James (Eddie) Watkins

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James (Eddie) Watkins

Professor of Biology

Department/Office Information

129 Ho Science Center


The work in my lab focuses on organismal plant ecology and physiology with a primary focus on ferns and their allies. Lab members work across many disciplines to include molecular biology, systematics, ecology, physiology, anatomy, and reproductive biology. I run a diverse ship, but the lab mainly focuses on how the natural world shapes the distribution of species in both modern landscapes and over evolutionary time frames. Central to our work are studies on plant stress physiology and developing a better understanding of the mechanisms behind stress tolerance and recovery.

Learn more about my lab

BS, Auburn University, 1997
MS, Iowa State University, 2000
PhD, University of Florida, 2006
Post Doc, Harvard University, 2008

  • OTS Specialty Course: Ferns and lycophytes of Costa Rica, Harvard University
  • Graduate teaching assistantships at University of Florida, Iowa State University, and Auburn University

Biological sciences; botany; plant ecology; plant physiology

2018: Understanding the effects of ploidy level on responses to global change in plants. $760,800 (with Emily Sessa, University of Florida and Clayton Visger, Sacramento State). NSF Award Number: 1725458

2017: The Cenozoic radiation of Eupolypod ferns: did selection for drought tolerance drive the evolutionary physiology of sporophytes and gametophytes? $642,000 (with J.Pitterman, University of California, Santa Cruz). NSF Award Number: 1656801

2013: Novel lessons from ancient plants: water transport in the earliest tracheophytes. $380,000 (with J. Pitterman, University of California, Santa Cruz) NSF Award Number: 1258186

2010: Finnish Cultural Foundation: Examination of functional traits of ferns across a rainfall gradient in Panama.Co-PI Mirkka Jones, University of Turku, Finland (€6,000)

  • 2012-2014: President of the American Fern Society
  • 2010-2014: Council member to the American Institute of Biological Sciences to represent the American Association of Plant Taxonomists
  • 2008-Current: Associate Editor of the America Fern Journal
  • 2008-2014: Council member to the American Institute of Biological Sciences to represent the American Fern Society.
  • 2007-2009: Secretary of the Pteridological Section of the Botanical Society of America

Book Chapters

Pitterman, J.P., J.E. Watkins, Jr. and E. Schuettpelz. 2015. The Structure and Function of Xylem in Seed-Free Vascular Plants: An Evolutionary Perspective. In Functional and

Ecological Xylem Anatomy. Editor U. Hacke. Springer Farrar, D.R., C. Dassler, J.E. Watkins, Jr., C. Skelton, 2008. Gametophyte Ecology. In Biology and Evolution of Ferns and Lycophytes. Editors, T. Ranker and C. Haufler. Cambridge University Press. Pp 235-267.

Refereed Publications

(*Represents undergraduate student author)


Watts, J.L. and J.E.Watkins, Jr. In Press. Crossing the Divide: An Exploration of Functional Traits in Ferns
that Grow Across Terrestrial, Epipetric, and Epiphytic Habitats. American Fern Jounrnal

Nitta, J.H., J.E. Watkins, Jr., N.M. Holbrook, R. Taputuarai, T. Wang, C.C. Davis. 2021. Ecophysiological differentiation between life stages in filmy ferns (Hymenophyllaceae). Journal of Plant Research. oi: 10.1007/s10265-021-01318-z

Campany, C., J. Pittermann, A. Baer, H. Holmlund, E. Schuettpelz, K. Mehltreter, J.E. Watkins, 2021. "Leaf water relations in epiphytic ferns are driven by drought avoidance rather than tolerance mechanisms" Plant Cell and Environment 44(6):1741-1755


Ondřej H., W.L. Testo  E. B. Sessa  J.E. Watkins Jr.,  C. E. Campany, J. Pittermann  L. Ekrt. 2020. Insights into the evolutionary history and widespread occurrence of antheridiogen systems in ferns. New Phytologist. https://doi.org/10.1111/nph.16836

