hopkins. 2009. Moderate Echinostoma trivolvis infection has no effects on physiology and fitness-related traits of larval pickerel frogs (Rana palustris). Journal of Parasitology 95: 787-792.

Belden, L.K., P.D. Widder, L.R. Fischer*, A.B. Carter*, J.M. Wojdak. 2009. Hatching of Echinostoma trivolvis miracidia in response to snail host and non-host chemical cues. Parasitology Research 105: 883-885.


Budischak, S.A., L. K. Belden, and W.A. Hopkins. 2008. The effects of malathion on embryonic development and latent susceptibility to trematode parasites in ranid tadpoles. Environmental Toxicology and Chemistry 27: 2496-2500.

Griggs, J.L. and L.K. Belden. 2008. Effects of atrazine and metolachlor on the survivorship and infectivity of Echinostoma trivolvis trematode cercariae. Archives of Environmental Contamination and Toxicology 54: 195-202.

Rubbo, M.J., L.K. Belden, and J.M. Kiesecker. 2008. Differential responses of aquatic consumers to terrestrial carbon subsidies. Hydrobiologia 605:37-44.


Belden, L.K. and R.N. Harris. 2007. Infectious diseases in wildlife: the community ecology context. Frontiers in Ecology and the Environment 10(5): 533-539.

Belden, L.K., M.J. Rubbo, J.C. Wingfield and J.M. Kiesecker. 2007. Searching for the physiological mechanism of density-dependence: does corticosterone regulate tadpole responses to density? Physiological and Biochemical Zoology 80(4): 444-451.

Culp, C.E., J.O. Falkinham III and L.K. Belden. 2007. Identification of the natural bacterial microflora on the skin of Eastern newts, bullfrog tadpoles and Redback salamanders. Herpetologica 63: 66-71.


Belden, L.K. 2006. Impact of eutrophication on wood frog, Rana sylvatica, tadpoles infected with Echinostoma trivolvis cercariae. Canadian Journal of Zoology 84(9): 1315-1321.

Rubbo, M.J., R.S. Mirza, L.K. Belden, J.J. Falkenbach V, S. I. Storrs and J.M. Kiesecker. 2006. Evaluating a predator-prey interaction in the field: the interaction between beetle larvae (predator) and tadpoles (prey). Journal of Zoology 269: 1-5.


Belden, L.K. and J.M. Kiesecker. 2005. Glucocorticosteroid treatment of larval treefrogs (Hyla versicolor) increases infection by Alaria sp. trematode cercariae. Journal of Parasitology 91(3): 686-688.

Belden, L.K., I.T. Moore, J.C. Wingfield and A.R. Blaustein. 2005. Corticosterone and growth in Pacific treefrog (Hyla regilla) tadpoles. Copeia 2005(2): 324-330.


Kiesecker, J.M., L.K. Belden, K. Shea and M. J. Rubbo. 2004. Amphibian decline and emerging disease. American Scientist 92: 138-147.


Belden, L.K., I.T. Moore, R.T. Mason, J.C. Wingfield and A.R. Blaustein. 2003. Survival, the hormonal stress response, and UV-B avoidance in Cascades frog tadpoles (Rana cascadae) exposed to UV-B radiation. Functional Ecology 17: 409-416.

Blaustein, A.R. and L.K. Belden. 2003. Amphibian defenses against UV-B radiation. Evolution and Development 5: 89-97.


Belden, L.K. and A.R. Blaustein. 2002. Population differences in sensitivity to UV-B radiation for larval long-toed salamanders. Ecology 83: 1586-1590.

Belden, L.K. and A.R. Blaustein. 2002. Exposure of red-legged frog embryos to ambient UV-B radiation in the field negatively affects larval growth and development. Oecologia 130: 551-554.

Belden, L.K. and A.R. Blaustein. 2002. UV-B induced skin darkening in larval salamanders does not prevent sublethal effects of exposure on growth. Copeia 2002: 748-754.


Blaustein, A.R., L.K. Belden, D.H. Olson, D.M. Green, T.L. Root and J.M. Kiesecker. 2001. Amphibian breeding and climate change. Conservation Biology 15: 1804-1809.

Marco, A., D. Cash, L.K. Belden, and A.R. Blaustein. 2001. Sensitivity to urea fertilization in three amphibian species. Archives of Environmental Contamination and Toxicology 40:406-409.

Hatch A.C., L.K. Belden, E.A. Scheessele and A.R. Blaustein. 2001. Juvenile amphibians do not avoid potentially lethal levels of urea on soil substrate. Environmental Toxicology and Chemistry 20: 2328-2335.

Blaustein, A.R., E.L. Wildy, L.K. Belden, and A. Hatch. 2001. The influence of abiotic and biotic factors on amphibians in ephemeral ponds with special reference to long-toed salamanders (Ambystoma macrodactylum). Israel Journal of Zoology 47: 333-345.

Kiesecker, J.M., A.R. Blaustein, and L.K. Belden. 2001. Complex causes of amphibian population declines. Nature 410: 681-684.


Belden, L.K., E.L. Wildy, and A.R. Blaustein. 2000. Growth, Survival and Behaviour of Larval Long-toed Salamanders (Ambystoma macrodactylum) exposed to Ambient Levels of UV-B radiation. Journal of Zoology 251: 473-479.

Belden, L.K., E.L. Wildy, A.C. Hatch, and A.R. Blaustein. 2000. Juvenile western toads, Bufo boreas, avoid chemical cues of snakes fed juvenile, but not larval, conspecifics. Animal Behaviour 59: 871-875.


DeVito, J., D.P. Chivers, J.M. Kiesecker, L.K. Belden and A.R. Blaustein. 1999. Effects of snake predation on aggregation and metamorphosis of Pacific treefrog (Hyla regilla) larvae. Journal of Herpetology 33: 504-507.

Chivers, D.P., J.M. Kiesecker, E.L. Wildy, L.K. Belden, L.B. Kats, and A.R. Blaustein. 1999. Avoidance response of post-metamorphic anurans to cues of injured conspecifics and predators. Journal of Herpetology 33: 472-476.


Blaustein, A.R., J.M. Kiesecker, D.P. Chivers, D. G. Hokit, A. Marco, L.K. Belden and A. Hatch. 1998. Effects of Ultraviolet Radiation on Amphibians: Field Experiments. American Zoologist 38: 799-812.