Posted on Authorea 4 Feb 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161240006.66602471/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. Schuettpelz Campany Courtney drought mechanisms by tolerance driven than are rather ferns avoidance epiphytic in relations water Leaf than-tolerance-mechanisms 507201-leaf-water-relations-in-epiphytic-ferns-are-driven-by-drought-avoidance-rather- epiphyte_water_relations_main.pdf likely with the Even endure file successfully stress. Hosted to water ferns leaf epiphytic avoid allowed that have traits habitat. avoidance canopy structural drought epiphytic the and both for of in of stresses anatomical adaptations conservatism characteristics water for function, share of selection physiological prevalence tissue that the lower via canalized traits demonstrate and exhibited clearly ferns, saturation, ferns findings hemi-epiphytic to at hemi-epiphytic Our compared potential and Overall, species. (-46%) osmotic epiphytic ferns. density higher and reduced terrestrial terrestrial stomatal points, had than both reduced loss ferns loss and turgor Epiphytic explore turgor similar (+41%), status. after to exhibited laminae water capacitance forests thicker ferns leaf tropical Epiphytic (-56%), regulate Rican that lengths hemi-epiphytic, ferns. traits Costa stipe terrestrial, terrestrial functional in in shorter deficits to (-63%), species water findings area to fern these role responses xylem 39 related leaf transitionary surveyed and examined potential We ferns we epiphytic the context. Here, phylogenetic and and a unclear. canopy in remain the divergence However, trait form in epiphytic landscapes. establishment life dominated allowed hemi-epiphytic angiosperm function the in ecophysiological niches of how epiphytic into of radiate understanding to our ferns allowed has diversification Opportunistic Abstract 2021 4, February 6 5 4 3 2 1 ogt University Colgate Ecologia de Instituto Institution Smithsonian University Cruz Pepperdine Santa California of University University Shepherd 4 lu Mehltreter Klaus , 1 aml Pittermann Jarmila , 5 n ae Ede Watkins (Eddie) James and , vial at available 2 lxBaer Alex , 1 https://authorea.com/users/393602/articles/ 2 ee Holmlund Helen , 6 3 Eric , Posted on Authorea 4 Feb 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161240006.66602471/v1 — This a preprint and has not been peer reviewed. Data may be preliminary.

4 Xylem fraction x 10 Stipe length (cm) 0.00 0.05 0.10 0.15 0.20 0 20 40 60 80 Terrestrial Terrestrial a a Hemi−epiphyte Hemi−epiphyte b b Epiphyte Epiphyte b b A A 2 2 Stipe length (cm) Lamina area (cm )

0 20 40 60 80 10 100 1000 0.0 1 Epiphyte Hemi−epiphyte Terrestrial Epiphyte Hemi−epiphyte Terrestrial 0.2 T o t a l

Stipe length(cm) x y l e m

a 10 0.4 r e a s t i p e

( m m − 0.6 2 ) R R R R m 2 m 2 c 2 c 2 o o a a n n r r g d g d = = = = 0.88 0.24 0.88 0.33 100 B 0.8 B Posted on Authorea 4 Feb 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161240006.66602471/v1 — This a preprint and has not been peer reviewed. Data may be preliminary.

− Stomatal density (stomata mm 2) −2 Foliar Nitrogen (%) LMA (g m ) 0 50 100 150 0.2 0.5 1 2 5 0 100 200 300 400 500 600 10 Terrestrial Terrestrial Wright etal.2004 Wright Epiphyte Hemi−epiphyte Terrestrial a a Hemi−epiphyte Hemi−epiphyte b ab Epiphyte Epiphyte b b A A L M 3

2 A 100 ( ) 13

Stomatal size um δ

( C (‰) g

500 1000 1500 2000 2500 m

− −38 −36 −34 −32 −30 −28 −26 2 ) Terrestrial Terrestrial a a Hemi−epiphyte Hemi−epiphyte b ab R R m 2 c 2 Epiphyte Epiphyte o a n r d g ab = = b 0.22 0.79 1000 C B B Posted on Authorea 4 Feb 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161240006.66602471/v1 — This a preprint and has not been peer reviewed. Data may be preliminary.