Nitta, J.H., J. E. Watkins, Jr., R. Taputuarai, and C. Davis. 2020. Life in the canopy: community trait assessments reveal previously undetected diversity among fern epiphytes. New Phytologist https://doi.org/10.1111/nph.16607

Kattge, J., G. Bönisch, S. Díaz, S. Lavore, I. C. Prentice, P. Leadley, S. Tautenhahn… J.E. Watkins, Jr. with 729 other authors. 2020. TRY plant trait database – enhanced coverage and open access. Global Change Biology


Watts*, J.L., R.C. Moran, J.E. Watkins, Jr. 2019. The first report of hemiepiphytism in Aspleniaceae. Annals of Botany. https://doi.org/10.1093/aob/mcz124. Editor’s Choice Award

Watkins, Jr., J.E. and R.C. Moran. 2019. The gametophytes of the fern genera Dracoglossum and Lomariopsis (Lomariopsidaceae) and their phylogenetic significance. International Journal of Plant Sciences. doi/10.1086/705587

Li-Yaung, K., Y. Chang, Y. Huang, W.L. Testo, A. Ebihara, G. Rouhan, L.Quintanilla, J. E. Watkins Jr, F. Li, Y. Huang. 2019. A global phylogeny of Stegnogramma sensu lato ferns (Thelypteridoideae, Thelypteridaceae, Aspleniineae): generic and section-level revision, historical biogeography, and evolution of leaf architecture. Cladistics. Online Early at https://doi.org/10.1111/cla.12399

Krieg, C.P., J.E. Watkins, Jr., K.A. McCulloh. 2019. A new protocol for psychometric pressure-volume curves of fern gametophytes. Applications in Plant Sciences 7(5): e1248. doi:10.1002/aps3.1248


Campany, C., J.E. Watkins, Jr. and *Martin, L. 2018. Convergence of ecophysiological traits drives floristic composition of early lineage vascular plants in a tropical forest floor. Annals of Botany. 123(5):793–803.  https://doi.org/10.1093/aob/mcy210

Gabriel y Galan, J. M., A. Murciano, L. Sirvent, A. Sanchez, Watkins, Jr. J.E. 2018. Germination fitness of two temperate epiphytic ferns shifts under increasing temperatures and forest fragmentation. PLOS One. https://doi.org/10.1371/journal.pone.0197110

Mahley*, J. Pittermann, J. Rowe, N., Baer, A. Watkins, Jr, J.E., Schuettpelz, E. Wheeler, J. Mehltreter, K., Windhan, M. Testo, W., Beck, J. 2018. Geometry, allometry and biomechanics of fern leaf petioles: their significance for the evolution of functional and ecological diversity within the Pteridaceae. Frontiers in Plant Science.  https://doi.org 10.3389/fpls.2018.00197


Krieg*, C., C. Husby, J.E. Watkins, Jr. 2017. Sex-specific differences in functional traits and resource acquisition in five cycad species. AoB Plants doi: https://doi.org/10.1093/aobpla/plx013

Chambers, S., J.E. Watkins, Jr., E. Sessa. 2017Differences in dehydration tolerance among populations of a gametophyte-only fern. American Journal of Botany 104:1-10


Haufler, C.H. , K. Pryer, E. Schuettpelz, E. Sessa, D.R. Farrar, R. Moran, J. Schneller, J.E. Watkins, Jr., M. Windham. 2016. Sex and the single gametophyte: Revising the homosporous vascular plant life cycle in light of contemporary research. Bioscience doi: 10.1093/biosci/biw108

Britton*, M.R. and J.E. Watkins, Jr. 2016. The economy of reproduction in dimorphic ferns. Annals of Botany. 118 (6): 1139-1149 DOI: 10.1093/aob/mcw177

Watkins, Jr., J.E., A.C. Churchill*, N.M. Holbrook. 2016. A site for sori: ecophysiology of fertile-sterile leaf dimorphy in ferns. American Journal of Botany. doi:10.3732/ajb.1500505

