Welcome to the Watkins Lab

@ Colgate University!

A place for Pteridological truth.


Here in the Watkins Lab we focus broadly on plant ecology and physiology with an emphasis on ferns. We ask questions about reproductive ecology and how plants balance reproduction with growth and maintenance. Ultimately we hope to understand better the evolutionary and ecological processes that shape plant diversity using ferns as a model system. While the lab specializes in fern ecophysiology, members of the group work on a diverse array of questions from heavy metal hyperaccumulation in Brassicaceae to reproductive ecology in Cycads and functional convergence in Lycophytes. We are a diverse team with many ongoing projects!


Nitta, J.H., J. E. Watkins, Jr., R. Taputuarai, and C. Davis. In Press. Life in the canopy: community trait assessments reveal previously undetected diversity among fern epiphytes. New Phytologist

Kattge, J., G. Bönisch, S. Díaz, S. Lavore, I. C. Prentice, P. Leadley, S. Tautenhahn… J.E.Watkins, Jr. with 75 other authors. In Press. Twelve years of TRY – towards a third generation of plant trait data assimilation and sharing. Global Change Biology.

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

Watkins, Jr., J.E. and R.C. Moran. Online ahead of print. The gametophytes of the ferngenera 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 psychometricpressure-volume curves of fern gametophytes. Applications in Plant Sciences 7(5): e1248.  https://doi.org.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: XX, 1-11.  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. doi: 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. 2017. Differences in dehydration tolerance among populations of a gametophyte-only fern. American Journal of Botany 104:1-10 https://bsapubs.onlinelibrary.wiley.com/doi/full/10.3732/ajb.1600279

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

NP Cover Sessa


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 doi: 10.1111/nph.13985


EDITOR’S CHOICE 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. doi: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., 2013. Understanding 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.
Front 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., 2011. Comparative 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 articulata. American Fern Journal 101(1):50-56

AJP XS Cover.jpg


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.



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 back 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. Kaiser. 2006. 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. 2006. Species 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 racemiflora. Southeastern 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 Journal 92(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.

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.