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\begin{document}

\title{Leaf water relations in epiphytic ferns are driven by drought avoidance
rather than tolerance mechanisms}



\author[1]{Courtney Campany}%
\author[2]{Jarmila Pittermann}%
\author[2]{Alex Baer}%
\author[3]{Helen Holmlund}%
\author[4]{Eric Schuettpelz}%
\author[5]{Klaus Mehltreter}%
\author[6]{James (Eddie) Watkins}%
\affil[1]{Shepherd University}%
\affil[2]{University of California Santa Cruz}%
\affil[3]{Pepperdine University}%
\affil[4]{Smithsonian Institution}%
\affil[5]{Instituto de Ecologia}%
\affil[6]{Colgate University}%


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  \date{June 14, 2024}


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\begin{abstract}
Opportunistic diversification has allowed ferns to radiate into
epiphytic niches in angiosperm dominated landscapes. However, our
understanding of how ecophysiological function allowed establishment in
the canopy and the potential transitionary role of the hemi-epiphytic
life form remain unclear. Here, we surveyed 39 fern species in Costa
Rican tropical forests to explore epiphytic trait divergence in a
phylogenetic context. We examined leaf responses to water deficits in
terrestrial, hemi-epiphytic, and epiphytic ferns and related these
findings to functional traits that regulate leaf water status. Epiphytic
ferns had reduced xylem area (-63\%), shorter stipe lengths (-56\%),
thicker laminae (+41\%), and reduced stomatal density (-46\%) compared
to terrestrial ferns. Epiphytic ferns exhibited similar turgor loss
points, higher osmotic potential at saturation, and lower tissue
capacitance after turgor loss than terrestrial ferns. Overall,
hemi-epiphytic ferns exhibited traits that share characteristics of both
terrestrial and epiphytic species. Our findings clearly demonstrate the
prevalence of water conservatism in both epiphytic and hemi-epiphytic
ferns, via selection for anatomical and structural traits that avoid
leaf water stress. Even with likely canalized physiological function,
adaptations for drought avoidance have allowed epiphytic ferns to
successfully endure the stresses of the canopy habitat.%
\end{abstract}%



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