Ψ ( −1) Ψo (MPa) 1 MPa −2.0 −1.5 −1.0 −0.5 0.0 0 5 10 15 20 0 Terrestrial a 5 100−RWC (%) Hemi−epiphyte ab 10 Epiphyte Epiphyte Hemi−epiphyte Terrestrial b 15 C A 4 ( −1) Cft MPa ΨTLP (MPa) 0.05 0.10 0.15 0.20 0.25 −2.0 −1.5 −1.0 −0.5 0.0 0 Terrestrial 10 a 20

−1 CTLP (MPa ) 0.0 0.2 0.4 0.6 0.8 1.0 1.2 Hemi−epiphyte ε 30

( M Terrestrial a P a a ) 40 Hemi−epiphyte ab 50 Epiphyte Epiphyte a b 60 D B Posted on Authorea 4 Feb 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161240006.66602471/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. 100 E2 100 0 51 E1 100 84 100 100 100 100 Stipe length(cm) 75 100 98 92 100 98 100 100 100 72 Goniopteris curta Goniopteris Goniopteris nicaraguensis Goniopteris 100 Diplazium atirrense Diplazium prominulum Diplazium striatastrum Polybotrya osmundacea Olfersia cervina Dryopteris patula Dryopteris Christella dentata Christella 100 62.043 Parapolystichum excultum 100 Blechnum occidentale Parablechnum schiedeanum Salpichlaena volubilis 100 79 96 100 73 Cyclopeltis semicordata 98 Nephrolepis rivularis 100 Nephrolepis brownii Nephrolepis biserrata Tectaria incisa 5 Oleandra costaricensis Epiphyte Hemi−epiphyte Terrestrial Oleandra articulata 100 100 100 100 Phlebodium aureum 96 Mickelia nicotianifolia Campyloneurum brevifolium 100 Niphidium crassifolium 99 Lomariopsis maxonii Lomariopsis 100 Lomariopsis japurensis Lomariopsis Serpocaulon triseriale Serpocaulon Serpocaulon fraxinifoliumSerpocaulon Elaphoglossum decursivum Serpocaulon loriciforme Serpocaulon Pecluma pectinata Elaphoglossum herminieri Elaphoglossum latifolium Asplenium uniseriale Pleopeltis bradeorum Elaphoglossum amygdalifolium nucleotide substitutionspersite 0 Asplenium serratum Asplenium juglandifolium Asplenium auritum 0.02 Posted on Authorea 4 Feb 2021 — The copyright holder is the author/funder. All rights reserved. No reuse without permission. — https://doi.org/10.22541/au.161240006.66602471/v1 — This a preprint and has not been peer reviewed. Data may be preliminary. 100 E2 8.7 100 51 E1 100 84 S t o m 100 100 a 100 t 100 a l 75

d e n s 100 98 i t y 92

( s t o 100 m 98 100 100 100 a t a

m 72 m Goniopteris curta Goniopteris − Goniopteris nicaraguensis Goniopteris 2 ) 100 Diplazium atirrense Diplazium prominulum Diplazium striatastrum Polybotrya osmundacea Olfersia cervina Dryopteris patula Dryopteris Christella dentata Christella 100 121.6 Parapolystichum excultum 100 Blechnum occidentale Parablechnum schiedeanum Salpichlaena volubilis 100 79 96 100 73 Cyclopeltis semicordata 98 Nephrolepis rivularis 100 Nephrolepis brownii Nephrolepis biserrata Tectaria incisa 6 Oleandra costaricensis Epiphyte Hemi−epiphyte Terrestrial Oleandra articulata 100 100 100 100 Phlebodium aureum 96 Mickelia nicotianifolia Campyloneurum brevifolium 100 Niphidium crassifolium 99 Lomariopsis maxonii Lomariopsis 100 Lomariopsis japurensis Lomariopsis Serpocaulon triseriale Serpocaulon Serpocaulon fraxinifoliumSerpocaulon Elaphoglossum decursivum Serpocaulon loriciforme Serpocaulon Pecluma pectinata Elaphoglossum herminieri Elaphoglossum latifolium Asplenium uniseriale Pleopeltis bradeorum Elaphoglossum amygdalifolium nucleotide substitutionspersite 0 Asplenium serratum Asplenium juglandifolium Asplenium auritum 0.02