New Phytologist Cover

Sessa, E.B., W.L. Testo, and J.E. Watkins, Jr. 2016. On the widespread capacity for and functional significance of extreme inbreeding in ferns. New Phytologist 211:761–764. doi: 10.1111/nph.13670


Zier*, J., B. Belanger*, G. Trahan*, J.E. Watkins Jr., 2015. Ecophysiology of four co-occurring lycophyte species: an investigation of functional convergence. AoB Plants 7: plv137; doi:10.1093/aobpla/plv137

Testo, W.L., J.E. Watkins, Jr., J. Pitterman, and R. Momin*. 2015. Pteris ×caridadiae, a new hybrid fern from Costa Rica. Brittonia DOI 10.1007/s12228-015-9370-8


Testo, W.L., J.E. Watkins, Jr., D.S. Barrington. 2014. Dynamics of asymmetrical hybridization in North American wood ferns: reconciling patterns of inheritance with gametophyte ecology. New Phytologist. 206:785–795. 10.1111/nph.13213

Canestraro, B.K., R.C. Moran, and J.E. Watkins, Jr. 2014. Reproductive and physiological ecology of climbing and terrestrial Polybotrya (Dryopteridaceae). International Journal of Plant Sciences. 175:432-411.


Testo*, W.L. and Watkins, J.E., Jr., 2013Understanding mechanisms of rarity: competition and climate change threaten a rare fern species. American Journal of Botany. 100:2261-2270

Pitterman, J., C. Brodersen, and J.E. Watkins, Jr., 2013. The physiological resilience of fern sporophytes and gametophytes: advances in water relations offer new insights into an old lineage. Invited Review. Frontiers in Plant Biophysics and Modeling. doi: 10.3389/fpls.2013.00285


Testo*, W.L. and J.E. Watkins, Jr., 2012. Influence of plant size on the ecophysiology of the epiphytic fern Asplenium auritum Sw. from Costa Rica. American Journal of Botany. 99:1840-1846.

International Journal of Plant Sciences Cover

Watkins, Jr., J.E. and C. Cardelús.  2012. Ferns in an angiosperm world: Cretaceous radiation into the epiphytic niche and diversification on the forest floor. Invited review for the International Journal of Plant Sciences for the special issue: Major Transitions in Angiosperm Ecology and Functional Biology. 173:695-710.


Laurence, W. … J.E. Watkins, Jr., and 48 other authors. 2011. Global warming, elevational ranges, and the vulnerability of tropical biota. Conservation Biology. 144:548-557. doi: 10.1016 /j. biocon.2010.10.010.

Testo*, W.L. and J.E. Watkins, Jr., 2011Comparative development and gametophyte morphology of the hart's-tongue fern,  Asplenium scolopendrium L. J. Torrey Bot. Soc. 138(4):400-408. 2011 

Watkins, J.E., W.L. Testo*, and R. Merkhofer*. 2011. In situ gametophyte morphology of the tropical epiphyte: Oleandra articulataAmerican Fern Journal 101(1):50-56


American Journal of Botany Cover

Watkins, Jr., J.E., N.M. Holbrook, M. Zwieniecka. 2010. Hydraulic properties of ferns: consequences for ecological and evolutionary diversification. American Journal of Botany. 97(12):2007-2009. Featured cover article


Watkins, Jr., J.E. and C.L. Cardelús. 2009. Habitat differentiation of ferns in a lowland tropical rain forest. American Fern Journal 3:162–175.


Watkins, Jr., J.E., C.L. Cardelús, M.C. Mack 2008. Ants mediate nitrogen relations of an epiphytic fern. New Phytologist 150:5-8.


New Phytologist Cover

Watkins, Jr., J.E., M.C. Mack, T. Sinclair and S.S. Mulkey. 2007. Ecological and evolutionary consequences of desiccation tolerance in tropical fern gametophytes. New Phytologist 176:708-717. Featured cover article with commentary

Watkins, Jr., J.E., M.C. Mack, and S.S. Mulkey. 2007. Gametophyte ecology and demography of epiphytic and terrestrial tropical ferns. American Journal of Botany 94(4):701–708. Featured cover article

Watkins, Jr., J.E., P. Rundel, C.L. Cardelús. 2007 The influence of life form on carbon and nitrogen relationships in tropical rainforest ferns. Oecologia 153(2): 225-232.


Watkins Jr., J.E. A.Y. Kawahara, S.A. Leicht, J.R. Auld, A.J. Bicksler and K. Kaiser2006. Fern laminar scales protect against photoinhibition from excess light. American Fern Journal 96(3):83–92.

Watkins, Jr., J.E., C.L. Cardelús, R. Moran, R.K. Colwell. 2006Species richness and distribution of ferns along an elevational gradient in Costa Rica. American Journal of Botany 93(1):144-156.


Cardelús, C.L., R.K. Colwell, J.E. Watkins, Jr., 2005. Vascular epiphyte distribution patterns: explaining the mid-elevation richness peak. Journal of Ecology. 93(6):73-83.

Watkins Jr., J.E. and D.R. Farrar. 2005. Origin and taxonomic affinities of the disjunct fern Thelypteris burksiorum (Watkins and Farrar). Brittonia 57(2):183-201.


Dute, R.R., D.R. Folkerts, J.E. Watkins, Jr., M.E. Saxon, and R.S. Boyd. 2004. Floral anatomy and pollination biology of Cyrilla racemifloraSoutheastern Naturalist 3(2):309–326.

Moran, R. and J.E. Watkins, Jr., 2004. Lomariopsis X farrarii: a new hybrid fern between L. japurensis and L. vestita (Lomariopsidaceae) from Costa Rica. Brittonia 56(3):205-209.


Watkins Jr., J.E. and D.R. Farrar. 2002 A new name for an old fern from North Alabama. American Fern Journal92(2):171-178.

Moran, R. and J.E. Watkins, Jr., 2002. The occurrence and morphology of Adiantum × variopinnatum Jermy & T. G. Walker (=A. latifolium Lam. × A. petiolatum Desv.; Pteridaceae). Brittonia 54:57-60.

Yoko Y., J.E. Watkins, D.R. Farrar and N. Murakami. 2002. Genetic variation in populations of the morphologically and ecologically variable fern Stegnogramma pozoi subsp. mollissima (Thelypteridaceae). Japanese Journal of Plant Research 115:29-38.


Boyd, R.S., M.A. Wall, and J.E. Watkins, Jr., 2000. Correspondence between Ni tolerance and Ni hyperaccumulation in Strepanthus (Brassicaceae). Madroño 47(2):7-105.

Watkins, J.E. Jr., A. Diamond, R.R. Dute. 2000. Distribution and new county records of the fern: Botrychium lunarioides (Michaux) Sw. in Alabama. Castanea 65(1):64-68.

Watkins, J.E. Jr., and R.R. Dute. 1998. Spore morphology of Botrychium lunarioides (Michaux) Sw. American Fern Journal 88:86-92.

  • 2012: Elected as President of the American Fern Society
  • 2010: Elected Fellow of the Linnaean Society
  • 2009: Nominated for the Phi Eta Sigma Professor of the Year award, Colgate University
  • 2006: Mercer Post-Doctoral Fellow, Harvard University
  • 2006: McLaughlin Dissertation Fellowship, University of Florida
  • 2005: Edgar T. Wherry Award for best talk presented to the Pteridological section of the Botanical Society of America (Austin, Texas)
  • 2004: Nominated for University of Florida teaching assistant award
  • 2003: Nominated for University of Florida teaching assistant award
  • 1999: Edgar T. Wherry Award for best poster presented to the Pteridological section of the International Botanical Congress (St Louis, Missouri)
  • 1997: Young Botanist Award, Botanical Society of America
  • 1997: Outstanding undergraduate poster presentation, Alabama Academy of Science